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sea otter studies - 1970 - alaska

135
SEA OTTER STUDIES - 1970 by Karl Schneider March, 1971 This report is a compilation of rough reports from the marine mammals staff files. It summarizes most of the work accomplished on sea otters during 1970. The analysis of much of the data is incomplete and many of the conclusions are tentative. This report is intended to present the data collected in a useful form and to indicate the progress and stage of thinking at the time it was prepared. It is intended for staff use only. The contents of the report are as follm.,;rs: Surveys Alaska Peninsula - south side Alaska Peninsula - north side Afognak - Barren Islands Kenai Peninsula Prince Willia% Sound Sitka - Cape Spencer ants Sexual Identification of Hale Areas by Pup Counts Kanaga Island Tanaga Island Identification of Male Areas from Ha1.vests Delarof Isla..'1ds Adak Island Amchitka Island Composition of Female Areas Segregation Hithin Female Areas Composition of Hale Areas Population Sex Ratio Summary of .A..nnual Cycle Magnitude of Breeding and Pupping Peaks Gestation Period Fetal Gro,;;th Rate Age of Sexual Maturity Frequency of 07ulation Maximum Breeding Ovulation Significance of Delayed Implantation Fetal Orientation

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Page 1: Sea otter studies - 1970 - Alaska

SEA OTTER STUDIES - 1970

by Karl Schneider March 1971

This report is a compilation of rough reports from the marine mammals staff files It summarizes most of the work accomplished on sea otters during 1970 The analysis of much of the data is incomplete and many of the conclusions are tentative This report is intended to present the data collected in a useful form and to indicate the progress and stage of thinking at the time it was prepared It is intended for staff use only

The contents of the report are as follmrs

Surveys

Alaska Peninsula - south side Alaska Peninsula - north side Afognak - Barren Islands Kenai Peninsula Prince Willia Sound Sitka - Cape Spencer

ants

Sexual

Identification of Hale Areas by Pup Counts Kanaga Island Tanaga Island

Identification of Male Areas from Ha1vests Delarof Isla1ds Adak Island Amchitka Island

Composition of Female Areas Segregation Hithin Female Areas Composition of Hale Areas Population Sex Ratio

Summary of Annual Cycle Magnitude of Breeding and Pupping Peaks Gestation Period Fetal Groth Rate Age of Sexual Maturity Frequency of 07ulation Maximum Breeding Ovulation Significance of Delayed Implantation Fetal Orientation

Sea Otter Studies - 2 - ]1archt 1971

Location of Pregnancy Birth Weight Sex Ratio at Birth Sex Ratio of the Population Litter Size In Utero Mortality Mortality at or near Parturition Recovery of the Uterus after Parturition Frequency of Breeding Conclusions Literature Cited

AERIAL COUNT OF SEA OTTERS- SOUTHSIDE OFTHE ALASKA PENINSULA

March 1970

Between March 16 and March 27 1970 an aerial count of sea otters was rnade along the south side of the Alaska Peninsula from Shaw Island to Cape Lazaref on Unlmak Island Sanak Island the Sandman Reefs the Pavlof Islands northern Shumagin Islands and Sutwick Island were included A BLM Aero Commander N6392U was flown by Cal Ward at an altitude averaging 150 feet and an airspeed bullgtf 100 mphmiddot Observers were Karl Schneider next to the pilot and Jim Faro behind the pilot This is the same technique aircraft pilot and observers as were used in the 1969 Aleutian count The count on March 27 was made in a Gruman Goose 1r1ith Robert Wolfe substituting for Faro Pups with their mothers were not counted

Because of conditions of visibility and distribution of animals all of the counts made are considered to be low

In general the variability in factors influencing this type of count is so great that the results are not rei iable for population estimates or for determining short-term fluctuations in dense populations However aerial counts should be useful to determine general distribution and abundance and to follow large changes in population size Such changes are occurring where relatively dense populations are expanding into habitat that is either unshyoccupied or sparsely occupied Aerial counts are also the quickest way to familiarize new personnel with large areas of habitat

The following is a 1ist of definitions of terms used to describe counting conditions Application of these terms is completely subjective and based on the writers experience

Excellent- Surface of water relatively calm Usually overcast with 1ittle surface glare Single animals easy to spot at a distance

Very good - Surface may be slightly choppy but not enough confusing reflection to prevent spotting of animals off shore

Good - Off shore animals may be difficult to spot but single animals near shore and in kelp beds and small pods everywhere are readily spotted

Fair -Usually choppy or surface glare Single animals ~in kelp beds or in the lee of rocks or shore and most groups of animals relatively easy to see Other single animals and some pods in deep bays in the shadow of cliffs or off shore may be missed

Poor -Single animals difficult to spot and many pods may be missed but conditions still good enough to get a rough idea of the distribution of animals

The conditions encountered for each area are listed below

Description of Conditions During March 1970 Aerial Count of Sea Otters

March 20 Aniakchak Bay to Chignik Lagoon (1022 AM- 1240 PM) Conditions were generally fair with a heavy chop The south side of Sutwick Island and the area between the island and mainland were poor with heavy surf Visibility was very good inside Kujulik Bay however many animals were scattered throughout the area On two passes ~cross the bay 12 and 27 were seen therefore many were probably missed Similar scattering occured in Chignik Bay The overall count for the area is probably low

Pavlof Bay to Cold Bay (130PM-200PM) Conditions poor with heavy chop and squalls The count was superficial however a second superfi cia 1 count of this a rea was made on March 21 and only one otter was seen in Cold Bay

Cold Bay to Caee Lazaref (310PM -420PM) Conditions very poor fair in a few spots Heavy chop dense kelp

March 21 Northern Shumagin Islands (943AM- 1137 AM) Poo~ with chop on windward sides Fai_r on leeward sides (westerly wind) however air turbulence forced us away from high cliffs Heavy surf created very poor conditions on the south side of Unga

Pavlof Islands (1145 AM- 125ff PM) Conditions similar to those in the Shumagins

Chignik Lagoon to Pavlof Bay (305 PM- 535 PM) Conditions varying from fair to poor with chop and surface glare Count concentrated on points and better looking areas Most deep bays not completely surveyed

March 22 Sanak Islands (849AM- 1020 AM) Conditions fair to poor with chop on south side and surface glare on north side Animals scattered and very difficult to see Many otters on rocks Count is probably very low

Sandman Reefs ( 1025 AM~ 1115 AM) Conditions similar to Sanak With the exception of one pod of 450+ the animals were widely scattered and very difficult to pick out Only those very close to the aircraft could be seen Bad squalls moved in and the wind increased preventing a complete count of Deer Island and the eastern half of the reefs

March 23 Wide Bay to Aniakchak Bay (1123 AM- 140PM) Conditions generally good at first becoming fair after Cape Providence with a light chop and some surface glare Otters in Amber and Aniakchak Bays were widely scattered and many may have been missed

Port Heiden to Naknek_ Superficial counts were made from Ugashik Bay to Naknek on March 19 and from Port Heiden to Egegik on March 23 These were made under poor conditions by flying outside the surf line The water was choppy and full of silt No sea otters were seen

11arch 27 Shaw Island to Puale Bay (1030 AM - 1205 PM) Conditions very good to excellent High overcast and very little wind down to Katmai Bay After this the conditions deteriorated rapidly to fair and the count had to be stopped because of a combination of bad weather and lack of fuel As shown on the charts the count was concentrated on the points and islands however the good visibility allowed us to see well into the bays We feel certain no concentrations were missed

The numbers of sea otters counted are presented in the Table Specific locations and numbers are shown on the charts Where coverage of an area was not complete the approximate path of the aircraft is shown

Discussion of Results

Surveys along the Alaska Peninsula have been fragmentary and most of those made were done under less than ideal conditionsmiddot There are a number of reasons for this Weather tends to be poor along the south shore where squall5 form along the mountains Areas where an aircraft can refuel are north of the mountains and there are relatively few passes therefore range of the aircraft is a problem Also there is much shallow water off shore in the area from the Shumagin Islands to Sanak Island Coverage of this area is difficult and many animals are undoubtably missed

The present survey covered most of the area however conditions were such that most of the counts are probably quite low Despite these problems and a lack of continuity we have a fairly good picture of the recovery of sea otter populations in this area

Reports from various individuals are available from the 1930s and 1940s 11ajor surveys by the Fish and Wildlife Service in 1951 (Jones) 1957 (Lensink) 1962 and 1965 (Kenyon and Spencer) covered at least portions of the area In 1969 the southern Shumagin Islands were counted by the Alaska Department of Fish and Game and the present survey covered most of the rest of the area

Basically there are four distinct population nuclei in the area These centered around the Sanak Island-Sandman Reef area the Shumagin Island area the Sutwick Island area and the Augustine Island-Cape Douglas area The following discussion is based on information from reports by Lensink and Kenyon and from Alaska Department of Fish and Game surveys middot

~anak Island-Sandman Reefs

Small numbers of sea otters have been reported at Sanak Island since 1922 No reports came from the Sandman Reefs until 1942 In 1948 27 were sighted at

----Cherni Island The 1951 aerial survey showed 65 around Sanak and 97 in the Sandman Reefs In 1957 251 were seen around Sanak and 508 around the Sandman Reefs In addition two were seen in the Pavlof Islands however few were on the mainland In 1962 548 were counted in the Sanak area and 638 in the Sandman Reefs None were reported from the mainland or Pavlof lsland~however

much of the increase was in the northern area around Deer Island The 1962 survey was probably the best being made under excel lent conditions

AERIAL COUNT OF SEA OTTERS MARCH 1970

Partial or Location Count Date Camp 1ete Count Visibility

Cape Lazaref - Cold lsecty_

Cape Lazaref 3 320 Complete Poor

lkatan Peninsula 71 320 Complete Poor

False Pass - Amagat I 66 320 Camp 1ete Poor

Cold Bay 321 Partial Poor

TOTAL 141

Sanak Islands 239 322 Camp 1ete Fair to Poor

Sandman Reefs

Clubbing Rocks 12 322 Complete Fair to Poor

Cherni I amp Rocks 495+ 322 Complete fair to Poor

Goose I 7 322 Complete Fair to Poor

Hay I 18 322 Camp lete Fair to Poor

Hunt I 2 322 Complete Fair to Poor

Deer I 322 Partial Fair to Poor~

TOTAL 568+

Pavlof Islands

Inner 11iasik 2 321 Camp 1ete Fair to Poor

Outer lliasik 16 321 Camp 1ete Fair to Poor

Goloi 2 321 Complete Fair to Poor

Dolgoi 67 321 Complete Fair to Poor

Poperechnoi 29 321 Complete Fair to Poor

Ukolnoi 2 321 Complete Fair to Poor

yenWosnesenski 4 321 Complete Fair to Poor

TOTAL 122

Partial or Location Count Date Comp 1ete Count vis ib j 1i ty

Northern Shumagin Islands

Unga 184 321 Complete Fair to Poor

Popof 52 321 Complete Fair to Poor

Korovin 46 321 Complete Fair to Poor

Karpa __2t 321 Complete Fair to Poor

TOTAL 286

Cold ~ to Beaver ~ 0 320-21 Partial Poor

Beaver Bay to Kupreanof Pen

Beaver Bay 2 321 Partial Fair to Poor

Cape A 1 iaks in 4 321 Partial Fair to Poor

Guillemot I 16 321 Partial Fair to Poor

__1Kupreanof Peninsula 321 Partial Fair to Poor

TOTAL 23

Kupreanof Pen to Castle Cape 0 321 Partial Fair to Poor

AntJrrCastle Cape to-1-Ji e ~

Castle Cape-Castle Bay 16 321 Partial Fair to Poor

Chignik Bay 118 320 Complete Fair

Nakchamik I 5 320 Complete Fair

Hook Bay 13 320 Complete Fair

Cape Kum 1i un 88 320 Complete Fair

Kujul ik Bay 1199 320 Complete Very Good

Univikshak I 62 320 Complete Fair

Cape Kuml ik 54 320 Complete Fair to Poor

Sutwick I 14 320 Complete Fair to Poor

Cape Agutka 323 Complete Fair~

TOTAL 1766

Partial or Location Count Date Comp 1ete Count vis i b i 1i ty

Cape Kunmik 13 323 Complete Fair

Naka 1i kok Bay amp offshore islands 16 323 Complete Fair

Chiginagak Bay 5 323 Complete Fair

Agripina amp lmuya Bay 105 323 Complete Good

Wide Bay to Puale Bay N 0 T S U R V E Y E D

Puale Bay to c Kubugakl i __]_ 327 Partial Fair

TOTAL 146

Kashvik Bay to L Chiniak 0 327 Partial Very Good to Excellent

Cape Chiniak to Shaw _I

Shakun Is amp Rocks 71 327 Partial Very Good to Exce 11 ent

Kiukpal ik J to Shaw 1 0 327 Partial Very Good to Excellent

TOTAL 71

In the present survey which was made under relatively poor conditions 239 were seen in the Sanak area and 568+ in the Sandman Reefs The latter count Was incomp 1ete and conditions were such that the number of otters was more a function of time spent counting rather than area covered Obviously many were missed As a result not much can be said about total numbers The main change evident is that 141 were seen along the mainland and eastern portion of Unimak Island and 122 were seen in the Pavlof Islands This indicates that a fairly extensive movement northward and along the Peninsula is occuring

A remnant population probably remained along the south side of Sanak Island in the early 1900bulls By the late l940 1 s they began spreading into the Sandman Reefs This northward expansion has continued to the present time There is still unoccupied or sparsely populated habitat along the mainland shore and in the Pavlof Islands There should be continued expansion of this population for a number of years although the numbers around Sanak Island and the western Sandman Reefs may have already reached a peak

With the arrival of substantial numbers in the Pavlof Islands this population is probably on the verge of mixing with the Shumagin Island populshyation No doubt some individuals have moved back and forth in the past but the two populations are almost continuous at the present time

As in past surveys few otters were seen around the north side of Sanak Island and Caton Island This is probably poor habitat for otters The main population is around the south and west sides The higher count on the west end is probably more a result of intensive searching rather than a higher population

Shumagin Is 1 ands

This has been considered the largest of the four populations in the Alaska Peninsula area The most recent counts have not been adequate enough to show any major increase in numbers in the last decade however major changes in distribution have occured which are very similar to the pattern mentioned in the Sanak-Sandman population

There were occasional reports of sea otters in the Shumagins in the 1930s By 1947 Victor Scheffer estimated that 500 1 ived around Simeonof Island In 1953 Hooper counted 633 around Simeonof and Little Koniuj i Island The 1957 survey showed 1829 Most of these were- in the southern islands Five individuals were scattered in the northern islands and one was on the mainland shore near Elephant Point The 1962 survey totaled only 1352 The animals were scattered well off shore and the count was low However the count on Nagai increased from 149 to 338 indicating a continued northward expansion

In 1969 1510 were counted in the southern islands under relatively poor conditions An additional 286 were counted in the northern islands in the present survey and 23 were seen along the mainland north of the Shumagins Again survey conditions were not good

There is a very clear expansion of the population from a center near Simeonof Substantial numbers now occur on all the islands and repopulation of the mainland is occuring The population in the northern islands should continue to increase and most of the habitat on the mainland is not occupied

There is no evidence of an increase in total numbers since 1957middot However the last two surveys have been less than ideal and the large shallow off shore areas have not been covered adequately Considering the survey conditions and the obvious extentions of range it is almost certain that the populations from San-ak Island to the Shumagins have continued to increase In general it appears that the southern islands and reefs have become fully populated and the northern areas are just developing significant populations The entire a rea may be comp 1ete 1 y repopu 1 a ted in the next 10 years This wi 11 depend largely on the status and potential of the off-shore areas

Sutwi ck Area

Reports of sea otters near Sutwick Island are available since 1936 This population was far removed from other populations and is probably a remnant left after hunting ceased In 1951 Jones counted 388 between Cape Kuml iun and Cape Kunmik Most were near Sutwick Island In 1957 889 were counted Nineteen of these were between Cape Kunmik and Cape Providence indicating some northeastward expansion In 1962 949 were seen with individuals straying as far as Cape lgvak and one between there and Cape Kuliak In 1965 a stray otter was seen at Kinak Bay A major shift in the population from Sutwick into Kujulik Bay occured This may be the result of time of year and weather

In the present survey 1766 were counted between Castle Cape and Cape ~~~gtf~aip the majority were in Kujulik Bay Large numbers have moved

~~ranging as far as Cape Kubugakl i The area from Wide Bay to Puale Bay was not surveyed because of weather

The total count for the population was 1912 even though conditions were less than ideal throughout the bcunt and we feel many were missed It is possible that the increase in numbers is entirely due to reproduction assuming a 10 percent annual increase However it is difficult to compare surveys

There is still room for expansion in both directions from this population The greater movement to the northeast is probably the result of better habitat The population around Kujulik Bay is very dense and a large movement of animals may occur if competition for food becomes serious Otherwise we should expect to see continued steady expansion into adjacent areas for a number of years

to come

Augustine Island-Cape Douqlas

For some time a population has existed around the Augustine Island area at the mouth of Cook In 1 et In 1948 approxImate I y 50 sea otters vvere reported from Augustine Island Reports of sightings have been made from Shaw Island to Tuxedni Bay In 1957 Spencer counted 40 at Augustine and one at Shaw Island Lensink counted 52 on Augustine in 1959 but he considered it a poor count In 1965 Kenyon counted 18 on Augustine and 101 in the Shaw Island-Cape Douglas area In March of 1969 Loren Flag saw 62 around Augustine and in May 1969 Jim Faro counted 130 another observer saw about 30 some of which were probably not tallied by Faro

On the present survey Augustine Island could not be surveyed because of weather No otters were seen around Cape Douglas but 71 were seen in Shakun Islands and rocks near the abandoned village of Kaguyak These were probably from the Cape Douglas group

It appears that the animals move about in the area To date no complete survey of the area has been made at one time Until this is done 1ittle can be said about the population other than that several hundred probably occur there

The following table is a summary of sightings and counts made by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game These data were collected by different individuals using different types of aircraft under varying conditions of weather A strict comparison of numbers should not be made from this table

SUM

MAR

Y OF

SE

A OT

TER

SURV

EYS

(Com

pile

d by

p

op

ula

tio

n f

rom

L

ensi

nk

(196

2 T

hes

is

K

enyo

n 0

96

2 amp

196

5 R

epor

ts

amp M

anu

scri

pt)

an

d AD

FampG

Dat

a)

Lo

cati

on

P

re

1951

19

51

1957

19

59

1962

19

65

1969

19

70

AU

GU

STIN

E -

C

DOUG

LAS

Aug

usti

ne

Isla

nd

A

ppro

xim

atel

y 50

(1

948)

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1

amp

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a S

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tin

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from

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Is

to

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i B

ay

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40

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41

52

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18

_lQ

J

119

130+

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130+

J

71

ALAS

KA

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INSU

LA

c

Kul

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-

c

lgva

k

c

lgva

k -

c

Kun

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Ani

akch

ak

Bay

amp

Am

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Sutw

i ck

Are

a M

isce

llan

eous

R

epor

ts

Kuj

ul i

k B

ay

amp c

K

uml

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Sin

ce

1936

C

Kum

1 i u

n

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N

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lan

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8

355 12

13

0 19 6

581

103

180

1

22

47

109

684 86

Not

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7(+

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139

197 14

1 2

53

101 62

5

I

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1951

19

51

1957

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59

1962

19

65

1969

19

70

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(Co

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Chi

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amp

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1969

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430

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75

~-

MARINE MAMMALS SURVEY - BRISrOL BAY TO AKUTAN ISLAND June 2 1970

OBSERVER J Vania J Faro

PILOT George Tibbets Jr

AIRCRAFr Piper Azetex

middotWEATdER Partly cloudy throughout flight wind 10 miles per hour or less visibility generally excellent One bad area and that was at Akutan Island Air was turbulant there and we were unable to go completely around the island because of fog in Akutan Pass Water had slight ripple throughout flight and ocassionally there was glare but overall conditions for sightinmiddot=r animals was good

Departed King Salmon 1115 AM Returned to King Salmon 745 PM

FLIGHT PA~1 middotAfter leaving King Salmon we went over to the north side of the Alaska Peninsula and proceeded down the coast flying at 300 to 600 feet of altitude Generally we flew aboumiddott one mile off the beach At Port Heiden Moller and Cinder River we checked the outer sand bars for seals and some of the inner bars where I knew seal hauled out

We did not cover Izembeck Laroon very well Instead we flew offshore and checked Amak Island We then fueled up at Cold Bay Leaving Cold Bay we continued dmvn the north side of the peninsula (sea otter sighted) down to Cape Sharichef and crossed over to Akutan Island flying along the south side of it We were able to get around Cape Morgan and fly doNn the beach a few miles but then had to turn back becausmiddote of fo9

The flight path was then eastward to the north side of Tigalda Island At Ugamak Island we circled it completely flying at about 300 to 500 feet We completed the survey at this point and returnt~d to King Salmon generally following the route we had taken on the flight down

SEA OTTER SIGHTINGS

Seven pods of sea otter were seen south of Amak Island These were photographed and later counted from middotthe photographs Beshycause of the density in the larger pods and the fact that some animals in the smaller pods were diving when the pictures were taken some individuals may have been missed

The number counted in each pod is as follows

1071 520 357 68 36 75 30

TOTAL 2157

Several hours later the pods had drifted four or five-miles northeastward (see map) bull In addition 1 one sea otter was seen at Amak Island and sixteen near Tigalda Island

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middot~Q

middotmiddotshy

53

S

Sit 51

53

54 ~

_~

I middot

bull

_

bull

46

bull

49

)

~middotmiddot

middot 5

(

47

middot~~_

middot

38

5

6

535

3

middot

48

45

39

61

61

ti

de r

ip

58

56

54

54

56

n 1

1

73

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41

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gt ~51

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gt1lt gt1lt

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6

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shy

SEA OTTER COUNT BARREN ISLANDS and SHUYAK - AFOGNAK ISLANDS

June 9 1970

On June 9 1970 an aerial survey of sea otter was flown in the Barren Islands and in the AfognakShuyak Islands area A Gruman Goose was used with Schneider serving as the only observer Offshore coverage was poor Conditions of visibility were as follows

Barren Islands- 1140 AM to 1225 PM- Water calm moderate surface glare Conditions verygood Count complete

Ban Island to Pernosa Bay- 1255 to 135PM- Conditions similar to Barren Islands but surface glare worse in spots Conditions generally good

Marmot Island to Latax Rocks - 255 to 345PM- Increased ripples on surface Glare Bad Conditions fair Count around Sea Otter Island complete although some otter may have been scattered offshore Remainder of count very superficial

Area

BARREN ISLANDS East Amatuli Island West Amatuli Island Sugarloaf Island Ushagat Island Rocks south of Ushacat Sud Island Nord Island

AFOGNAK-SHUYAK ISLANDS Ban Island-Shuyak Strait Pernosa Bay Marmot Island Sea Otter Island area East side of Shuyak Jest side of Shuyak Latax Rocks-Dark Island

TOTAL

TOTAL

Number of Sea Otters

0 2 0

76 200

29 0

307

18 6 3

121 0

33 26

207

Complete or Visibility Partial Count

Very good Complete ll If II

It II

II

II IJ

Good Complete II

Fair Partial If Complete II Partial II II

II

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lt6gt iltj- cgtcgt Jgt

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9

I

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J

64

7

4

49middot

675

6

57

55

-

7

4

50

5

6

61(

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55

5

467

61

62

5

3

53

49

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5

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6

4

66

48

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-

52

62

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5

5

48

2

4

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46

4

3

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48

49

)

46

9~

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7

4-2

47

41

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63

5

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62

6

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4

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48

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6

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)

middotshy

59

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3~-6

74

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_

20

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1 5

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34

3

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48

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3

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93

52

26

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66

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43

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6

8

60

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middot~

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3

74

12

(+

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30

5

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46

)

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deg

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36

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3

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3

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3

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3

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73

97

7

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19

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0

13

83

31

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0

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46

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38

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29

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24

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ll

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37

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77

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49

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27

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37~~~

32

66

72

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1 2

2

3~

3

0

149

51

SEA OTTER COUNTS - KENAI June J970 to January 1971

Carl Divinyi flew aerial surveys of seals along the outside of the

Kenai Peninsula on a monthly basis He recorded all sea otters sighted

on these surveys The surveys did not cover the entire coast line and

the type of aircraft weather conditions and observers varied from one

survey to another Therefore the seperate counts cannot be compared

However when viewed in total they indicate the areas used by sea otters

and the relative abundance of otters in each area

The attached table presents the counts by broad areas

--

SEA

OTT

ERS

CO

UN

TED

ON

A

ER

IAL

SUR

VEYS

O

F K

EN

AI

PEN

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AERIAL SURVEY Prince William Sound

April 1970

Between April 15 and 18 1970 Carl Divinyi and Julius Reynolds fletr an aerial survey of most of Prince William Sound using a Cessna 185 The survey tras primarily to locate seals however sea otter were counted The following is a summary of the survey

I departed Anchorage on April 14 via Alaska Airlines and arrived in Cordova at 600 AM tJeather conditions prevented flying so Julius and I drove the local road network looking for anything in the way of wildlife

April 15 vleather was a little better than yesterday so we decided to try surveying along the east side of the Sound up to Valdez Arm We started the survey at Bomb Point and flew the coast and offshore islands up to Galena Bay

Observations Seal - Scattered small groups with the majority of the animals being in Port Fidalgo There were no noticeable concentrations (50 on up) of seals

Sea Otter - A total of 105 otter were counted with the majority (60) being located in St Matthews Bay The remainder of the animals were concentrated between Red Head and Knmvles Head outside of Port Gravina

Sea Lion- Small scattered groups in Port Fidalgo (60-80 total)

April 16 Weather inclement - no flying

April 17 We flew Hawkins Hinchinbrook Montague and Green Islands Very few seal- many otter and two large concentrations of sea lions (See maps for numbers and locations of seals sea otter and sea lion) We also counted 5 otter around Kayak Island

April 18 Flew those islands from Montague Strait to Icy Bay and from Port Bainbridge to Knight Island Passage (See maps for numbers and locations of seals sea otter and sea lions)

Julius Reynolds continued the survey along the mainland from Knight Island Passage to Esther Island and around Knight Ingot and Eleanor Islands

The area from Esther Passage to Valdez Arm and Culross Perry Lone Naked Peak and Storey Islands were not surveyed

SEA OTTER COUNTED Prince William Sound

April 1970

Number of Area Sea Otter Date

PORT BAINBRIDGE - ICY BAY Bainbridge Island 30 418 Evans Island 14 418 Elrington Island 4 418 Latouche Island 46 418 Whale Bay - Icy Bay 39 418

CHENEGA ISLk~D - MAIN BAY Chenega Island and Dangerous Passage 33 52 Crafton Island 2 52

PORT NELLIE JUAN - ESTHER ISLAND Blackstone Bay 1 52 Culross Island NS

ESTHER PASSAGE - VALDEZ ARM NS

GALENA BAY FISH BAY 103 415

FISH BAY - GRAVINA POINT 1 415

GRAVINA POINT - CORDOVA 1 415

HAWKINS ISLfu~D 1 4l7

HINCHINBROOK ISLfu~ 99 417

EGG ISLAND 2 417

MONTAGUE ISLAND 259 417

GREEN AND LITTLE GREEN ISLANDS 102 4J7

KNIGHT ISLAND 136 52

INGOT ISLAND 6 52

ELEANOR ISLAND 3 52

SMITH ISLAND 4 52

SEAL ISLAND 0 52

APPLEGATE ROCK 1 52

PERRY ISLAND NS

(continued) SEA OTTER COUNTED Prince William Sound

April 1970

Number of Area Sea Otter Date-

LONE ISLAND NS

NAKED ISLAND NS

PEAK ISLAND NS

STOREY ISLAND NS

KAYAK - WINGHAM ISLAND 5 417

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PRE-TRANSPLANT SURVEY OF SEA OTTERS Green 1 andMontague 1 -July 171970

On July 17 1970 an aerial survey of sea otters was made in the area

around Green Island and the adjacent coast of Montague Island The purpose

of the survey was to locate concentrations of animals prior to a capturing

operation which began the next day A Dornier twin-engine arrcraft was used

with Schneider Reynolds and Somerville acting as observers The sky was

overcast almost no wind and the sea calm However heavy rain showers intershy

fered with forward visibility and the counting conditions were only fair on

the average Conditions vmiddotere especially poor in Port Chalmers and Zaikof Bay

The count began at 210pm and ended at 345 pm

The numbers and locations of otters are shown on the map In addition

16 sea otters were seen on a superficial look at Constantine Harbor on Hinchshy

in brook Island and a pod of 35 were seen three miles east of Johnstone Pt

Most of the area included in the survey was traveled repeatedly in a

skiff over the next four days Durlng this time the weather became calm and

clear for the first time in several days and the bulk of the sea otters shifted

from the coast of Montague out into the shallow areas between there and Green 1

and to Green and Channel Islands It was obvious that many otters had been missed

Fn the aerial sur~ey and that the population in the area is quite dense A

total of 48 sea otters were captured in three days of netting

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SEA OTTER SURVEY Salisbury Sound to Cape Spencer

August 23-25~ 1970

On August 23 1970 an aerial search for sea otters was made from Salisbury Sound to Torch Bay in the area north of Sitka A Helie Courier aircraft was used with Karl Schneider Alan Courtright and

middotWalt Cunningham serving as observers

The water on the outside coast was too rough for good visibility however it was calmer in the lee of rocks and islands and visibility was fair to good

On the initial survey only two otters were seen These were both in Surge Bay on Yakobi Island While returning after completing the count we located approximately 25 in the same area we saw the first two About 15 of these including at least one pup were in a single pod This illustrates how relatively large numbers of otters can be missed from the air The fact that none were seen in other areas does not mean that established populations do not exist elsewhere

On August 24 and 25 a search of the release area north of Khaz Bay was made by the skiff While flying into Kla) Bay for this seach two otters were seen near the old mine of Chichagof Two miners who have been working there reported that two otters have been there off and on since May

Rough weather and outboard problems restricted tl the search area~

and the effectiveness of the search in that area The following sightings were made

82470 I Adult West of Granite Islands 1 Adult South of Granite Islands

11 0A~dgtzl~4JVflup) middot1 In rocks SE of ranite Islands 82570 I Adult -n Southwest of Gran~te Islands

These sightings represent from four to seven animals

Reports over the last year indicate that sea otters regularly move in and out of Kla~ Bay and occasionally as far in as Sisters Lake bull

The fact that the count in the Khaz Bay area Has less than that made in 1969 does not mean much The animals appeared to be scattered during the present count the search did not include all of the nearby sea otter habitat and survey conditions were less than ideal

The population in Surge Bay appears to be separate and is probably well established Two otters were reported in the same location in January 1966 after only 23 otters had been released near Khaz Bay In 1969 two were seen in the same spot from the ait

While no estimate of the population of sea otters in the area can be made from the available information is evident that sea etters can be found along the entire west coast of Chichagof Island and that concentrations occur near Black Island and the Granite Islands north of Khaz Bay and in Surge Bay on Yakobi Island

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HARVEST AND TRANSPLANTS - 1970

Harvest

In May of 1970 a sea otter harvest was conducted on Tanaga Island the middotDelarof Islands and Amchitka Island The numbers to come from each island

and the distribution of the kill around each island were planned in advance Conservative estimates of the population tvere made and the total harvest numbers were based on these estimates The harvest level for Tanaga and the Delarof Islands was set at 15 percent assuming that the next harvest in these areas would be three years later so the rate of harvest would be 5 percent per year Amchitkas harvest level was set at 10 percent assuming an annual harvest and a harvest rate of 10 percent per year

The harvest plan was based on information collected on a June 1968 skiff survey for Tanaga on the 1965 USFWS aerial survey and the 1969 ADFampG aerial

survey for the Delarofs and on 1968 helicopter survey by the USFWS for Amchitka There is good evidence that all the population estimates tvere low especially for the Delarofs

Hunting was done from avo skiffs with two men each Both men shot when possible A 117-foot vessel was used for skinning and living quarters Table 1 shows the schedule of the hunt with the daily harvest numbers

Table 2 presents the projected and actual harvest numbers by area Table 3 presents the numbers killed pelts saved specimen and pelt numbers used and the sex ratio of the kill for each area Figures 1 2 and 3 show the actual numbers and locations in more detail tveather was the main factor causing deviations from the original harvest plan Possible male areas were identified on the 1968 skiff survey on Tanaga An attempt was made to concentrate more pressure on these areas to achieve a more balanced sex ratio This effort is reflected in the sex ratio of the kill A high degree of sexual segregation occurs at this time of year and without specific pressure on male areas we could expect 85 percent females to be taken as occurred on Amchitka With the effort the female percentage was reduced to 65 percent on Tanaga In the Delarofs a male area which had not been identified previously was hit and the effect was much the same

Transplants

Two transplant operations took place in 1970 The first was done in the same manner as the 1968 and 1969 transplants A total of 86 otters tvere captured at Amchitka One aircraft load was shipped out Twenty-nine were released in Oregon and 30 in vashington The animals were held in floating pens at the release sites for several days to recover from the trip Survival appeared to be excellent

The second operation took place in Prince William Sound The Canadian research vessel G B Reed came to the Green Island-Port Chalmers area with tanks built on deck Forty-six sea otters were captured Approximately 44 were alive when the vessel left Prince William Sound however only 14 survived to be released off Vancouver Island

Table 1

April 20

middotMay 1

May 2

May 3

May 4

May 5

May 6

May 7

May 8

May 9

May 10

May 11

May 12

May 13

Schedule of the 1970 harvest

1030 pm depart Homer

Midnight arrive Adak MV Active

Refuel and prepare boat

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Complete skinning pack hides etc in pm

Delarofs

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88

131

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Table 2 Projected and actual harvest numbers May 1970

Projected Actual Tana~a Island Harvest Harvest

Bumpy Point - Hot Springs Bay 200 191

Hot Springs Bay - Twin Bays 50 50

Twin Bays - Cape Sasmik 50 ) ) 199

Cape Sasmik - Tower 120 )

Tower Tanaga Bay 180 166

TOTAL 600 606

Delarof Islands

Gareloi 20-25 0

Kavalga Ogliuga and Skagul 75-100 125

Ulak and Amatignak 35-55 19

TOTAL 150 144

Amchitka Island saved for

USFWS Counting Units 1 amp 2 100 transplant

3 30 82

4 50 36

5 120 87

TOTAL 300 205

Table 3 Details of sea otters harvested in May 1970

Tanaga Island

Total kill 606

Number of pelts saved 598

Number of small pup pelts discarded 8

Specimen numbers SO 70-4 to SO 70-609 Pelt numbers 3198-3769 and 3771-3796

(Seal 3770 void - damaged)

Approximate sex ratio - 220 males 386 females or approximately 64 percent females

Delarof Islands

Total kill 144

Number of pelts saved 138

Number of pups discarded 6

Specimen numbers SO 70-610 to SO 70-753 Pelt numbers 3797 to 3934

Sex ratio- 44 males 100 females or approximately 69 percent females

Amchitka Island

Total kill 205

Number of pelts saved 194

Number of pups discarded 11

Specimen numbers SO 70-754 to SO 70-958 Pelt numbers 3935 to 4128

Sex ratio- 27 males 178 females or approximately 867 percent females

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A combination of bad weather a lack of time to let the otters recover from their capture and problems with the holding facilities probably caused the high rate of mortality

Summary of Harvests and Transplants

Table 4 summarizes the sea otters removed from Alaskan populations in the middot last 20 years An attempt has been made to remove 300 otters from Amchitka each year since 196 7 Originally this number was set as 10 percent of the population which was conservatively estimated to toal 3000 Recent USFtfS helicopter counts of up to 3927 show that this estimate definitely tvas low In the past the removal of animals was from the southeas tern half of the island The 1970 harvest was purposely concentrated toward the northwestern end in order to spread out the pressure and to learn something about the population in that area

The numbers of animals released in all transplant attempts by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game are presented in Table 5

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30

SEXUAL SEGREGATION IN SEA OTTER POPULATIONS

by Karl Schneider March 1971

Workers have recognized for some time that sea otters tend to segregate by sex Lensink (1962) described this segregation around the southeastern end of Amchitka Island and identified three male areasn and three female areas He speculated that females used areas of more favorable habitat and that younger males were excluded by territorial males scattered throughout the female areas The implication is that male areas contain younger or at least nonterritorial males

Marakov (1965) mentions sexual segregation around Medny Island in the USSRs Commander Islands

Kenyon (1969) gives a more complete description of the sexual segregation around the southeastern end of Amchitka and supports it with quantitative data from harvested animals

Both Lens ink (1962) and Kenyon (1969) w_ere primarily describing hauling grounds used by different sexes While Kenyons harvested animals included otters near shore neither study provided much information on the use of off-shore areas

A knmmiddotlledge of the degree of sexual segregation and the location of male and female areas is important to the management of sea otter populations Harvests must be regulated to avoid putting too much pressure on one segment of the population Then capturing animals for transplants it may be necessary to set nets in specific areas to obtain the desired sex ratio case of a localized kill of otters such as when oil spills occur it is important to know the distribution of sexes to evaluate the importance of kill to the population

In

the

By the same token much can be learned about the distribution of sexes through harvest and capture operations The area from which each sea otter was taken during the harvests of 1967 1968 and 1970 was recorded Because a single hunting party might kill 30 otters over up to 10 miles of coast in a few hours it wasnt possible to record the precise locations Therefore the coast was broken up into areas and the numbers killed in each area ltJere totaled

Figs 1 - 5 show the locations of the areas letters correspond to those in Tables 1 - 4 which present the sex and age composition of animals harvested in each area The ages of the 1968 animals 1vere based on cementum deposition and reproductive condition The other samples were broken down using body size In general all males under 25 lb females under 20 lb and both sexes shorter than 100 em were considered dependent pups Females under 35 lb and 120 em and males under lb and 130 em were considered subadults These criteria probably underestimate the age of some animals Cementum deposition information for the 1967 and 1970 samples is not available yet

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Sexual Segregation - 2 - March 7 1971

Information from the 1967 Amchitka harvest and the 1968 1969 and 1970 Amchitka transplant capturing operations is not comparable and is not presented here However knmv-ledge gained from these operations has contributed to our understanding of sea otter distribution and this knowledge will be used in the following discussion In general this data confirms Kenyons (1969) observations for the Amchitka area although these summer and fall collections indicate a higher percentage of males in female areas than Kenyon reported from his winter and spring samples

Kenyon (1969) states that female areas are more numerous and less discrete than male areas He identified three male hauling grounds and 13 female hauling grounds on the southeastern end of Amchitka Island Recent studies involving netting and collecting animals off~hore indicate that male areas remain discrete off shore Usually these areas are off major points of land and pods of males often form in kelp beds directly off shore from the hauling ground

Female areas on the other hand are not discrete at all Any area that is not a male area can be considered a female area Some areas may be more attractive to females with pups or certain areas may be more favorable for hauling out but more females than males vill be found almost everyvthere except in the discrete male areas Therefore the main problem is to locate the male areas in et population He have no evidence of any shift in the location of male areas over a period of time so once an area is identified the information should remain useful for management purposes for many years Lensink (1962) and Kenyon (1969) correctly identified all of the male areas on southeastern Amchitka Extensive harves and netting have not turned up any new areas

Identification of Hale Areas by Pup Counts

In June 1968 a survey of most of Tanaga and Kanaga Islands was made by skiff to gather information to be used in planning harvests from those islands During this survey females with pups were counted separately from single animals No special effort was put into identifying pups fuen a female obviously had a pup it vas recorded Large pups often were separate from the females and some otters were seen only briefly in vaves kelp or at a distance Therefore many pups probably were missed and the counts should be considered minimal 6 and 7 present the counta by area A similar count was attempted by helicopter around Adak but for various reasons the results are not considered useable

From this skiff survey a number of areas were judged to be possible male areas Usually these were points or more exposed aTeas vhere relatively large numbers of single animals but no pups were seen

Shoal Point Kanaga Pass north of Eddy Rock and possibly Naga Point and Cape Tusik were suspected male areas on Kanaga Island Cape Sudak and Cape Amagalik were considered male areas on Tanaga Island and the area around Hazard Point and Lltnnoy Rock in Kanaga Pass was believed to blend with the Eddy Rock area

The best way to confirm the presence of male areas is to collect animals from these areas Harvests were conducted on Kanaga in October 1968 and on

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7A

Sexual Segregation - 3 - March 1 1971

Tanaga in May 1970 The sex ratios of the animals harvested are compared with the skiff survey pup counts in Table 5 The harvest areas are fairly broad while male areas are usually only a few hundred yards wide As a result a harvest area may include both a male area and portions of female areas to either side

The hunting pressure is seldom even over an area Often a good portion of the kill in an area would come from one group of rocks Therefore the information in Table 5 must be tempered with some knmrledge of how the hunting pressure was distributed

Kanaga Island

Shoal Point - While a few more females than males were taken in this area the percentage of males is considerably higher than one would expect in a female area Because weather vas less than ideal much of the hunting pressure ~vas directed to areas other than Shoal Point Most of the females probably came from these areas Therefore Shoal Point is considered to be a male area$

Naga Poin~- There is no strong evidence that this is a male area While no pups were recorded on the survey the total number of otters counted was not high Fairly extesive hunting of the area under good conditions yielded a sex ratio that would be typical of a female area Therefore it is assumed that there is no male area near Naga Point

Cape Tusik - Cape Tusik was suspected as a male area more because of the nature of the area than arry counts The count included areas to both sides of the Cape which could contain females Very little of the hunting was done at the Cape Therefore judgement on this area will have to wait until more information is available Extensive hunting indicates that no male areas exist east of Cape Tusik to Cape Chlanak

Kanaga Pass - From the count it appears that a male area exists in the Pass however the harvest area extended to Hive Rock and included a large portion of female area (Fig 6) The area is too broad as is shown in Table 5 gtvhere the number of pups counted for the whole area is relatively high The results of the harvest confirm the conclusions drawn from the count While the sex ratio for the entire area is roughly equal most of the males came from Kanaga Pass Some females without pups came from that general area but the majority of the females came from the area northeast of the Pass

The Tanaga harvest showed the male areas more distinctly for several reasons The total harvest was higher the entire count area was covered in the harvest and the hunting pressure was recorded more precisely Also as will be shown later sexual segregation is more pronounced in the spring

Cape Sudak - Most of the hunting pressure was concentrated on the tip of the Cape Some pressure was exerted on the area back to~mrd Pendant Point which is a female area as shmvn by the count The high percentage of males harvested from this area clearly confirms that the tip of the Cape is a male area

Tab

le

5

Com

pari

son

of

Pup

s C

ou

nte

d w

ith

S

ex R

atio

o

f H

arv

est

Jun

e

1968

A

rea

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rvey

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cto

ber

19

68 H

arv

est

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ps

10

0

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ps

10

0

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ult

s T

ota

l KA

NAGA

IS

LAN

D

Ind

ep

Ott

er

Ind

ep

Ott

er

M

M

F

Rou

nd

Hea

d-S

ho

al P

oin

t 0

85

3

45

6

55

4

71

5

29

Nag

a P

oin

t-K

anag

a B

ay

66

1

64

middot

18

5

81

5

20

5

79

5

Cap

e C

hla

nak

-Cap

e T

usi

k

51

1

93

2

40

7

60

3

53

6

47

Edd

y R

ock

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e R

ock

61

3

4

35

7

64

3

47

1

52

9

Jun

e

1968

A

rea

Su

rvey

H

ay

1970

Har

ves

t

Pu

ps

10

0

Pu

ps

10

0

Ad

ult

s T

ota

l TA

NA

GA

IS

LAN

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Ind

ep

Ott

er ~ter

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F

M

F

Cap

e S

ud

ak-P

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ant

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4

60

6

39

4

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4

33

6

Bar

nes

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run

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59

3

3

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83

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ays

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4

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77

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23

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60

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Bay

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th B

ay

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11

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3

92

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14

3

85

7

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uth

Bay

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ock

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8

31

6

7

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3

91

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09

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fern

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73

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0

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25

4

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em R

ock

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lak

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int

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35

6

68

4

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6

Ku

lak

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int-

SE

B

igh

t 6

9

82

1

32

8

68

1

51

8

49

Sexual Segregation - 4 - March~ 1971

Annoy RockHazard Point - The harvest did not confirm this area as a male area However the weather Vas marginal and this area was too rough to hunt Therefore the otters taken in the harvest ~vere from either side of the Point and almost none were taken from the portion that is suspected to be a male area Actually Kanaga Pass is narrow and shallmv Otters feed in all parts of the Pass when weather and tide rips permit This could be considered a single male a~ea that serves both islands

Cape Amagalik - Two harvest areas overlap this area Lash Bay-Inferno Reef and Harem Rock-Kulak Point The kill from both areas strongly reflects the presence of a male area Nost of the hunting pressure was concentrated on Inferno and this is probably the main male area

As shown above vle 1vere able to identify four major male areas from a relatively superficial pup count No male areas were located during the harvests that had not already been identified from the count ~mile more detailed observations including field identification of males would be desirable the pup counting technique appears to be an acceptable method

Identification of

Because male areas are usually on v1ell exposed points it is possible to pick likely areas from a chart and then confirm these areas by collecting animals This method avoids the expense time and danger involved in surface surveys of remote areas Actual collection of animals also provides the most concrete proof of the nature of an area

The follmving is a description of the sex composition of areas for which no skiff count data was available

Delarof Islands - No attempt 1vas made to predict the location of male areas in the Delarof Islands This is a difficult area to work in Ogliuga Skagul and a portion of Ulak Island v1ere hunted on Nay 9 1970 The hunting effort in the afternoon was concentrated around Tag Island and the rocks south of Skagul Island While the entire kill is lumped together most of the males bullJere taken in the afternoon The percentage of males taken for the day is higher than would be expected if only female areas were hunted Most likely there is a male area near Tag Island

Adak Island - Portions of Adak gtvere hunted in 196 7 and 1968 No attempt was made to identify male areas before or during either harvest The harvest figures indicate that no male areas have been hunted The Bay of Islands is a classical female area The results of extensive hunting in the Bay and around Eddy Island indicate that there is definitely no male area there The percentage of males taken between Mid Point and Blind Point vms higher than might normally be expected in a female area however the scarcity of subadult males indicates that no male area exists within this area A large congregation of otters is often seen near Finger Shoals This could be a male area but Ye have no information from there If this is a male area a few stray males from there could account for the slightly higher number of males in the kill

Sexual Segregation - 5 - March 1971

Amchitka Island - While much work has been done on Amchitka Island almost all of the effort has been concentrated on the southeastern end from Crmvn Reefer Point to a fegtv miles northvest of Rifle Range Point During the 1970 harvest an attempt was made to distribute the harvest over previously unstudied areas Two areas were suspected to be male areas These were the rocks off Chitka Point and either Bird Rock or Aleut Point Heather prevented us from hunting the area around Bird Rock and Aleut Point

The sex ratio of the kill does not indicate a male area near Chitka Point however heavy waves prevented hunting off the point Therefore a male area could be there but because no animals were killed there the kill figures would not reflect it

Composition of Female Areas

Kenyon (1969) found that 93 percent of the adult otters and 63 percent of the juvenile otters in female areas were females The May samples from the 1970 harvest agree with this The mean percent of adult females in female areas was 93 percent The mean percent of subadults not including pups was 73 percent Hmmiddot1ever in the September and October samples only 77 percent of the a-dults in female areas were female About 86 percent of the subadults were female

The seasonal difference in the sex ratio of adults is consistant for all samples These seasonal changes are most likely due to adult males moving into female areas to look for estrous females During late winter and spring breeding activity is at the lmmiddot1est point of the year and fev1er females are entering estrus In fall the number of estrous animals is at its peak and presumably more sexually mature males move into the female areas to breed It is interesting to note that there were about three times as many males in female areas in fall as in winter and spring Information from female reproductive tracts indicates that approximately three times as many females would be in estrus at any given time in fall It appears that the number of adult males in a female area is directly proportional to the number of estrous females From the data collected there appear to be 15 to 20 males for each female -vlith enlarged follicles at any given time hmmiddotrever hunter bias might influence this figure

Ages have been estimated for otters harvested in 1968 on the basis of cementum layering These age data indicate that virtually all of the males in discrete female areas are either pups (or very young subadults) or are 7 years old or older No males between the ages of 2 and 6 Here found This strongly implies some form of territorial behavior among actively breeding males Fighting among males has not been observed ofteci hmvever a mature male at the Tacoma Zoo became intolerant of a subadult male in the presence of females They frequently 11 fought 1 although neither seemed to attempt to injure the other Finally the young male had to be removed Under imiddotJild conditions the young male might have moved out of the area without repeated physical encounters

Kenyon (1969) described the breeding behavior of sea otters His description indicated that males patrolled an area and two to four males may pass a given point during a 3 or 4 hour period Vandevere (1970) observed some males

Sexual Segregation - 6 - March~ 1971

apparently defending a territory in California but from his observations t

it appeared that successful breeding males were more mobile Perhaps established breeding males are more tolerant of each other More likely they maintain a separation from each other which limits the number in an area In this way a number of males might be patrolling the sa~e area rather than each establishing a geographical territory This might be advantageous where female concentrations shift from place to place as weather conditions change

Why does the number of adult males in female areas decline when the number of estrous females declines The limiting factor may be competition for estrous females rather than territory size With fewer receptive females competition lvould be greater causing some animals to leave the area Another possible reason may be that males may have a seasonal fluctuation in fertility Lensink (1962) found that all adult males produced sperm at all seasons Limited studies since then support this however it is possible that most of these males were collected in or near female areas Also while a male may produce sperm at all seasons there may be less production at certain times of year

A contributing factor may be that males are capable of feeding in more exposed areas than pregnant females or females with pups With a reduced attraction to the female areas either middotbecause of fewer receptive females or a reduced fertility in the male the male may find it easier to obtain food away from the crowded female areas where competition for food is greatest

The fact that the number of males does appear proportional to the number of estrous females indicates that competition for these females may play an important part in regulating the number of males in female areas

The numbers of subadults in the samples is relatively low so fluctuations in sex ratio may be due to sampling errors However the greater number of subadult males found in female areas in winter and spring may be a true increase made possible by the reduction in the number of breeding males A subadul t male would have fewer encounters -vli th breeding males

There are areas within what would normally be considered female areas where subadult males tend to congregate particularly during the winter These are usually areas v7ith a ma-r-ginal food supply that are not heavily used by females Because there are fewmature females in these areas there are also very few breeding males The subadult males using these areas seem to be transients that find less pressure from breeding males there These areas are used less in summer probably because calrrer weather allows feeding farther off-shore reducing competition for food in the male areas

Examples of areas within female areas that are used by subadult males are Constantine Harbor on Amchitka and possibly Finger Bay on Adak

within Female

In the course of the harvest on Tanags in tIay of 1970 a large number of pregnant females tvith large fetuses were collected at one time Most of

Sexual Segregation - 7 - March 1971

these came from Tidgituk Island This raised the possibility that females may segregate according to reproductive condition Table 6 presents the numbers of females in each stage of the reproductive cycle by area on Tanaga Island More pregnant animals were collected in the South Bay area and within that area most of the females with fetuses weighing over 100 g were collected near Tidgiktuk Island This concentration was evident in the pup counts made in June 1968 when a very high percentage of pups was observed in the same area (Table 5) Host females ~vith large fetuses in May would pup by June

The data do not reveal any other areas of this type on Tanaga Harakov (1965) mentioned a bay on Hedny Island that tvas preferred by pregnant females and females with pups Obviously all females do not go to a specific area to pup but a female vi th a large fetus or small pup probably has more difficulty in exposed areas and seeks more protected areas This tendency can be seen by the casual observer Single animals are more likely to venture off-shore and remain in rougher waters In some areas such as Crown Reefer Point on Amchitka there is a male area near the tip of a point Males congregate directly off-shore However to either side of these male congregations there are females that are not accompanied by a pup and probab do not have large fetuses In this way we may find segregation in a single kelp bed off a point If we set a net near the outer margin of the kelp we will catch almost all males Ho~Vever if we set a net to the side of the bed and closer tmiddoto shore we will catch mostly single females Nets set in a nearby bay will catch more females with pups

Table 6 shoHs that an unusually high number of resorptions were found between Cape Sudak and THin Bays This is probably a reflection of poor physical condition of the otters in that area due to food Other data indicate a continuous decline in body condition from the east side of Adak to Kanaga Pass There is some evidence that condition improves west of Kanaga Pass The animals in the Kanaga Pass vicinity are probably in poorer condition than any of the other populations sampled Therefore the high incidence of resorptions in this area should not be considered an example of segregation within the population

ition of Male Areas

As mentioned earlier the harvest areas ivere broader than any male area Harvest areas containing male areas also contained portions of female areas Our identification of the location at rhich each animal vas collected is not precise enough to determine the exact composition of these rnale areas Kenyons (1969) data from winter and early spring harvests are more precise His data indicate that at that time of year 98 percent of the adults and 80 percent of the ju~veniles using male areas were males Of 128 males in male areas 100 gtvere adults and 28 were juveniles

Experience from netting in the summer and harvesting in fall indicates that the percentage of males remains very high all year While adult females may be very close to male areas it appears unlikely that a significant number regularly use these areas at any time year nile adult males enter female areas regularly and in predictable numbers for a specific purpose females probably ~nter male areas only occasionally in stray movements

le 6

R

epro

du

ctiv

e S

tag

e o

f S

exu

ally

Mat

ure

S

ea O

tters

Har

ves

ted

on

Tan

aga

Isla

nd

-by

Are

a -

May

1

97

0

Un

imp

lan

ted

Im

pla

nte

d

Pre

gn

ant

Pre

gp

ant

Fet

us

Wei

gh

t C

lass

A

rea

No

np

reg

nan

t N

o

No

1

2 3

4 5

Res

orp

tio

n

To

tal

A

Su

dak

-Pen

dan

t P

oin

t 12

7

26

8 30

0

2 0

3 2

2 27

( B

arn

es

Po

int-

Ho

t S

pri

ng

s B

ay

12

7 24

10

36

3

0 1

4 2

2 29

B

( (

Bar

nes

P

oin

t-T

run

k P

oin

t 1

1

6 25

7

29

0 3

1 3

0 2

24

( T

run

k P

oin

t-T

win

Bay

s 7

9 39

7

30

2 1

1 1

2 2

23

c ( (

Haz

ard

P

oin

t-T

win

Bay

s 4

3 27

4

36

0 0

1 1

2 11

( T

win

B

ays-

Her

d

Roc

k 1

2

8 28

9

31

1 1

1 Lf

2

29

D

( ( T

win

B

ays-

So

uth

Bay

2

9 45

9

45

3 0

1 1

Lf

1 20

( S

ou

th B

ay-L

ash

B

ay

6 9

29

16

51

1 0

0 9

6 1

31

E

( ((

So

uth

Bay

-Har

em R

ock

17

14

31

14

31

2 0

1 5

6 45

(

F ( (

L

ash

B

ay-I

nfe

rno

Ree

f 4

1 9

6 55

2

1 1

0 2

11

( G

(

Har

em R

ock

-Ku

lak

Po

int

12

6 29

3

14

1 0

1 1

0 21

H

Ku

lak

Po

int-

SE

B

igh

t 13

9

28

10

31

2 0

1 2

5 32

Bra

cket

s in

dic

ate

o

ver

lap

pin

g are

as

Sexual Segregation - 8 - March 1971

Juveniles of both sexes probably move more randomly than adults lihile definite sexual segregation exists among younger animals it is not as pronounced as in adults Because subadult males are tolerated less by breeding males they are more strictly confined to small areas than females of the same age This has been demonstrated in fall harvests where extensive hunting over a large area will turn up only an occasional subadult male Then ~v-hen the hunters move to a definite male area large numbers of subadult males are taken in a very short time and in a very small area

While Kenyons data indicate that 22 percent the males in male areas are subadults our experience indicates that this percentage may be quite a bit higher at least in fall Such a seasonal change in the age composition of males sounds reasonable We have demonstrated that adult males move into female areas in greater numbers in the fall perhaps tripling their numbers in these areas At the same time fewer subadult males are found in the female areas Presumably the adult males are coming from male areas and the subadults driven from the female areas move to the same male areas The result vmuld be a higher percentage of young males in male areas in fall than in spring

The total number of males of all ages in female areas doubles in fall About 13 percent of all animals taken in these areas in spring were ~ales while 25 percent ta-ken in fall were males This suggests that the total number of males in male areas declines in fall There are more adults leaving than subadults entering the area

Sex Ratio

With the sexes segregating and with the degree segregation changing seasonally and betmiddotween age groups it is extremely difficult to get a completely random harvest As a result we have not been able to determine the sex ratio of any population as a vhole Hith female areas being larger and in more protected areas where hunting is easier there is a strong tendency to take more females than males particularly in sp-ring ~Je have shown that through a concentrated effort the percentage of males harvested can be increased however even then the lowest percentage of females taken in recent harvests ivas 62 percent on Kanaga Island in October 1968 Similar results occur in capturing operations for transplants

On all islands studied there are fewer male areas thltm female areas The male areas are very small usually only a few hundred yards wide The male feeding areas off male areas are also very narrow although they may extend farther off-shore In short that portion of the total available sea otter habitat included in male areas is very smalL

All evidence indicates that there are more females in the population than males While the percentage of is probably greater than that indicated by most harvest statistics it appears that at least 60 perc2nt of the sea otters in the populations studied are females

There are probably several factors contributing to this uneven sex ratio First our reproductive data indicate that 56 percent of the pups are females at birth Secondly Kenyon (1969) has found a higher mortality rate among

Sexual Segregation - 9 - March 1971

newly weaned males There is also some evidence that males might not live as long as females These factors could easily account for the preponderance of females in the population

It is possible that the higher rate of mortality in juvenile males is in part due to the breeding behavior of adult males and the resulting sexual segregation Juvenile males tend to be restricted to male areas or areas of marginal feeding habitat where there is little competition from breeding males During the ltvinter the feeding activity of young males would tend to be confined to small areas close to shore in male areas Therefore they may be subjected to greater competition for food than females of the same age which have a broader area in ltvhich to feed Kenyon (1969) found that subadult males vere less hardy in captivity than subadult females Therefore we can not adequately evaluate the influence of segregation on mortality

The sex ratio probably varies from one population to the next Very little juvenile mortality occurs in expanding populations eliminating that source of sexual selection However there is some evidence that males are more numerous among the first animals to repopulate a new area While a high percentage of males might be found on a newly repopulated island the loss of these males tvould alter the sex ratio of the parent population even though juvenile mortality might not yet be a significant factor

If the nThuber of breeding males is regulated by the number of estrous females an even sex ratio might produce a surplus of sexually mature males This surplus would not be beneficial to the population and could be detrimental This would permit the situation of an unbalanced sex ratio to evolve

The segregation of sexes and ages and the composition of sea otter populations is complex Unless we can understand the situation we cannot fully evaluate the effects of mortality or habitat changes

REPRODUCTION IN THE FElIALE SEA OTTER

by Karl Schneider Narch 1971

A total of 1358 sea otter reproductive tracts collected between 1967 and 1970 have been examined Ovaries were sliced with a razor blade and examined macroscopically Uteri Here macroscopically examined both internally and Each tract vas classified as to its stage in the reproductive cycle

The follovJing criteria were used in classification

Nulliparou~- Horns of uterus small thin and smooth no corpora lutea or corpora albicantia although there is occasionally evidence of a past ovulation but no evidence of implantation

Nultiparous - Uterus thicker corpora lutea andor corpora albicantia present Includes primiparous animals (these are not readily separated from multiparous animals)

Anestrus - Largest follicle under 3 5 rnm no corpus luteum

Proestrus - Follicles over 35 mm

Es - Follicles approximately 10 rrm

Implan~elt_ Pregnant_ - Visible s~velling in uterine horn usually with fetal membranes embryo or fetus visible Corpus luteum usually over 10 mm

- Uterine horn sti11 noticeab enlarged fresh roughshy~=-=--=~

al scar newly formed corpus albicans with some luteal tissue remaining

The appearance of the uterus ~ras used to confirm the classification The thickness arrd of the horns thickness and consistency of the uterine walls coloration of the of the horns and condition of the rugae change with each stage In most cases it is possible to guess the condition of the tract by a superficial examination of its exterior This appearance was used only to confirm what was indicated by structures in the ovary hmvever

Table 1 smnmarizes the reproductive condition of the sexually mature females in the that are enough for comparison throughout the year In the following discussion reported by Sinha et (1966) and Kenyon (1969) are included 7here possible The January and Harch are from these studies

Tab

le

l Re

p iv

e co

nd

itio

n o

f se

xual

ly m

atur

e se

a o

tters

_

IF

etus

Wei

--------------+

__

_

ln~a

ctive

ov

arie

s 1

Act

ive ovari~

ht C

Jas

s

jmiddot

I P

ost

-I P

roes

tru

s U

nim

pl

lmpl

D

ates

L

ocat

ioQ

1A

nest

rus

Part

um

Res

orpt

ion

l +

E

stru

s P

reo

Pr

eQ

2 3

lJ

S

~

4-8

1970

T

anag

a 1

51

41

0

10

104

17

8 10

33

30

4 (1

68)

(1

35)

(3

3)

(3 3

) (2

89)

(34~

(5

6)

(27

) (3

3)

(11

4)(

10

9)

May

9

19

70

Del

arof

Is

18

14

1

4 13

27

10

0

3 6

8 77

4

) (1

82)

( l

3)

(5

(16

9)

( o

) (1

30

) (3

9)

(78

)(1

04

)

10-1

219

70 A

mch

ltka

I

34

18

3 8

27

48

5 6

2 23

12

13

8

( 0

(24

6)

(13

0)

(22

) (5

8)

(19

6)

(34

8)

(36

) (4

3)

(14

) (1

67)

(8

7)

~lune

23

middotmiddot ~middot~~middot~Amchitka

1

17

2 1

2 4

18

1 2

3 5

7 44

A

ugus

t 13

196

8 (3

86)

(4

5)

(23

) (4

5)

(90

) (4

09)

(2

3)

(45

) (6

8)

(11

3)(

15

8)

July

6-

~dgtAmchitka

1

17

1 0

4 14

10

0

1 1

2 6

46

Aug

ust

819

69

(36

9)

( 2

0 2

) (8

6)

(30

4)

(21

9)

(22

) (2

2)

(43

)(1

32

)

Sep

t 1

0-17

A

dak

72

5 0

16

31

40

6 6

8 9

11

164

1967

(4

39

) ( 3

1)

(98

) ( 1

89)

(

4)

(3 7

) (3

7)

(49

) (5

5)

(6

7)

Sep

t

25

-A

mch

itk

a l

55

6

0 18

19

17

4

0 0

0 3

115

OcL

6

19

67

(4

7~8)

(5

2)

(15

7)

(16

5)

(14

8)

(35

) (8

7)

(26

)

Oct

o 1

2-1

5

Kan

aga

I

58

6 1

13

31

31

4 4

7 11

5

140

1968

(4

13)

(4

3)

(o 7

) (9

3)

(22

2)

(22

2)

(29

) (2

9)

(50

) (7

8)

(36

)

Oct

18

-20

A

dak

l

46

6 0

1 24

13

2

3 0

4 4

100

1968

(L

16 0

) ( 6

0)

(11

0)

(24

0)

(13

0)

(20

) (3

0)

0)

(40

)

Num

bers

in

re

nth

esis

are

pe

rcen

t se

xu

ally

mat

ure

fem

ales

F

etus

wei

ght

clas

s 1

=

0-l

g

2 =

1-l

Og

3

= 1

0-lO

Og

4

= 1

00-lO

OO

g

5 =

Tra

nsp

lan

t m

ort

alit

ies

(all

o

ther

sam

ples

fr

om

har

ves

ts)

Reproduction - 2 - March 1971

There may be some problems associated with comparing samples taken from different islands and in different years but allowing for sampling errors the data for the most part fit a definite pattern indicating that the annual cycle remains fairly constant The one outstanding exception is the number of pregnant animals in the t1m summer samples In 1968 there were many more implanted pregnancies than unimplanted pregnancies and in 1969 the situation was reversed hmvever the total number pregnant was almost identical These are the t s being relatively small collected over a longer of time and consisting of transplant mortalities An average of the two samples fits the pattern established by the other samples Relatively large samples are necessary because all stages of reproduction are found at all times of the year The period from November to early January is not represented hmmiddotJever things are changing so rapidly during that period that any information from that time may be confusing unless a large sample was collected in a very short time

The information in Table 1 is shmvn graphically vJith Kenyons (1969) winter information in Figs 1 and 2 It has often been demonstrated that sea otters breed and pup at all times of the year A number of observers have noticed that roore pups are born in the spring aJd sumrrter than at other times and there have been conflicting reports about breeding times Kenyon (1969) was the first to demonstrate seasonal changes in pregnancy rates but only his January and Yarch sallples gtvere large enough to be at all reliable

Figs 1 and 2 demonstrate that there are cons le secsonal in the numbers of animals in each s reproductive cycle There is a definite period of increased bree activity and a definite period of increased pupping Hmmiddotlever ~ some anii1als are in each s at all times of the year and these periods of increased breeding and pupping do not have distinct boundaries

Each curve is influenced by two factors so it is difficult to isolate the magnitude of any single factoc For example the of implanted pregnancies vill increase ss blastocysts and decrease as births occur The birth rate may be increas but if the number of implantations increases at a greater rate the implanted curve will still rise The animal remains in the anestrus implanted pregnant~ and unimplanted pregnant categories for a relatively long time and there may be considerable carry-over from one sample to the next Because the animal a relatively short time in the proestrus-estrus and post partum categories there is little if any carry-middotover from one sample to the next and the changes betgtmiddotTeen are more likely to be absolute changes The mean fetus weight class prob does not mean much because it does not take the actual number of animals in each class into account ~hen there are many implanted pregnancies ard the mean lmiddotTeigh t class is lovJ there actually may be more Class 5 fetuses in the population than vhen there are fetv irr1planted pregnancies and the mean middotleight class is high Therefore the actual percentages of all sexually mature females middotlith Class 1 ald Class 5 fetuses have been plotted in 3 These curves represent the actual number of fetuses in the population Because tha sample sizes become quite small vhen the fetuses are divi(12d into classes and the surruner sample is believ2d

2

~ _s ~

~

c U)

~-1

r

~J

lt

c])

Reproduction - 3 - Narch 1971

to be biased tvo curves for each class are shmvn The solid curves approximate the data while the broken curves represent subjective adjustments to correct for sampling errors

cussion of the Annual

The number of animals entering estrus rises in the fall At this time the pregnancy rate is at its lowest point and a high percentage of anestrous (nonpregnant) animals are in the population Breeding activity increases and at this time the number of sexually mature males in 1female areas increases

The number of unimplanted pregnancies begins to rise sharply in October and November At the smne time the nlli~ber of anestrous animals drops as these animals enter estrus breed and become pregnant Thj_s is the peak breeding season

The birth rate is at its louest this period as is shovm by the lmv number of post-partum a1imals and the lovr number of large fetuses

By late November and December the pregnant rate is increas even though the birth rate is slightly This is due to ne7

blastocysts implanting as is shown the increase in Class 1 fetuses

By January the period of t breedLng activity has declined and the number of unimplanted animals ceaches a peak as the number of blastocysts implanting equals the number of The lanted pregnancy rate then begins to declLne as fevJer animals breed and more implant The greater rate of antation is reflected in a rise in the Class 1 fetuses as well as an overall increas in the er of lanted pregnancies However the rate of implantation is partially obscured by an increased birth rate removing animals from the total of lanted pregnancies This increase in the birth rate is shmm in the rise in post-partum females and a rise in Class 5 fetuses

This condition of a low rate of breeding high rate of implantation and increasing rate of birth lasts from February to Hay

At this point He come to the least reliable data The sumner are smaller gtJere collected over a period of seven weeks and were from animals that died as the result of capture and handling

Therefore tvhile the data shaH that the post-partum rate decreases sh indicating a reduction in the birth rate~ the number of Class 5 fetuses

a peak indicating a very high birth rate The true birth rate is probably some1middothere in betigtJeen Females 1-ith very young pups (hence are post-partum) are vlary and probably less likely to be caught in nets Females with fetuses appear to be more likely to from handling In several cases~ twisting of the reproductive tract by a fetus vJas the actual cause of death Anestrous females (including post partum) on the other hand are less likely to die

Reproduction - 4 - March 1971

There is no question that the birth rate remains high during June and July Kenyons (1969) sum11er sample is limited but shows a large number of Class 5 fetuses Again the sample is biased in favor of pregnampIt animals but not to large fetuses only His field counts as vJell as observations by most other observers indicate an increase in the nUIlber of dependent young througbout the surnmer The number of Lmted pregnancies declines throughout the suTimer tvhile the unimplanted pregnancy rate declines at a slmver rate and breeding remains fairly constant at a lm-v level This indicates that through June there are more implantations than conceptions hmvever the birth rate is substantially greater than the implantation rate By late July and August there are relatively fe1r implantations taking place as shmm by a leveling of the unimplanted pregnant rate and a lotr number of Class 1 fetuses The number of Class 5 fetuses declines as a result of births during June July and August

Therefore the season of greatest pupping may be quite broad running from April through August possibly with the peak occurring in late May or June Until a better summer sample is available tve cannot pin dmm the peak of this pupping season with certainty (See Fetal GrmmiddotJth Rate for further discussion on breeding and pupp peaks)

If the information were precise we could compare subtracting numbers of pups born and numbers of conceptions to deterrnine the actual percent of ferrales breeding implantLng ltmd pupping in each month Unfortunately the data are not Each curve is influenced by more than one factor and ue do not knmmiddotr for sure the magnitude of any of these factors Hmmiddotrever by comparing some of the major samples e may be able to get a rough idea of how much higher the birth rate is at one time of year than another

The combined Mty and the combined September and October samples provide the best comparison Both are large samples One occurs near the peak of pupp and a lmv point in breeding and the other is near the low point in pupping and the peak of b

The number of Height Class 5 fetuses~ post-partum femaless and proestrousshyestrous females come closest to represent a particular event The time spent in each category is comparatively short

There were approximately three times as many Class 5 fetuses in Hay thanmiddot in Septerqber-October There Here also about three times as wany postshypartum females in May In tember-October there were two and a half to three times as many proestrous-estrous animals as in Hay It ~middotwuld appear that the peaks of breeding and pupping are roughly three times as high as the lowest points

At the lmmiddot12st period of pupping 2 to 3 percent of the females have Class 5 fetuses and should pup middotJithin a month If vm assume the post-partum stage lasts L 5 to 2 months (see Recovery of the Uterus) it also appears that 2 to 3 percent pup in the ltwrest months of pupping

Reproduction - 5 - March 1971

Similarly about 3 percent of the females have enlarged follicles during the lmvest period of breeding He do not knmv how long this period lasts hovever

At the peak of pupping about 10 of the females have Class 5 fetuses and should pup in the next month About 15 are post partum indicating about 8 births per 100 females in the preceding month

These percentages are prob2bly slightly high because of hunter bias avoiding females with pups It appeatmiddots that at least 2 to 3 percent of the mature females breed and 2 or 3 percent have pups in any month of the year In the peak breedingmonths such as September and October perhaps 7 to 10 percent breed and similarly in peak pupping months sud1 as Nay 7 to 10 percent of the mature females pup

Gestation Period

Barabash-Nikiforov (1947) listed the gestation period of 8 to 9 months This apparently was based on the case where a fentele mated in captivity as reported by i-Ialkovich (19 37) According to Barabash-Nikiforov a pup was born 8 months later although it died While it was fully formed~ it may have been premature

A number of births have occurred in captive animiddotmals held by the of Fish and Game in recent years In all cases the pu-ps ~-ere sc-middot1all usually less tha1 1600 g but still than the smallest pups found in the -rild All died after birth

The gestation period of 9 months has been repeated in the literature but apparently these statemsnts are based on Barabash~Nikiforol (19lf7)

Lensink (1962) also estimated a gestation period of 8-9 months based on field observations He felt that the peak of breeding was in August or September and the of pupping ~Alas in l-1arch or April

It has been demonstrated that the gestation of sea otters a relatively long period of tation (Sinha 1966 Kenyon 1969 and present study) Therefore any attempt to estimate the length of gestation by examinatioD of tracts must take both the unimp1anted and Janted periods into accour1t

Kenyon (1969) presented an estimate of the implanted period by assuming that the cube roots of fetus weights fall in a straight line after the embryo is established (Hugget and Hiddas 1951) and by that the European otter and Aluerican river otter would have similar fetal growth rates By plotting tenn fetus on a line established by the European otter he estimated an gestation of about 120 days not including the period for establishment of the ewbryo

Kenyon (1969) then an estimate made by Dr D G Chapman using the method of Hugget and ~-Jiddas (1951) v7here a regression line is fitted

Reproduction - 6 - March 1971

to the cube roots of the raeail of the mean weights of fetuses in each of five vreight clas3es plotted t the mean time of year

From the regression coefficient ob he estimated an implanted period of 154 days By taking the mean of the pregnant animals that are unimplanted pregnant estimated the tmimplanted period as about 75 months 1Ulmv-ing a half month for establishment the embryo he estimates a total gestation of 12-13 months

The data used by Chapman included a January-February sample of 26 a March-April sample of 49 and a Nay-August of only 9 The surtmler sample is more biased than the others and quite small yet the estimate relies heavily on this sa~ple when fetus size is the greatest

Table 2 presents a neTw- estimate of the length of the implanted period using the method used by Chapman but including data from the present study in addition to Chapmans data Table 3 presents a similar estimate of the length of the unimplanted period

The following is a comparison of the two estimates

Unimplanted Period 230 days 66 days Establishment of Embryo 15 15 days Implanted Period 154_~~Yrgt~ _73 d~s

Total Gestation Period 399 d2ys 154 days or about 13 months 5 months

Obviously there is some problem tiith the the technique or both A similar calculation mcde before the Hay sample had been collected produced an estimate of 109 for the implanted period and about 8 months for the total ation period

This method of calculation assur1es that there are definite peaks in breeding and pupping If breeding and pupping occurred evenly throughout the year the average fetus size 1vould remain constant thruughout the year and the technique would not tvork An ideal situation v10uld be when all individuals breed and pup 1vi thin very short periods Sea otters do have but are not well defined Very large s v-ould be required in such a case because there are ahvays fetuses of all sizes and the periods of maximum breeding are broad

Obviously Chapmans sample vas inadequate llthile the number of fetuses available for the latest estimate seems large and are well distributed throughout the year the a_ctual nu~ober in any cne fetus vJeight class at any one time of year is small A look at Table 1 shows that there is little consistency in many of the classes (Fig 3 shmvs that Class 1 and 5 fetuses do fit a pattern)

Table 2 Estimated Length of Period

Percent of Fetuses in each Height Class

Time of Collection After January

1 month 31 15 15 38 0

3 months 18 12 22 35 12

5 months 18 8 8 36 30

7 months 3 8 16 29 45

9 months 20 7 10 40 23

10 months

14 18 12 33 23

---~--

r1ean Time in Nonths After January 50 57 53 58 70

Cube Root of Umveighted Mean 063 17 36 7lf lllf

Specific Grmth Velocity 0169day EstirGated Implanted Period = 73 days

Table 3 Length of Unimplanted Period

Months After Unimplanted Unimplanted Percent January Pregnant Pregnant Unimplanted

1 month 35 26 58

3 months 49 49 so

5 months 128 179 42

7 months 22 37 37

9 months 50 57 47

10 months 55 44 56

Unweighted

48

Reproduction - 7 - March 1971

All of the estimates very heavily on su111mer samples 1vhen the fetuses are largest These sauples are knmm to be biased probably in favor of larger fetuses There are more fetuses at this time of year but it is exaggerated in these SCilllples

The latest estimate is not necess better than Chapmans even though it uses more and larger samples The velocity is twice as high as that calculated for fur seal and a 5 month gestation period is not consistent with other information available

This technique for estimating the lanted gestation period is not capable of providing a reliable estimate with the available data Therefore the 12 to 13 month estimate and any calculations based on it are meaningless

Much of the discrepancy lies in the estimate of the unimplanted period The present estimate that 48 percent of all pregnancies throughout the year are unimplanted based on an even distribution of relatively large samples and is probably more accurate than Chaprnan 1 s estimate of 60 percent

There are several other ways of guessing at the length of gestation Kenyons (1969) estillate of an lanted period of 4 months using the cube roots of term fetus weightscannot be relied upon entirely but it may be closer than the other estimates If gtmiddotJe estimate the uninplanted period from this using the 48 percent ted factor and allow for the establishment of the embryo we get a total gestation of about 85 months

Lensinks (1962) method of detennining the and peak pupping periods and taking the time between the peaks as the total gestation period has some merit TJnile Lens ink tried to determine these peaks by field observations we can determine them more accurately using the condition of reproductive tracts

Lensink felt that the peak of breeding middotlas in August or September The data in Fig 2 indicate that he may seen the beginning of the period of est breeding activity but that the probably in October or possibly November Lensink estimated period in larch or April Again the data indicate that he v1as s the beginning of the period of greatest pupping but the actual peak is probably in June or July At this time the number of Class 5 fetuses is greatest The nucJber of post-partum females probably and the number of implanted pregnant females is dropping from its If the gestation period of

11 11the average animal lasts from October or November to June or July we get a period of 8 or 9 months tThile Lensink 1 s was early 1 the duration agrees vith the present data

The peak of implantation when the greatest number of blastocysts are implanting can also be roughly estimated from 1 and 3 In February the number of unimplanted middot is dropping and the number of implanted pregnancies at this time th2 number of Class 1 fetuses probably reaches a

Reproduction - 8 - March 1971

Therefore it appears that the unimplanted period lasts 4 to 5 months and the implanted period from 4 to 5 months 1vi th a total gestation period of 8 to 9 months It should be recognized that these estimates are based on general trends and not distinct Therefore they are approximate However the relative length of the unimp and implanted p2riods agrees with the percentages noted previous and this estimate agrees vlith the one based on Kenyons (1969) comparison vdth other species of otters

If the gestation period ~Jere 12 months the percentage of pregnant animals would remain constant throughout the year If it vJere only slightly longer or slightly shorter there would be only a slight fluctuation unless there were a distinct breeding period Fig 4 indicates a substantial fluctuation indicating a gestation period substantially shorter or longer than 12 months

The peak of pregnancy occurs in February after the period of t breeding activi The luH point on the pregnancy curve occurs after those animals giving birth dur the period of t pupping have pupped That is the t point on the pregnancy curve occurs after the main breeding and the lmves ~ point is after the main pupping period These periods actually last for 4 or 5 months so these points are well after the peaks of breedtng and pupping

The time betbulleen the peak when the pregnant and the lmr point Hhen the fewest are the average gestation period This is about 85 months (Fig 4) The anestrus curve (Fig 1) shmvs the same thing betng a negative of the pregnancy curve

Hith three different methods we have estimated a gestation period of 8 to 9 months Hith the unimplanted period roughly equal to the implanted period This agreas bullmiddotlith the observed period in captivity (BarabashshyNikiforov 194 7) and with field estimates Any gestation period differirg greatly from this does not fit the data Conceivably one could apply a 20 to 21 month period to the SaTEpound information however if this 1vere the case females would have to breed vJhile accomp by a pup or be on a four year reproductive cycle This prospect stretches the imagination

There is a definite possibility that the gestatton period varies in sea otters as in land ot te~cs This most likely -middotwuld occur through variations in the unimplantec1 period The above discussion refers to the average gestation period Until better information to the contrary is obtained we must continue to assume that the average period is 8 to 9 months

Fetal Rate

the estimates of the gestation curve are corrects Kenyons (1969) Fig 4 should approximate the grouth curve of the fetus Fig 5 duplicates this and shmvs the actual Heights Fig 6 shous the same information more graphically It is possible to esttmgtte the of time in each v7eight classlt Fig 6 does not take the period for establishment of the embryo into account so Height Class 1 may be longer than shmm perhaps 15 days if we accept Chapcnans (Kenyon 1969) es e

--

3 0

~

shy

----- -shy --middotmiddot~middotmiddot

___ ___1__

_

(middot-)

_ ft -) J ~)___

~-

3

Reproduction - 9 - IYarch 1971

The short time span of Classes 2 and 3 explains the relatively small numbers and the lack of a consistent pattern found in those classes (Table 1)

Using this curve it should be possible to predict the time of birth and with less precision the time of Luplantation and conception of any particular fetus By plot the estimated birth dates of a large number of fetuses rve should be able to drav curves Ttlhich approximate the breeding implantation and pupping rates of the population throughout the year

Unfortunately there is overlap bet1veen samples collected at different times of year Some of the samples are too small to provide a comparison We vould have to give eight to each in order to combine them Therefore only the ttay September and October samples are used lfuere the September and October samples overlap t1lt10 separate points are used Fig 7 presents the estimated birth dates grouped by month Fig 8 separates the dates into half month groups February and Narch are not represented and April is an estimate based on post-partum females rather than fetus size (see Recovery of the Uterus after Parturition)

The peak in this curve may be exaggerated It has been shmvn previously that fetus size is less consistent bet-Jeen samples than the other categories (Table 1) because the sample size of each grouping is small Therefore the exact shape of the curve may not be correct If however we assume the approximate of the peak is correct we can construct a model of the based on our estimates of the gestation period besed on a fetal grmJth rate estimated from this estimated gestation Therefore our reasoning would be some~hat circular if -Je used thjs to prove the estimate Hmvever we can see how this model fits the rest of the data A good fit would tend to support the model and the estimates on which it is based

For this model He use the curve in 8 to represent births We assume a gestation period of 8 months -rith the unimplanted and implanted periods each lasting 4 months TheTefore v8 draw curves identical to the birth curve but moved backward 4 months for implantation of the blastocyst and 8 months for conception (Fig 9)

A comparison of the peaks in Fig 9 with the data in Figs 1 2 and 3 and the narrative b on those indicates a fairly close fit The data indicat that the peaks may ~ctually occur slightly later than indicated in 9 but in the timing and distances bet7een the peaks fit

The sharpness of the birth curve indicates that the breeding in1plantatimiddotm and pupping peaks mey be more distinct than indicated by the curves in Figs 1 and 2 It is possible to fit curves with more abrupt changes to the data Fig 10 shows the same data lith the curves drawn to more closely fit the model tfuether these curves are more accurate or not is open to debate The data on Hhich the birth curve is based is not that strong The numbers in each month are relatively snall If such distinct did occur it seems likely that field observers ~vould have seen the effects Nost field observations indicate broader peaks that are not gtvell defined

2

G

X

middot ~ -- (-- ) r~ ~ - - -- ~

yen~ r- ~--~

3

i__middot -~---

~

gt

- cshy

Reproduction - 10 - March 1971

~e of _Sexual Maturity

Tables 4 - 7 list the frequency of nmnbers of corpora albicantia and corpora lutea found in females of different areas It should be recognized that the ages may not al-1ays be exact Animals ATTichi tka to become sexually mature Hhen about 4 years old It appbull~ars that more animals may become mature earlier on Adak The information is too limited to dratmiddotJ any definite conclusions from this hmmiddotJever the food availabili ty is probably better at Adak and an earlier age of sexual 1naturity may be a response to better nutrition

Ovulation

From Tables 4 - 7 it can be seen that the maximum number of corpora albicantia for each age roughly the numb~r of years after sexual maturity that some animals ovulated once every year after maturity It can be assumed that many corpora albicantia are invisible macroscopically after several years particularly if they are not remnants of a corpus luteum of pregnancT Therefore it Ls possible that most females ovulate every year This poss ili ty is also supported by the high incidence of large follicles in lactating females (TabL~ 10)

faximum

Tables 4 ~ 7 indicate that at t some females continue to breed at a very old age The samples are teo small to determine 1rhether a t number become unproductive at any point

Ovulation

Many mustelids are induced ovulatoTs

On September 14 1967 a female vJas tsd Hhile copulating She appeared to have Oilnlated shortly before Because sea otters may copulate several times over a period of two or three (Kenyon 1969) ovulation may have been induced by a previous

ficance of

Delayed lon is a charact8ristic of the reproductive cycle of most mustelids Several theories on the ccdv of a delay have been advanced Most assume that spring is the most favorable time of year for survival of nemiddotJborn young but that either a wir1ter breeding season is not favorable or that the presence of a Ttlale vhich occurs only during the

season is for survival of the young of the previous pregnancy (1963) discusse the evolution of delayed implantation in mustelids and points out that there are exceptions even in arctic areas middotJhere time of birth be consLdered more may be born at any tin1e Scheduling the and made possible by delayed mey be advantageous but is not necessary

----

a - o lt bull y bull bull l w laquo bull _ ~ ~

Table 4 Frequency of occurance of Numbers of Corpora Albicantia Amchitkll - June-August 1968

N U M B E R 0 F C 0 R P 0 R A A L 8 I C A N T A

I

~I -AGE -~ 1 I 1084 6 92 5 7(Years) 1 middot shy

- 0-l - ~~ ~- middot~middot- middotmiddotmiddotmiddotmiddot

1-2

2

3

4 1 I

2 (1)

6

middot5

1 (1)1

2 (l) 17

1 ( 1 )

2 ( 1)

18

l (1) I ( 1 ) 9

10 1 ( 1) L ~j( 2 1 ( 1 )

2 ( l) 1

12

11

1 (1)

I I (I)

1

2 ( 1)

13

1 ( 1 )

2

114

115 I116

17

I 18

I I

I

119 I

Numbers inside parentheses =Number of individuals with corpus luteum

11

Table s Frequency of oc~urance of Numbers of Corpora A1bicantfa _Kanaga - October 1968

N U M B E R 0 F C 0 R P 0 R A Al B CANT IA - I I I

AGE 84 I I

5 I l 6 7z 3(Years) 1

l rbull ~middot--

--- shy

0-l

J l-2

~ 2

3

4 2 ( 1)

5 3

Ibullbull 6 6 (3)

4 (4)7

8 4

3 9 2 ( 1 )

j iI 10

I 1 (1)11

12

13

14

1 ( 1)

16

15

17

18

Numbers irisi

-

J

(Plus 2 with corpus 1 uteum but no corpora a 1 b i cant i a)4(1)

1

1 (1)

5 (1)

10(5)

3

3 1

3 (2)

2 ( 1) 3 (3)

1 ( 1)

2

1 ( 1 ) 5 ( 1)

1 ( 1 ) 3 ( 1)

1

3

2 ( 1 )

1 (l)

1 (1)4 ( 1)

2) 1 (1) 3 (3) 4 (1) 12 ( 1 ( 1 )

1 ( 1)

1

2

1 ( 1)

1 ( 1)

1

I I parentheses = Number of individuals with corpus luteum

11

Table 6 Frequency of occurance of Numbers of Corpora Albicantia Mid Pt-~Blind Pt~ Adak- October 1968

N U ~1 B E R 0 F c 0 R P 0 R A A L B I C A NT I A

-1AGE

~

9 1084 75 6Years) 1 2 3 - shy ----- --middot shy~--

- _ Q~L

1-2

(Plus one with_corpus luteum but no corpus a1bicans)2 1 l I I I I I ) (One with corpus luteum but no corpus albicans 3

4 2 (1)

l5

16 1 (1)

1 1)7

8

2 (1)9

10

111

12

I13

14

15 1 (1) 16

17

18

1 ( 1)

1 (l)

1

1

1

2

1

1

1

1(1 )

1

1 (1)

1 (1)

1

I

1

1

1

1

-I

Numbers Inside rentheses = Number of individuals with corpus luteum

1

Table 7- Frequency of occurance of Numbers of Corpora A1bicantia Three Arm Bay - Bay of Islands Adak - October 1968

AGE (Years)

Q-1

t-2

2

3

4

s 6

7

8

9

10

11

12

13

14

15

16

17

18

Numbers

N U ~ B E R 0 F C 0 R P 0 R A A l B C A N T I A

~L 3 4 slE - 1middotshy

7 1~8 __J~l_11 shy

1-

One -Ji th corpus 1uteum but no corpora _a 1bicantia I I I I I I I

One vJi th corpus luteum but no corpora al bicantia

12 I I (Plus one with corpumiddots luteum but no corpora albicantia)1 (1) 3

1 (1) 2 (1)

1 2 (2) l 1

1 (1) 4 (1)

2 (1)

2 1 )

1 ( 1 ) 1

2 (1)

2 (2)

l22 3

3 ( 1)2 21 ( 1)

2 ( 1)

2 ( 1)

l ( 1)1

1 1

1 I ll

I

I I

inside parentheses Nurnber of individuals with corpus luteurn

Reproduction - 11 - March 1971

The sea otter lives in an area of relatively uniform temperatures However storms tend to be more frequent and violent in fall and winter Survival may be better in the late spring and sunrmer Therefore there may be some advantage to birth in the late spring However there does not appear to be a great deal of advantage to a breeding season at any particular time of year

Wright (1963) pointed out that most musteHds lant after daylight begins to increase usually around February This is the period when sea otters implant at the t rate If the time of implantation is influenced by daylight the period when the greatest number of births occur could be shorter than the equivalent breeding period The data do not show this although the possibility cannot be ruled out

The fact that substantial numbers of sea otters are breeding implanting and pupping at all times of the year indicates that hile there may be some advantage to certain events taking place at a particular time of year that advantage is not great enough for a distinct breeding season to evolve

Sea otters exhibit certain characteristics that are comnon to most mustelid breeding cycles but these chfracteris tics are not as well developed as in most es

It is interesting to note that Hright (1963) correlated the molt of Mus with the implanted period TI1e molt began around the

ion and ended around parturi tioD

Sea otters molt all year and some are implanted pregnant at all times At this time e cannot say uhether any between molt and pregnancy exists

Fetal

Kenyon (1969) found that the number of caudally presented fetuses roughly equalled the nunber of cephali presented fetuses He suggests that this indicates a lack of tatim1 for birth occurring in water and infers that sea otters must give birth on land

Fetuses of all 1veights in the present study also appear to be randomly oriented however 97 Class 5 fetuses (1000 g and over) showed 60 percent cephalic ion while only 40 percent shovJed caudal presentation

The orientation of large fetuses should be a better indicator of orientation at birth Smaller fetuses may position Therefore it appears that more sea otters are born head first than tail first

If you accept Kenyons premise that caudal presentation is necessary or at least beneficial to birth in water this more recent information supports the supposition that birth occurs on land The little information available indicates that birth does occur on lando but there have been unconfirmed reports of birth in water

Reproduction - 12 - tltIarch 1971

At least some sirenians are born head first indicating that caudal presentation is not necessary North of Unimak Island a large population of sea otters lives off-shore There is little evidence that any numbers come ashore It seems likely that many of these animals are born in the water

Of 305 implant pregnancies 138 fetuses (45 percent) had implanted in the left horn and 167 (55 percent) in the right horn Most of the difference was in the 1970 Amchitka sample ~fithout this sarnple 48 percent implanted in the left horn and 52 percent in the right horn Kenyon (1969) found an equal number in each horn If a real difference exists it is probably exaggerated by the 1970 A~1chitka sample

Out of 305 implanted pregnencies the point of implantation was on the same side of the reproductive tract as the corpus luteum in all but three instances In one case the corpus lutaum was in the left ovary and the fetus in the right porn In the second case the corpus luteum in the right ovary but the fetus was in the left horn In the third case there was a corpus luteurn in each ovary and tHo fetuses (of different sexes) in the left horn

It appears that blastocysts rarely cross from one horn to the other

Because there is usually a long period of time parturition and the next est_rus is little if any inhibitory effect on ovulation caused by the corpus of the previous pregnancy The ovary producing the largest follicle appears to be random and there does not appear to be the tendency for a pregnancy to be from the opposite ovary from the previous pregnancy In many cases there are many more corpora albicantia in one ovary than in the other

When a loses a pup shortly after birth and enters estrus before the uterus has conpletely recovered and before the corpus luteum has completely degenerated there may be some inhibitory effect Of the ll+ such cases identified six had large follicles in the same ovary as the new corpus albicans and in the opposite ovary It appears that any inhibitory effect by the corpus luteum only lasts for a short time

A few cases bullv-ere found vJhere one ovary was not developed or othenvise not capable of producing ova Because one ovary may support repeated pregnancies such animals may be as productive as those ~vith both ovaries active

Kenyon (1969) reported that most implantations occur within the central third of the uterine horn This held true for the animals in the present study they may implant In one case with a near term fetus the placenta covered the at the base of the horn blocking the exit of the fetus

Reproductive - 13 - March 1971

Birth iltleigh t

Barabash-Nikiforov (1947) listed the vJeight of a ne-v1born sea otter as 2000 g Kenyon (1969) presents the best quantitative data published to this time He found a maximum fetus weight of 1869 g 1-Thi1e he found pups as small as 1020 g those under 3 lb (14 kg) had died shortly after birth He estimated that successful birth weights range from 3 to 5 1b dr 1 4 to 2 3 and for purposes of calculation assumes an average birth weight of 1850 g to 1900 g

The weights of Weight Class 5 fetuses the smallest pups found living in the wild and three pups born in captivity (all died within 24 hours) are presented in Fig 11 These data tend to support Kenyons (1969) estimate of the ~veight at birth Few living pups middotHeighing less than 1350 g or about 3 lb and fevJ fetuses over 2000 g ( 4 5 lb) are found Kenyons upper estimate appears to be slightly high although there are extreme cases in both directions One pup weighed only 2 lb (900 and one fetus could not be weighed accurately on available equipment but weighed over 2500 g

One female with a 4 lb (1816 g) pup still had the placenta attached to the uterine wall and must have given birth shortly before being collected

From 11 it appears that most pups are born when they weigh in the neighborhood of 1800 to 1900 g in most of the populations sanpled Hmvever the and Delarof s indicate a bith Jei of between 1600 and 1700 g The Delarapound sample is srEall but the Tanaga sample the largest containing over a third of the large fetuses collected and was collected at a time of year when the birth rate is very high

This may be a reflection of the phys condition of the adult females and indirectly a reflection of food availab ty There are other factors indicating that the population is declining because of overpopulation

This lov1er birth yeight raises the question of middotwhether the pups are born earlier or whether are in poorer condition when born In an effort to gain some insight into this the fetal of Class 4 and 5 fetuses from Tanaga were plotted against their total lengths (Fig 12) middot Fetuses from other eastern Adak which appears to be a very healthy population were checked this curve fonaed by the Tanaga fetuses In all cases they fell Jell Jithin the limits of the Tanaga data indicating that there is no difference in the of fetuses of a given length It seems unlikely that a reduction in the rate of weight gain vould be completely proportional to a reduction of linear growth It appears that the rate of growth of fetuses is relatively constant but that females in poor physical condition may not be able to support as large a fetus as those in good condition so parturition occurs earlier

This may have sone effect on information on this

Fig 12 demonstrates some other interesting points There is no difference in the ratio and the maximum fetus size of males and females

~

(

~

-s 2 0 C

U)

0

--

ez ()

~) I) middot _t t- (

j -~middot -~

Reproduction - 14 - March 1971

This infeTs although dolt2s not prove an identical grmgtlth rate afld birth weight for males and females The circles on Fig 12 show the ~eight-length ratio of pups These fall below the curve formed by fetus weight-length ratios indicating that pups of a given length are some-Jhat heavier than fetuses of the sarne length This indicates an increase in weight immediately after parturition

Sex Ratio at Birth

Kenyon (1969) found that of 58 fetuses for which sex could be determined~ 45 percent vJere male and percent were female He assumed an equal sex ratio until a larger sample could be obtained

In the present study 111 fetuses were male and 144 ~vere female When this sample is combined with Kenyonts the sex ratio of 313 fetuses is 44 percent male and 56 percent female

Sex Ratio of the

In the process of harvesting capturing animals for transplants and delineating male and femalen areas it has become apparent that there are more females in the population than males Our general ion is that perhaps 60 to 65 percent of the sea otters are females Kenyon (1969) demonstrated that more juvenile males than females are found dead on the beaches of Amchitka lhe age structure of harvested in 1968 indicates that males may not live as long as fetnales

The lower birth rate higher juvenile mortality rate and slightly shorter life expectancy of males could account for the unbalanced sex ratio in the population

The sex ratio may vary from one island to another Presumably there -vmuld be more females in a population that has reached the carrying capacity of the habitat where juvenile mortality is higher There is some indication that males are the first to expand into new areas of unoccupied habitat Therefore in recently populated anas one might find more males However this situation vould be

Lit

The normal litter size of the sea otter is one This has been recorded by almost every observer since Steller Barabash-Nikiforov (194 7) reported twins in utero and mentioned other reports of twin fetuses from the early days of SnoF (1910) recorded a set of neHborn tdns In more recent studies~ both Sinha et al (1966) and Kenyon (1969) found reproductive tracts with two corpora lutee but in all cases only one fetus las present In thousands of hours of observation by many observers no tbullvin pups have been reported

In the present study 24 instances of multiple ovulations were recorded Five of these resulted in hv-in and one in triplet fetuses The information is summarized in Table 8

Table 8 Multiple Corpora Lutea Fetuses

Total Pregnant Females 565

Implanted Pregnancies 316

Unimplanted with 2 CL (corpora lutea) 9

Implanted Pregnancies with ) L CL and 1 Fetus 8

Implanted Pregnancies Vlith 3 CL and 1 Fetus 1

Implanted Pregnancies gtvith 2 CL and 2 Fetuses 5

Implanted Pregnancies middotwith 3 CL and gt Fetuses 1

Total Nultiple Ovulations

I

Percent Nultiple Ovulations 42

Percent of Pregnancies tvith Nultiple Fetuses 19

Corpora Lutea per Pregnancy 105

Fetuses per 102

Reproduction - 15 - lvlarch 1971

From these data it appears that in over 4 percent of the estrus cycles more than one ovulation takes place Of these about half result in the development of more than one fetus In most cases the were of a relatively large size and probably would have been born normally

All multiple fetuses appeared to be the result separate ovulations All had separate placentas and in some cases were different sexes Four tracts with twins had both fetuses in the same horn one had one in each horn and the tract with triplets had one fetus in one horn and t~vo

in the other Another tract had skeletal remains of triplets in one horn that were being resorbed Another tract appeared to have had five fetuses in one horn but they were almost completely resorbed One might infer that physically there is not enough room in one horn to accomn10date more than two fetuses for any length of time

With the exception of the newborn tvins reported by Snow (1910) there are no docu_mented records of female sea otters with tvin pups There are occasional unconfirmed reports of twin pups but as Kenyon (1969) points out a female may tolerate a pup in addition to her ovm but it is unlikely that a female could care for more than one pup It is unlikely that more than one pup survi~es In any case orily 2 percent of the pregnancies produce more than one pup 1middot1ultiple births cannot be a significant factor in the productivity of sea otter populations

The presence of a corpus luteum without gross evidence of implantation t-7as assumed to be an unimplanted pregnancy No attempt ~vas made to locate unimplanted blastocysts As a result e have little information concerning the survival of the blastocyst Of 206 reproductive tracts classified as nulliparous on the basis of the appearance of the uterus 12 had one corpus albicens and one had two Some of these may have been large attritic follicles Others may have ovulated and even conceived but implantation did not occur All of these cases represent the failure of the antmals first estrus

A total of 16 resorptions or possible abortions were identified (Table 1) The follmJing is a brief descdption of these ~7i th possible causes of some

Des

Resorption of blastocyst or ovum (corpus luteum 4 degenerating no evidence of ion)

Failure at time of (possibly because 1 of growmiddotth inside horn adjacent to implantation site)

Resorbed or aborted fetus (implantation at end of horn) 3

Resorbed fetus (umbilicus tvTisted tightly) 1

Resorbed fetuses (three or more fetuses in one horn) 2

Resorbed fetus (cause unknmm) 5

Reproduction - 16 - March 1971

Six of these resorptions may have been caused by a lack of room for growth of the placenta because of crowding from other placentas a growth or the end of the uterine horn These six and the one vith the twisted umbilicus are probably the result of random events or accidents

The numbers of resorptions are too small to accurately measure the frequency of such occurrences although the relatively large number in the Tanaga sample (Table 1) may be a reflection of the physical condition of the animals Part of this number is probably due to the fact that there are more pregnant animals in the sample In general the samples with the most pregnancies also conta~~ned the most resorptions Half of the Tanaga resorptions can be attributed to random events but five vere still due to unknown causes Eight of the 10 resorptions fo1md on Tanaga came from a 20-mile stretch of shore about 25 percent of the harvest area

While concrete conclusions cannot be drawn from these data they do raise the possibiliy that under certain conditions 5 percent of the pregnancies may fail

near Parturition

Collecting methods were such that the presence of a pup with a female was often overlooked Therefore the lack of a pup with a post-partum or lactating female did not mean that the pup had died However 14 females shmving of recent pregnancy were estrus presumably having lost their pup shortly before or shortly parturition

Again the magnitude of this mortality cannot be determined but it appears to be of the same order as the in utero mortality

after

Reproductive tracts were subjectively classified as post-partum or anestrous on the basis of degeneration of the corpus luteum (or appearance of the corpus albicans) the size of the horn of pregnancy and appearance of the placent scar In an effort to get an idea of how long it takes for the tract to return to normal the size of pups accompanying females in each category is listed in Table 9

It appears that females pass the subjective boundary from post-middotpartum to anestrus Hhen the pup about 10 lb and is around 75 to 80 em long

We do not know a great deal about the early growth of a sea otter pup but judging from the cur1e e9tablished based on cementum deposition of older animals and from tooth eruption a 10 lb pup is probably in its second month of life

Table 9 Pups Accompanying Pos artum Females

Sex llleight

M 2 lb F 3 F 4 M 5 F 55 F 625 F 775 M 10 F 11

Total Length

47 em 52 56 60 65 65 70 81 (borderline post partum-anestrous) 82 (late post-partum)

Pups Accompanying Anestrous Females

F 10 lb 78 em M 11 79 M 12 8l F 13 83 M 13 92 H 16 83 M 17 91 1 19 87 M 19 90 F 21 107 F 22 96 1 22 101 M 2Llmiddot 97 1 26 100 F 28 103

Reproduction - 17 - Narch 1971

Frequency of Breeding

Steller (1751) reported female sea otters with ne~vborn pups also accompanied by another pup that seemed to be no more than a year old Nurie (1940) recorded an instance of copulation by a female accompanied by a pup and Jones (1952) caught a young pup simultaneously gtvith a copulating pair Lensink (1962) felt that few females breed until the pup is a year old but mentions females with small pups also accompanied by large pups

I believe that some of thes2 observations may be misinterpreations of the situation Females will occasionally leave small pups for a time often near other animals The pups Lensinkmentions would be 2 years old and probably veigh 30-35 lb Sea otters are gregarious and subadults may remain near other animals appearing to be accompanying their mother I suggest that most of Stellers and Lens inks large pups ere such cases

Females gtvi th pups probably do breed occasionally but the most recent evidence indicates that this rarely occurs Kenyon (1969) records no instances of females accompanied by pups copulating and states that he found no females- knm7n to be accompanied by a pup that were This holds true for the animals collected in the present study However the nature of the harvesting operation is such that many females accompanied by pups were not recorded as such Therefore we are forced to look at lactating and assume that they were accompanied by a pup at the time of collection or shortly before le 10 sm1marizes the reproductive condition of the lactat females In the 1967 and 1968 samples some animals vith enlarged marmary middotmre listed as lactating As a result some females with near tenrr fetuses Here included Because this condition is considered to be associated with the unborn pup rather than a previous pup these animals were with the anestrous animals in le 10 In 1969 and 1970 the presence of milk was used as the sole criterion and no term animals we-rmiddote in the sample

Of 246 lactating females only one had an ted fetus (excluding the near term animals from 1967-68) Several others had small corpora lutea indicating that they recently become or possibly had large

vhich had luteinized but tvere not actually pregnant The remainder had follicles

This information that accompanied by a pup rnay enter proestrus and even ovulate but that they only rarely mate That they enter estrus is subs iated by the f2ct that the number of corpora albicantia in many tracts equals the age of the amplimal minus the three years prior to nBturity (see les 4-7) This indicates that large follicles tend to develop year after sexual nl8turi ty is reached whether the female is accompanied by a pup or notlt

Some of those lactating females with s1all corpora lutea may have recently weaned or othenvise lost a pup and are already pregnant again Malkovich (1937) took a pup least 3 months old) away in cap She mated 12 days later here she had the male It appears that a female enters estrus shortly after Heaning

Table 10 Reproductive Condition of Lactating Females

Location

Bay of Is Adak I Sept 1967 9 1 100

Allchitka I Sept-Oct 1967 17 2 105

Mid Pt - Blind Pt Oct 1968 19 3 136 Adak I

Bay of Is Adak I Oct 1968 22 4 154 3

Kanaga I Oct 1968 32 s- 135

Al1chitka I July-Aug 1969 4 2 333

iTanaga I May 1970 56 10 152

Delarof Is May 1970 18 3 143

Amchitka I May 1970 36 3 77

---~--middot--middot----~-~----~----middot~-middot--~----~----- -------~-~ middotmiddot--middot-shy

TOTAL 213 33 134

Anestrous or pregnant with term fetus

]_ Large follicles or corpus luteum present

3 Includes one implanted pregnancy with small fetus

Reproduction - 18 - March 1971

The question of hml long it takes for a female to enter estrus again after losing a small pup arises Eleven tracts of the 1970 sample and ttvo of the 1968 Kanaga Sampnple shmJed signs of being pregnant recently) but were in proestrus These animals had an enlarged horn usually middotlith a slightly pigmented area but no actual placental scar a11d a recently fonned corpus albicans However they also had enlarged follicles and the tvalls of the uterus were typical of proes trous animals Another animal was similar but appeared to have already ovulated and a corpus luteum was present In these cases the female has lost a pup either through abortion or vithin the first veeks after parturition and has started into another estrus cycle Three or four of these had follicles vhich may have started to become attritic This may be because the uterus had not recovered sufficiently from the last pregnancy

The implication of this information is that with a normal pregnancy and successful rearing of a pup the female becomes pregnac1t only after the pup has been weaned This n~y be a year after parturition However if the pregnancy fails or if a pup dies even at birth the female may become pregnant again in a few weeks rather than _waiting the full year

Kenyon (1969) suggests that the period middotbetween and the next estrus may long because he found animals that tvere not lactating but had a faint placental scar and an indistinct but recent corpus albicans These animals ltJere cLtssi as anestrus in the present study They are undoubtedly bet1veen veaning of their last pup and the next estrus The number of suh animals is t durirg Sspte12~ cnd October At this time the rate of estrus increases a1d the number of anestrous animals decreases sharply the next feJ months Therefore many of these anestrous anirrals become pregnant 1lithin a month or two Kenyon (1969) suggests that if the female keeps the pup for about a year that the period between births may be smreltmiddotJhat greater than two years He is assuming a 12-13 month gestation period However the present study shows that the estimate of a 12-13 month tation period was based on an inadequate sample Estimates in the present study put this period at closer to 8 or 9 months Presumably the anestrous nonlactating females are in the 3 to 4 month period

This does not change Kenyons (1969) point that the and the next estrus may long It indicates that in a normal cycle it may be months Hmvever as it vvas shmvn above~ it is possible for a female to enter estrus very shortly after losing a pup

We have demonstrated that sea otters breed at all times of year and it is possible for a female to become pregnampJt again after losing a pup sooner than she vould have if the pup had survived HmmiddotJever we have also demonstrated that annual of b and pupping occur and that these are similar in different years and in different populations The assumption can be made that the average length of the entire reproductive cycle is some multiple of a year

Kenyon (1969) assumes that tbe pup remains with its mother for about a year ~-Ieights and measurements cementum deposition skull changes~

Reproduction - 19 - March 1971

observations in the wild and in captivity all support this assumption although it cannot be said with certainty that it averages 12 months It could be s tly ITDre or less

Assuming about 12 months for rearing the pup 8 to 9 months gestation plus a fev months rest bet~veen and the next estrus the average reproductive cycle takes two years

This is supported by Fig 4 which shows that slightly over half the mature females are pregnant in a year Kenyon (1969) pointed out that his January and Harch samples gtvere biased against females with pups a1d therefore in favor of pregnant females because of selective hunting This bias applies to the fall sa~ples as well The summer samples were biased similarly because pregnant females appeared to be less resistant to handling during capture Therefore the percent of pregnant females is shown to be higher than lt actually Has -rhile no correction factor can be applied to all samples the evidence indicates that the percent of pregnaJt females is usually around 10-middot15 lmrer than in the sample For example the 1970 Alnchitka sltLrnple t-ras less biased and was about 9 percent lower than the Tanaga sample vrhich is l)nm-rn to be more biased Also on a June skiff survey 7 percent of the animals counted had pups that were readily identified If ~re assune that this is the nunber of animals avoided it can be calculated that 15 of the sexually mature females all of vhich are t middotTen~ avoided Actually this will vary with the hunter weather time of year etc Hmvever it indicates the order of magnitude of the bias

Using this as a rough correctton for the data in 4 we find that about half or slightly fe-Jer of the sexually mature females are pregnant each year or as indicated ly the average female has a pup every two years

Reproduction - 20 - March 7 1971

Conclusions

1 While sea otters may breed or pup at any time of year there are seasonal fluctuations in the illliTlbers of females in each reproductive stage The pattern of these fluctuations remains the same from year to year and population to population

2 Breeding activity is highest in September October and November

3 More implantations occur during January February and March than at other times of year

4 The birth rate is highest during April Y~y and June

5 Tlvo and a half to three times as many females breed during the peak breeding months as during the months of lowest breeding activity

6 Similarly two and a half to three times as many females pup during peak pupping months

7 During the period of lovest b activity seually mature females breed at a rate of 2 to 3 percent per month During the peak of breeding 7 to 10 percent per month breed

8 During the period of lov1est pupping activity 2 or 3 percent per month pup and during the peak of pupping 7 to 10 percent per month pup

9 The gestation period about 8 to 9 months on the average About half of this time is ~~ ted period

10 Female sea otters become sexually mature ihen 3 to 4 years old

11 Females may ovulate on an average of once a year after sexual maturity is reached

12 Females continue to breed at Cn old age

13 Sea otters retain certain characteristics common to the reproductive cycles of many other mustelids however exceptions are common Scheduling of the reproductive cycle during the year may have some advantages or it may be a trait that has not been entirely lost through evolution

14 Sixty percent of the near term fetuses are cephalicly oriented and 40 percent are candllly oriented

15 Sea otters probably birth both on land and in the water

16 Blastocysts rarely cross from one uterine horn to the other

17 Consecutive pregnanctes may occur in the same horn The side of pregnancy appears random

Reproduction - 21 - March 1971

18 Implantation usually occurs in the central third of the horn but it may occur any~rhere in the horn

19 Birth weights may range from 900 g to over 2500 g however most pups weigh 1800 g to 1900 g at birth

20 Females from areas gtvhere the population is declining because of food shortages may pup earlier and as a result the pups 1 birth weight may average 200 g lmver than nonnal

21 Male and female fetuses gr01middot1 at the same rate ampJd are the same size at birth

22 Body condition of the female has little effect on the growth rate of the fetus

23 There is a rapid ~Jeight gain shortly after parturition

24 The sex ratio at birth is 44 percent male and 56 percent female

25 The normal litter size is one hmmiddotever in 2 percent of the implanted pregnancies two or fetuses survive to near term

26 Multiple ovulations occur in 4 percent of the estrus cycles

27 es do not appeErc- to be able to more than two fetuses in a single horn

28 Up to 5 percent of the p may fail dne to in after implantation

29 An unknown but probably significant number of first pregnancies fail before implantation

30 The uterus is usually considered recovered from a pregnancy when the pup vleighs about 10 lb and is probably in its second month of life

31 Sexually matu1middote females normally have one pup every two years

32 Females vrith a pup may ovulate but rarely mate

33 Slightly less than half of the sexually mature females are in any given year

34 A female may enter estrus within a few weeks of losing a pup Consequently she may become agaln gtvi thout waiting the same length of time as if the pup hacl survived

35 Hhen a pup is weaned normally there is a longer period perhaps 3 or 4 months between weaning and the next estrus

Reproduction - 22 - March 1971

CITED

Barabash-Nikiforov I I 1947 The sea otter (Kalan) Soviet Ministrov RSFSR Translated from Russian by Dr A Birron and Z S Cole Israel Program for Scientific Translation 1962 227 pp

Hugget A St G and W F Widdas 1951 The relationship betlveen marmnalian foetal -reight and conception age Jour Physiology 144306-317

Kenyon K W 1969 The sea otter in the eastern Pacific Ocean USFWS North American Fauna No 68

Malkovich T A 1937 The sea otter in captivity Priroda No 381-87 Translated by J T Naximovitch 1966 Fisheries Research Board of Canada Nanaimo BC Translation Series 657

Sinha A A C H Conavay and K t-J Kenyon 1966 Reproduction in the female sea otter J Hildl Ngrrit 39(1)121-130

Snow H J 1910 In forbidden seas Edward Arnold London 303 pp

Wright P L 1963 Variations in reproductive cycles in North Arrierican mustelids In Enders A C ed Delayed tation Univ of Chicago Press 318 pp

  • SEA OTTER STUDIES - 1970
    • Surveys
    • Harvests and Transplants
      • Sexual Segregation in Sea Otter Populations
      • Reproduction in the Female
        • AERIAL COUNT OF SEA OTTERS- SOUTHSIDE OF THE ALASKA PENINSULA
          • Description of Conditions During March 1970 Aerial Count of Sea Otters
          • Discussion of Results
          • Sanak Island-Sandman Reefs
          • Shumagin Islands
          • Sutwick Area
          • Augustine Island-Cape Douglas
            • SUMMARY OF SEA OTTER SURVEYS
            • Amagat 1 to False Pass Ikatan Peninisula Cape Lazaref
            • Sanak Is
            • Deer 1
            • Pavlof Is
            • Northern Shumagin Is
            • Cold Bay 1
            • Kupreanof Peninsula
            • Mitrofania 1 to Kupreanof Pennisula
Page 2: Sea otter studies - 1970 - Alaska

Sea Otter Studies - 2 - ]1archt 1971

Location of Pregnancy Birth Weight Sex Ratio at Birth Sex Ratio of the Population Litter Size In Utero Mortality Mortality at or near Parturition Recovery of the Uterus after Parturition Frequency of Breeding Conclusions Literature Cited

AERIAL COUNT OF SEA OTTERS- SOUTHSIDE OFTHE ALASKA PENINSULA

March 1970

Between March 16 and March 27 1970 an aerial count of sea otters was rnade along the south side of the Alaska Peninsula from Shaw Island to Cape Lazaref on Unlmak Island Sanak Island the Sandman Reefs the Pavlof Islands northern Shumagin Islands and Sutwick Island were included A BLM Aero Commander N6392U was flown by Cal Ward at an altitude averaging 150 feet and an airspeed bullgtf 100 mphmiddot Observers were Karl Schneider next to the pilot and Jim Faro behind the pilot This is the same technique aircraft pilot and observers as were used in the 1969 Aleutian count The count on March 27 was made in a Gruman Goose 1r1ith Robert Wolfe substituting for Faro Pups with their mothers were not counted

Because of conditions of visibility and distribution of animals all of the counts made are considered to be low

In general the variability in factors influencing this type of count is so great that the results are not rei iable for population estimates or for determining short-term fluctuations in dense populations However aerial counts should be useful to determine general distribution and abundance and to follow large changes in population size Such changes are occurring where relatively dense populations are expanding into habitat that is either unshyoccupied or sparsely occupied Aerial counts are also the quickest way to familiarize new personnel with large areas of habitat

The following is a 1ist of definitions of terms used to describe counting conditions Application of these terms is completely subjective and based on the writers experience

Excellent- Surface of water relatively calm Usually overcast with 1ittle surface glare Single animals easy to spot at a distance

Very good - Surface may be slightly choppy but not enough confusing reflection to prevent spotting of animals off shore

Good - Off shore animals may be difficult to spot but single animals near shore and in kelp beds and small pods everywhere are readily spotted

Fair -Usually choppy or surface glare Single animals ~in kelp beds or in the lee of rocks or shore and most groups of animals relatively easy to see Other single animals and some pods in deep bays in the shadow of cliffs or off shore may be missed

Poor -Single animals difficult to spot and many pods may be missed but conditions still good enough to get a rough idea of the distribution of animals

The conditions encountered for each area are listed below

Description of Conditions During March 1970 Aerial Count of Sea Otters

March 20 Aniakchak Bay to Chignik Lagoon (1022 AM- 1240 PM) Conditions were generally fair with a heavy chop The south side of Sutwick Island and the area between the island and mainland were poor with heavy surf Visibility was very good inside Kujulik Bay however many animals were scattered throughout the area On two passes ~cross the bay 12 and 27 were seen therefore many were probably missed Similar scattering occured in Chignik Bay The overall count for the area is probably low

Pavlof Bay to Cold Bay (130PM-200PM) Conditions poor with heavy chop and squalls The count was superficial however a second superfi cia 1 count of this a rea was made on March 21 and only one otter was seen in Cold Bay

Cold Bay to Caee Lazaref (310PM -420PM) Conditions very poor fair in a few spots Heavy chop dense kelp

March 21 Northern Shumagin Islands (943AM- 1137 AM) Poo~ with chop on windward sides Fai_r on leeward sides (westerly wind) however air turbulence forced us away from high cliffs Heavy surf created very poor conditions on the south side of Unga

Pavlof Islands (1145 AM- 125ff PM) Conditions similar to those in the Shumagins

Chignik Lagoon to Pavlof Bay (305 PM- 535 PM) Conditions varying from fair to poor with chop and surface glare Count concentrated on points and better looking areas Most deep bays not completely surveyed

March 22 Sanak Islands (849AM- 1020 AM) Conditions fair to poor with chop on south side and surface glare on north side Animals scattered and very difficult to see Many otters on rocks Count is probably very low

Sandman Reefs ( 1025 AM~ 1115 AM) Conditions similar to Sanak With the exception of one pod of 450+ the animals were widely scattered and very difficult to pick out Only those very close to the aircraft could be seen Bad squalls moved in and the wind increased preventing a complete count of Deer Island and the eastern half of the reefs

March 23 Wide Bay to Aniakchak Bay (1123 AM- 140PM) Conditions generally good at first becoming fair after Cape Providence with a light chop and some surface glare Otters in Amber and Aniakchak Bays were widely scattered and many may have been missed

Port Heiden to Naknek_ Superficial counts were made from Ugashik Bay to Naknek on March 19 and from Port Heiden to Egegik on March 23 These were made under poor conditions by flying outside the surf line The water was choppy and full of silt No sea otters were seen

11arch 27 Shaw Island to Puale Bay (1030 AM - 1205 PM) Conditions very good to excellent High overcast and very little wind down to Katmai Bay After this the conditions deteriorated rapidly to fair and the count had to be stopped because of a combination of bad weather and lack of fuel As shown on the charts the count was concentrated on the points and islands however the good visibility allowed us to see well into the bays We feel certain no concentrations were missed

The numbers of sea otters counted are presented in the Table Specific locations and numbers are shown on the charts Where coverage of an area was not complete the approximate path of the aircraft is shown

Discussion of Results

Surveys along the Alaska Peninsula have been fragmentary and most of those made were done under less than ideal conditionsmiddot There are a number of reasons for this Weather tends to be poor along the south shore where squall5 form along the mountains Areas where an aircraft can refuel are north of the mountains and there are relatively few passes therefore range of the aircraft is a problem Also there is much shallow water off shore in the area from the Shumagin Islands to Sanak Island Coverage of this area is difficult and many animals are undoubtably missed

The present survey covered most of the area however conditions were such that most of the counts are probably quite low Despite these problems and a lack of continuity we have a fairly good picture of the recovery of sea otter populations in this area

Reports from various individuals are available from the 1930s and 1940s 11ajor surveys by the Fish and Wildlife Service in 1951 (Jones) 1957 (Lensink) 1962 and 1965 (Kenyon and Spencer) covered at least portions of the area In 1969 the southern Shumagin Islands were counted by the Alaska Department of Fish and Game and the present survey covered most of the rest of the area

Basically there are four distinct population nuclei in the area These centered around the Sanak Island-Sandman Reef area the Shumagin Island area the Sutwick Island area and the Augustine Island-Cape Douglas area The following discussion is based on information from reports by Lensink and Kenyon and from Alaska Department of Fish and Game surveys middot

~anak Island-Sandman Reefs

Small numbers of sea otters have been reported at Sanak Island since 1922 No reports came from the Sandman Reefs until 1942 In 1948 27 were sighted at

----Cherni Island The 1951 aerial survey showed 65 around Sanak and 97 in the Sandman Reefs In 1957 251 were seen around Sanak and 508 around the Sandman Reefs In addition two were seen in the Pavlof Islands however few were on the mainland In 1962 548 were counted in the Sanak area and 638 in the Sandman Reefs None were reported from the mainland or Pavlof lsland~however

much of the increase was in the northern area around Deer Island The 1962 survey was probably the best being made under excel lent conditions

AERIAL COUNT OF SEA OTTERS MARCH 1970

Partial or Location Count Date Camp 1ete Count Visibility

Cape Lazaref - Cold lsecty_

Cape Lazaref 3 320 Complete Poor

lkatan Peninsula 71 320 Complete Poor

False Pass - Amagat I 66 320 Camp 1ete Poor

Cold Bay 321 Partial Poor

TOTAL 141

Sanak Islands 239 322 Camp 1ete Fair to Poor

Sandman Reefs

Clubbing Rocks 12 322 Complete Fair to Poor

Cherni I amp Rocks 495+ 322 Complete fair to Poor

Goose I 7 322 Complete Fair to Poor

Hay I 18 322 Camp lete Fair to Poor

Hunt I 2 322 Complete Fair to Poor

Deer I 322 Partial Fair to Poor~

TOTAL 568+

Pavlof Islands

Inner 11iasik 2 321 Camp 1ete Fair to Poor

Outer lliasik 16 321 Camp 1ete Fair to Poor

Goloi 2 321 Complete Fair to Poor

Dolgoi 67 321 Complete Fair to Poor

Poperechnoi 29 321 Complete Fair to Poor

Ukolnoi 2 321 Complete Fair to Poor

yenWosnesenski 4 321 Complete Fair to Poor

TOTAL 122

Partial or Location Count Date Comp 1ete Count vis ib j 1i ty

Northern Shumagin Islands

Unga 184 321 Complete Fair to Poor

Popof 52 321 Complete Fair to Poor

Korovin 46 321 Complete Fair to Poor

Karpa __2t 321 Complete Fair to Poor

TOTAL 286

Cold ~ to Beaver ~ 0 320-21 Partial Poor

Beaver Bay to Kupreanof Pen

Beaver Bay 2 321 Partial Fair to Poor

Cape A 1 iaks in 4 321 Partial Fair to Poor

Guillemot I 16 321 Partial Fair to Poor

__1Kupreanof Peninsula 321 Partial Fair to Poor

TOTAL 23

Kupreanof Pen to Castle Cape 0 321 Partial Fair to Poor

AntJrrCastle Cape to-1-Ji e ~

Castle Cape-Castle Bay 16 321 Partial Fair to Poor

Chignik Bay 118 320 Complete Fair

Nakchamik I 5 320 Complete Fair

Hook Bay 13 320 Complete Fair

Cape Kum 1i un 88 320 Complete Fair

Kujul ik Bay 1199 320 Complete Very Good

Univikshak I 62 320 Complete Fair

Cape Kuml ik 54 320 Complete Fair to Poor

Sutwick I 14 320 Complete Fair to Poor

Cape Agutka 323 Complete Fair~

TOTAL 1766

Partial or Location Count Date Comp 1ete Count vis i b i 1i ty

Cape Kunmik 13 323 Complete Fair

Naka 1i kok Bay amp offshore islands 16 323 Complete Fair

Chiginagak Bay 5 323 Complete Fair

Agripina amp lmuya Bay 105 323 Complete Good

Wide Bay to Puale Bay N 0 T S U R V E Y E D

Puale Bay to c Kubugakl i __]_ 327 Partial Fair

TOTAL 146

Kashvik Bay to L Chiniak 0 327 Partial Very Good to Excellent

Cape Chiniak to Shaw _I

Shakun Is amp Rocks 71 327 Partial Very Good to Exce 11 ent

Kiukpal ik J to Shaw 1 0 327 Partial Very Good to Excellent

TOTAL 71

In the present survey which was made under relatively poor conditions 239 were seen in the Sanak area and 568+ in the Sandman Reefs The latter count Was incomp 1ete and conditions were such that the number of otters was more a function of time spent counting rather than area covered Obviously many were missed As a result not much can be said about total numbers The main change evident is that 141 were seen along the mainland and eastern portion of Unimak Island and 122 were seen in the Pavlof Islands This indicates that a fairly extensive movement northward and along the Peninsula is occuring

A remnant population probably remained along the south side of Sanak Island in the early 1900bulls By the late l940 1 s they began spreading into the Sandman Reefs This northward expansion has continued to the present time There is still unoccupied or sparsely populated habitat along the mainland shore and in the Pavlof Islands There should be continued expansion of this population for a number of years although the numbers around Sanak Island and the western Sandman Reefs may have already reached a peak

With the arrival of substantial numbers in the Pavlof Islands this population is probably on the verge of mixing with the Shumagin Island populshyation No doubt some individuals have moved back and forth in the past but the two populations are almost continuous at the present time

As in past surveys few otters were seen around the north side of Sanak Island and Caton Island This is probably poor habitat for otters The main population is around the south and west sides The higher count on the west end is probably more a result of intensive searching rather than a higher population

Shumagin Is 1 ands

This has been considered the largest of the four populations in the Alaska Peninsula area The most recent counts have not been adequate enough to show any major increase in numbers in the last decade however major changes in distribution have occured which are very similar to the pattern mentioned in the Sanak-Sandman population

There were occasional reports of sea otters in the Shumagins in the 1930s By 1947 Victor Scheffer estimated that 500 1 ived around Simeonof Island In 1953 Hooper counted 633 around Simeonof and Little Koniuj i Island The 1957 survey showed 1829 Most of these were- in the southern islands Five individuals were scattered in the northern islands and one was on the mainland shore near Elephant Point The 1962 survey totaled only 1352 The animals were scattered well off shore and the count was low However the count on Nagai increased from 149 to 338 indicating a continued northward expansion

In 1969 1510 were counted in the southern islands under relatively poor conditions An additional 286 were counted in the northern islands in the present survey and 23 were seen along the mainland north of the Shumagins Again survey conditions were not good

There is a very clear expansion of the population from a center near Simeonof Substantial numbers now occur on all the islands and repopulation of the mainland is occuring The population in the northern islands should continue to increase and most of the habitat on the mainland is not occupied

There is no evidence of an increase in total numbers since 1957middot However the last two surveys have been less than ideal and the large shallow off shore areas have not been covered adequately Considering the survey conditions and the obvious extentions of range it is almost certain that the populations from San-ak Island to the Shumagins have continued to increase In general it appears that the southern islands and reefs have become fully populated and the northern areas are just developing significant populations The entire a rea may be comp 1ete 1 y repopu 1 a ted in the next 10 years This wi 11 depend largely on the status and potential of the off-shore areas

Sutwi ck Area

Reports of sea otters near Sutwick Island are available since 1936 This population was far removed from other populations and is probably a remnant left after hunting ceased In 1951 Jones counted 388 between Cape Kuml iun and Cape Kunmik Most were near Sutwick Island In 1957 889 were counted Nineteen of these were between Cape Kunmik and Cape Providence indicating some northeastward expansion In 1962 949 were seen with individuals straying as far as Cape lgvak and one between there and Cape Kuliak In 1965 a stray otter was seen at Kinak Bay A major shift in the population from Sutwick into Kujulik Bay occured This may be the result of time of year and weather

In the present survey 1766 were counted between Castle Cape and Cape ~~~gtf~aip the majority were in Kujulik Bay Large numbers have moved

~~ranging as far as Cape Kubugakl i The area from Wide Bay to Puale Bay was not surveyed because of weather

The total count for the population was 1912 even though conditions were less than ideal throughout the bcunt and we feel many were missed It is possible that the increase in numbers is entirely due to reproduction assuming a 10 percent annual increase However it is difficult to compare surveys

There is still room for expansion in both directions from this population The greater movement to the northeast is probably the result of better habitat The population around Kujulik Bay is very dense and a large movement of animals may occur if competition for food becomes serious Otherwise we should expect to see continued steady expansion into adjacent areas for a number of years

to come

Augustine Island-Cape Douqlas

For some time a population has existed around the Augustine Island area at the mouth of Cook In 1 et In 1948 approxImate I y 50 sea otters vvere reported from Augustine Island Reports of sightings have been made from Shaw Island to Tuxedni Bay In 1957 Spencer counted 40 at Augustine and one at Shaw Island Lensink counted 52 on Augustine in 1959 but he considered it a poor count In 1965 Kenyon counted 18 on Augustine and 101 in the Shaw Island-Cape Douglas area In March of 1969 Loren Flag saw 62 around Augustine and in May 1969 Jim Faro counted 130 another observer saw about 30 some of which were probably not tallied by Faro

On the present survey Augustine Island could not be surveyed because of weather No otters were seen around Cape Douglas but 71 were seen in Shakun Islands and rocks near the abandoned village of Kaguyak These were probably from the Cape Douglas group

It appears that the animals move about in the area To date no complete survey of the area has been made at one time Until this is done 1ittle can be said about the population other than that several hundred probably occur there

The following table is a summary of sightings and counts made by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game These data were collected by different individuals using different types of aircraft under varying conditions of weather A strict comparison of numbers should not be made from this table

SUM

MAR

Y OF

SE

A OT

TER

SURV

EYS

(Com

pile

d by

p

op

ula

tio

n f

rom

L

ensi

nk

(196

2 T

hes

is

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enyo

n 0

96

2 amp

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5 R

epor

ts

amp M

anu

scri

pt)

an

d AD

FampG

Dat

a)

Lo

cati

on

P

re

1951

19

51

1957

19

59

1962

19

65

1969

19

70

AU

GU

STIN

E -

C

DOUG

LAS

Aug

usti

ne

Isla

nd

A

ppro

xim

atel

y 50

(1

948)

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1

amp

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glas

Are

a S

igh

tin

gs

from

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aw

Is

to

Tux

edn

i B

ay

TOTA

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40

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41

52

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18

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119

130+

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130+

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71

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INSU

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c

Kul

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-

c

lgva

k

c

lgva

k -

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mik

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amp

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a M

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llan

eous

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ts

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ay

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uml

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ce

1936

C

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355 12

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139

197 14

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53

101 62

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1962

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65

1969

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70

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1969

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Hook Bay Chignik Bay

13 118

-1 shy

shy

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I +

Castle Bay - Castle Cape 16 Nakcharriik I 5 -- Katmai Reef 0

Total 152

middot

~ middot-

bull -middot

bull

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Cape Agutka Cape Kum 1 i k Sutwick I~ Kujul ik Bay Cape Kum 1 i un Unavikshak 1

Total

X

-------8

197 54 14

1199 88 62

1614

X

E c 1 ~

(l) a 10

u

0 ---_ +-

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gt-sect OJ al middotshy

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)

r r -middot

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i

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I I

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l bullt

middot

99

10

i

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9

75

~-

MARINE MAMMALS SURVEY - BRISrOL BAY TO AKUTAN ISLAND June 2 1970

OBSERVER J Vania J Faro

PILOT George Tibbets Jr

AIRCRAFr Piper Azetex

middotWEATdER Partly cloudy throughout flight wind 10 miles per hour or less visibility generally excellent One bad area and that was at Akutan Island Air was turbulant there and we were unable to go completely around the island because of fog in Akutan Pass Water had slight ripple throughout flight and ocassionally there was glare but overall conditions for sightinmiddot=r animals was good

Departed King Salmon 1115 AM Returned to King Salmon 745 PM

FLIGHT PA~1 middotAfter leaving King Salmon we went over to the north side of the Alaska Peninsula and proceeded down the coast flying at 300 to 600 feet of altitude Generally we flew aboumiddott one mile off the beach At Port Heiden Moller and Cinder River we checked the outer sand bars for seals and some of the inner bars where I knew seal hauled out

We did not cover Izembeck Laroon very well Instead we flew offshore and checked Amak Island We then fueled up at Cold Bay Leaving Cold Bay we continued dmvn the north side of the peninsula (sea otter sighted) down to Cape Sharichef and crossed over to Akutan Island flying along the south side of it We were able to get around Cape Morgan and fly doNn the beach a few miles but then had to turn back becausmiddote of fo9

The flight path was then eastward to the north side of Tigalda Island At Ugamak Island we circled it completely flying at about 300 to 500 feet We completed the survey at this point and returnt~d to King Salmon generally following the route we had taken on the flight down

SEA OTTER SIGHTINGS

Seven pods of sea otter were seen south of Amak Island These were photographed and later counted from middotthe photographs Beshycause of the density in the larger pods and the fact that some animals in the smaller pods were diving when the pictures were taken some individuals may have been missed

The number counted in each pod is as follows

1071 520 357 68 36 75 30

TOTAL 2157

Several hours later the pods had drifted four or five-miles northeastward (see map) bull In addition 1 one sea otter was seen at Amak Island and sixteen near Tigalda Island

171

-amp

7

7

7 58

57

53

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8 A

l

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51 ~9

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87

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t 32

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54

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middotmiddotmiddot-

53

51

52

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48

bull9

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54

58

54

53

51

56

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s 43

36 ~

44

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39

3

6

313

8

4

44

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53

54 ~

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61

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shy

SEA OTTER COUNT BARREN ISLANDS and SHUYAK - AFOGNAK ISLANDS

June 9 1970

On June 9 1970 an aerial survey of sea otter was flown in the Barren Islands and in the AfognakShuyak Islands area A Gruman Goose was used with Schneider serving as the only observer Offshore coverage was poor Conditions of visibility were as follows

Barren Islands- 1140 AM to 1225 PM- Water calm moderate surface glare Conditions verygood Count complete

Ban Island to Pernosa Bay- 1255 to 135PM- Conditions similar to Barren Islands but surface glare worse in spots Conditions generally good

Marmot Island to Latax Rocks - 255 to 345PM- Increased ripples on surface Glare Bad Conditions fair Count around Sea Otter Island complete although some otter may have been scattered offshore Remainder of count very superficial

Area

BARREN ISLANDS East Amatuli Island West Amatuli Island Sugarloaf Island Ushagat Island Rocks south of Ushacat Sud Island Nord Island

AFOGNAK-SHUYAK ISLANDS Ban Island-Shuyak Strait Pernosa Bay Marmot Island Sea Otter Island area East side of Shuyak Jest side of Shuyak Latax Rocks-Dark Island

TOTAL

TOTAL

Number of Sea Otters

0 2 0

76 200

29 0

307

18 6 3

121 0

33 26

207

Complete or Visibility Partial Count

Very good Complete ll If II

It II

II

II IJ

Good Complete II

Fair Partial If Complete II Partial II II

II

i o j

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9

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64

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675

6

57

55

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61

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41

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13

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51

SEA OTTER COUNTS - KENAI June J970 to January 1971

Carl Divinyi flew aerial surveys of seals along the outside of the

Kenai Peninsula on a monthly basis He recorded all sea otters sighted

on these surveys The surveys did not cover the entire coast line and

the type of aircraft weather conditions and observers varied from one

survey to another Therefore the seperate counts cannot be compared

However when viewed in total they indicate the areas used by sea otters

and the relative abundance of otters in each area

The attached table presents the counts by broad areas

--

SEA

OTT

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CO

UN

TED

ON

A

ER

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F K

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20

70

AERIAL SURVEY Prince William Sound

April 1970

Between April 15 and 18 1970 Carl Divinyi and Julius Reynolds fletr an aerial survey of most of Prince William Sound using a Cessna 185 The survey tras primarily to locate seals however sea otter were counted The following is a summary of the survey

I departed Anchorage on April 14 via Alaska Airlines and arrived in Cordova at 600 AM tJeather conditions prevented flying so Julius and I drove the local road network looking for anything in the way of wildlife

April 15 vleather was a little better than yesterday so we decided to try surveying along the east side of the Sound up to Valdez Arm We started the survey at Bomb Point and flew the coast and offshore islands up to Galena Bay

Observations Seal - Scattered small groups with the majority of the animals being in Port Fidalgo There were no noticeable concentrations (50 on up) of seals

Sea Otter - A total of 105 otter were counted with the majority (60) being located in St Matthews Bay The remainder of the animals were concentrated between Red Head and Knmvles Head outside of Port Gravina

Sea Lion- Small scattered groups in Port Fidalgo (60-80 total)

April 16 Weather inclement - no flying

April 17 We flew Hawkins Hinchinbrook Montague and Green Islands Very few seal- many otter and two large concentrations of sea lions (See maps for numbers and locations of seals sea otter and sea lion) We also counted 5 otter around Kayak Island

April 18 Flew those islands from Montague Strait to Icy Bay and from Port Bainbridge to Knight Island Passage (See maps for numbers and locations of seals sea otter and sea lions)

Julius Reynolds continued the survey along the mainland from Knight Island Passage to Esther Island and around Knight Ingot and Eleanor Islands

The area from Esther Passage to Valdez Arm and Culross Perry Lone Naked Peak and Storey Islands were not surveyed

SEA OTTER COUNTED Prince William Sound

April 1970

Number of Area Sea Otter Date

PORT BAINBRIDGE - ICY BAY Bainbridge Island 30 418 Evans Island 14 418 Elrington Island 4 418 Latouche Island 46 418 Whale Bay - Icy Bay 39 418

CHENEGA ISLk~D - MAIN BAY Chenega Island and Dangerous Passage 33 52 Crafton Island 2 52

PORT NELLIE JUAN - ESTHER ISLAND Blackstone Bay 1 52 Culross Island NS

ESTHER PASSAGE - VALDEZ ARM NS

GALENA BAY FISH BAY 103 415

FISH BAY - GRAVINA POINT 1 415

GRAVINA POINT - CORDOVA 1 415

HAWKINS ISLfu~D 1 4l7

HINCHINBROOK ISLfu~ 99 417

EGG ISLAND 2 417

MONTAGUE ISLAND 259 417

GREEN AND LITTLE GREEN ISLANDS 102 4J7

KNIGHT ISLAND 136 52

INGOT ISLAND 6 52

ELEANOR ISLAND 3 52

SMITH ISLAND 4 52

SEAL ISLAND 0 52

APPLEGATE ROCK 1 52

PERRY ISLAND NS

(continued) SEA OTTER COUNTED Prince William Sound

April 1970

Number of Area Sea Otter Date-

LONE ISLAND NS

NAKED ISLAND NS

PEAK ISLAND NS

STOREY ISLAND NS

KAYAK - WINGHAM ISLAND 5 417

TOTAL 892

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PRE-TRANSPLANT SURVEY OF SEA OTTERS Green 1 andMontague 1 -July 171970

On July 17 1970 an aerial survey of sea otters was made in the area

around Green Island and the adjacent coast of Montague Island The purpose

of the survey was to locate concentrations of animals prior to a capturing

operation which began the next day A Dornier twin-engine arrcraft was used

with Schneider Reynolds and Somerville acting as observers The sky was

overcast almost no wind and the sea calm However heavy rain showers intershy

fered with forward visibility and the counting conditions were only fair on

the average Conditions vmiddotere especially poor in Port Chalmers and Zaikof Bay

The count began at 210pm and ended at 345 pm

The numbers and locations of otters are shown on the map In addition

16 sea otters were seen on a superficial look at Constantine Harbor on Hinchshy

in brook Island and a pod of 35 were seen three miles east of Johnstone Pt

Most of the area included in the survey was traveled repeatedly in a

skiff over the next four days Durlng this time the weather became calm and

clear for the first time in several days and the bulk of the sea otters shifted

from the coast of Montague out into the shallow areas between there and Green 1

and to Green and Channel Islands It was obvious that many otters had been missed

Fn the aerial sur~ey and that the population in the area is quite dense A

total of 48 sea otters were captured in three days of netting

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SEA OTTER SURVEY Salisbury Sound to Cape Spencer

August 23-25~ 1970

On August 23 1970 an aerial search for sea otters was made from Salisbury Sound to Torch Bay in the area north of Sitka A Helie Courier aircraft was used with Karl Schneider Alan Courtright and

middotWalt Cunningham serving as observers

The water on the outside coast was too rough for good visibility however it was calmer in the lee of rocks and islands and visibility was fair to good

On the initial survey only two otters were seen These were both in Surge Bay on Yakobi Island While returning after completing the count we located approximately 25 in the same area we saw the first two About 15 of these including at least one pup were in a single pod This illustrates how relatively large numbers of otters can be missed from the air The fact that none were seen in other areas does not mean that established populations do not exist elsewhere

On August 24 and 25 a search of the release area north of Khaz Bay was made by the skiff While flying into Kla) Bay for this seach two otters were seen near the old mine of Chichagof Two miners who have been working there reported that two otters have been there off and on since May

Rough weather and outboard problems restricted tl the search area~

and the effectiveness of the search in that area The following sightings were made

82470 I Adult West of Granite Islands 1 Adult South of Granite Islands

11 0A~dgtzl~4JVflup) middot1 In rocks SE of ranite Islands 82570 I Adult -n Southwest of Gran~te Islands

These sightings represent from four to seven animals

Reports over the last year indicate that sea otters regularly move in and out of Kla~ Bay and occasionally as far in as Sisters Lake bull

The fact that the count in the Khaz Bay area Has less than that made in 1969 does not mean much The animals appeared to be scattered during the present count the search did not include all of the nearby sea otter habitat and survey conditions were less than ideal

The population in Surge Bay appears to be separate and is probably well established Two otters were reported in the same location in January 1966 after only 23 otters had been released near Khaz Bay In 1969 two were seen in the same spot from the ait

While no estimate of the population of sea otters in the area can be made from the available information is evident that sea etters can be found along the entire west coast of Chichagof Island and that concentrations occur near Black Island and the Granite Islands north of Khaz Bay and in Surge Bay on Yakobi Island

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55

HARVEST AND TRANSPLANTS - 1970

Harvest

In May of 1970 a sea otter harvest was conducted on Tanaga Island the middotDelarof Islands and Amchitka Island The numbers to come from each island

and the distribution of the kill around each island were planned in advance Conservative estimates of the population tvere made and the total harvest numbers were based on these estimates The harvest level for Tanaga and the Delarof Islands was set at 15 percent assuming that the next harvest in these areas would be three years later so the rate of harvest would be 5 percent per year Amchitkas harvest level was set at 10 percent assuming an annual harvest and a harvest rate of 10 percent per year

The harvest plan was based on information collected on a June 1968 skiff survey for Tanaga on the 1965 USFWS aerial survey and the 1969 ADFampG aerial

survey for the Delarofs and on 1968 helicopter survey by the USFWS for Amchitka There is good evidence that all the population estimates tvere low especially for the Delarofs

Hunting was done from avo skiffs with two men each Both men shot when possible A 117-foot vessel was used for skinning and living quarters Table 1 shows the schedule of the hunt with the daily harvest numbers

Table 2 presents the projected and actual harvest numbers by area Table 3 presents the numbers killed pelts saved specimen and pelt numbers used and the sex ratio of the kill for each area Figures 1 2 and 3 show the actual numbers and locations in more detail tveather was the main factor causing deviations from the original harvest plan Possible male areas were identified on the 1968 skiff survey on Tanaga An attempt was made to concentrate more pressure on these areas to achieve a more balanced sex ratio This effort is reflected in the sex ratio of the kill A high degree of sexual segregation occurs at this time of year and without specific pressure on male areas we could expect 85 percent females to be taken as occurred on Amchitka With the effort the female percentage was reduced to 65 percent on Tanaga In the Delarofs a male area which had not been identified previously was hit and the effect was much the same

Transplants

Two transplant operations took place in 1970 The first was done in the same manner as the 1968 and 1969 transplants A total of 86 otters tvere captured at Amchitka One aircraft load was shipped out Twenty-nine were released in Oregon and 30 in vashington The animals were held in floating pens at the release sites for several days to recover from the trip Survival appeared to be excellent

The second operation took place in Prince William Sound The Canadian research vessel G B Reed came to the Green Island-Port Chalmers area with tanks built on deck Forty-six sea otters were captured Approximately 44 were alive when the vessel left Prince William Sound however only 14 survived to be released off Vancouver Island

Table 1

April 20

middotMay 1

May 2

May 3

May 4

May 5

May 6

May 7

May 8

May 9

May 10

May 11

May 12

May 13

Schedule of the 1970 harvest

1030 pm depart Homer

Midnight arrive Adak MV Active

Refuel and prepare boat

Arrive off Tanaga at noon

Complete skinning pack hides etc in pm

Delarofs

Spent am getting water at Amchitka

Clean up boat take down shelter pack gear

900 am arrive Amchitka fly to Anchorage

Killed 53

138

88

131

143

53

144

43

79

83

otter

(am)

(pm)

(by mid afternoon)

Table 2 Projected and actual harvest numbers May 1970

Projected Actual Tana~a Island Harvest Harvest

Bumpy Point - Hot Springs Bay 200 191

Hot Springs Bay - Twin Bays 50 50

Twin Bays - Cape Sasmik 50 ) ) 199

Cape Sasmik - Tower 120 )

Tower Tanaga Bay 180 166

TOTAL 600 606

Delarof Islands

Gareloi 20-25 0

Kavalga Ogliuga and Skagul 75-100 125

Ulak and Amatignak 35-55 19

TOTAL 150 144

Amchitka Island saved for

USFWS Counting Units 1 amp 2 100 transplant

3 30 82

4 50 36

5 120 87

TOTAL 300 205

Table 3 Details of sea otters harvested in May 1970

Tanaga Island

Total kill 606

Number of pelts saved 598

Number of small pup pelts discarded 8

Specimen numbers SO 70-4 to SO 70-609 Pelt numbers 3198-3769 and 3771-3796

(Seal 3770 void - damaged)

Approximate sex ratio - 220 males 386 females or approximately 64 percent females

Delarof Islands

Total kill 144

Number of pelts saved 138

Number of pups discarded 6

Specimen numbers SO 70-610 to SO 70-753 Pelt numbers 3797 to 3934

Sex ratio- 44 males 100 females or approximately 69 percent females

Amchitka Island

Total kill 205

Number of pelts saved 194

Number of pups discarded 11

Specimen numbers SO 70-754 to SO 70-958 Pelt numbers 3935 to 4128

Sex ratio- 27 males 178 females or approximately 867 percent females

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A combination of bad weather a lack of time to let the otters recover from their capture and problems with the holding facilities probably caused the high rate of mortality

Summary of Harvests and Transplants

Table 4 summarizes the sea otters removed from Alaskan populations in the middot last 20 years An attempt has been made to remove 300 otters from Amchitka each year since 196 7 Originally this number was set as 10 percent of the population which was conservatively estimated to toal 3000 Recent USFtfS helicopter counts of up to 3927 show that this estimate definitely tvas low In the past the removal of animals was from the southeas tern half of the island The 1970 harvest was purposely concentrated toward the northwestern end in order to spread out the pressure and to learn something about the population in that area

The numbers of animals released in all transplant attempts by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game are presented in Table 5

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Tab

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N

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N

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30

SEXUAL SEGREGATION IN SEA OTTER POPULATIONS

by Karl Schneider March 1971

Workers have recognized for some time that sea otters tend to segregate by sex Lensink (1962) described this segregation around the southeastern end of Amchitka Island and identified three male areasn and three female areas He speculated that females used areas of more favorable habitat and that younger males were excluded by territorial males scattered throughout the female areas The implication is that male areas contain younger or at least nonterritorial males

Marakov (1965) mentions sexual segregation around Medny Island in the USSRs Commander Islands

Kenyon (1969) gives a more complete description of the sexual segregation around the southeastern end of Amchitka and supports it with quantitative data from harvested animals

Both Lens ink (1962) and Kenyon (1969) w_ere primarily describing hauling grounds used by different sexes While Kenyons harvested animals included otters near shore neither study provided much information on the use of off-shore areas

A knmmiddotlledge of the degree of sexual segregation and the location of male and female areas is important to the management of sea otter populations Harvests must be regulated to avoid putting too much pressure on one segment of the population Then capturing animals for transplants it may be necessary to set nets in specific areas to obtain the desired sex ratio case of a localized kill of otters such as when oil spills occur it is important to know the distribution of sexes to evaluate the importance of kill to the population

In

the

By the same token much can be learned about the distribution of sexes through harvest and capture operations The area from which each sea otter was taken during the harvests of 1967 1968 and 1970 was recorded Because a single hunting party might kill 30 otters over up to 10 miles of coast in a few hours it wasnt possible to record the precise locations Therefore the coast was broken up into areas and the numbers killed in each area ltJere totaled

Figs 1 - 5 show the locations of the areas letters correspond to those in Tables 1 - 4 which present the sex and age composition of animals harvested in each area The ages of the 1968 animals 1vere based on cementum deposition and reproductive condition The other samples were broken down using body size In general all males under 25 lb females under 20 lb and both sexes shorter than 100 em were considered dependent pups Females under 35 lb and 120 em and males under lb and 130 em were considered subadults These criteria probably underestimate the age of some animals Cementum deposition information for the 1967 and 1970 samples is not available yet

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Sexual Segregation - 2 - March 7 1971

Information from the 1967 Amchitka harvest and the 1968 1969 and 1970 Amchitka transplant capturing operations is not comparable and is not presented here However knmv-ledge gained from these operations has contributed to our understanding of sea otter distribution and this knowledge will be used in the following discussion In general this data confirms Kenyons (1969) observations for the Amchitka area although these summer and fall collections indicate a higher percentage of males in female areas than Kenyon reported from his winter and spring samples

Kenyon (1969) states that female areas are more numerous and less discrete than male areas He identified three male hauling grounds and 13 female hauling grounds on the southeastern end of Amchitka Island Recent studies involving netting and collecting animals off~hore indicate that male areas remain discrete off shore Usually these areas are off major points of land and pods of males often form in kelp beds directly off shore from the hauling ground

Female areas on the other hand are not discrete at all Any area that is not a male area can be considered a female area Some areas may be more attractive to females with pups or certain areas may be more favorable for hauling out but more females than males vill be found almost everyvthere except in the discrete male areas Therefore the main problem is to locate the male areas in et population He have no evidence of any shift in the location of male areas over a period of time so once an area is identified the information should remain useful for management purposes for many years Lensink (1962) and Kenyon (1969) correctly identified all of the male areas on southeastern Amchitka Extensive harves and netting have not turned up any new areas

Identification of Hale Areas by Pup Counts

In June 1968 a survey of most of Tanaga and Kanaga Islands was made by skiff to gather information to be used in planning harvests from those islands During this survey females with pups were counted separately from single animals No special effort was put into identifying pups fuen a female obviously had a pup it vas recorded Large pups often were separate from the females and some otters were seen only briefly in vaves kelp or at a distance Therefore many pups probably were missed and the counts should be considered minimal 6 and 7 present the counta by area A similar count was attempted by helicopter around Adak but for various reasons the results are not considered useable

From this skiff survey a number of areas were judged to be possible male areas Usually these were points or more exposed aTeas vhere relatively large numbers of single animals but no pups were seen

Shoal Point Kanaga Pass north of Eddy Rock and possibly Naga Point and Cape Tusik were suspected male areas on Kanaga Island Cape Sudak and Cape Amagalik were considered male areas on Tanaga Island and the area around Hazard Point and Lltnnoy Rock in Kanaga Pass was believed to blend with the Eddy Rock area

The best way to confirm the presence of male areas is to collect animals from these areas Harvests were conducted on Kanaga in October 1968 and on

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Tanaga in May 1970 The sex ratios of the animals harvested are compared with the skiff survey pup counts in Table 5 The harvest areas are fairly broad while male areas are usually only a few hundred yards wide As a result a harvest area may include both a male area and portions of female areas to either side

The hunting pressure is seldom even over an area Often a good portion of the kill in an area would come from one group of rocks Therefore the information in Table 5 must be tempered with some knmrledge of how the hunting pressure was distributed

Kanaga Island

Shoal Point - While a few more females than males were taken in this area the percentage of males is considerably higher than one would expect in a female area Because weather vas less than ideal much of the hunting pressure ~vas directed to areas other than Shoal Point Most of the females probably came from these areas Therefore Shoal Point is considered to be a male area$

Naga Poin~- There is no strong evidence that this is a male area While no pups were recorded on the survey the total number of otters counted was not high Fairly extesive hunting of the area under good conditions yielded a sex ratio that would be typical of a female area Therefore it is assumed that there is no male area near Naga Point

Cape Tusik - Cape Tusik was suspected as a male area more because of the nature of the area than arry counts The count included areas to both sides of the Cape which could contain females Very little of the hunting was done at the Cape Therefore judgement on this area will have to wait until more information is available Extensive hunting indicates that no male areas exist east of Cape Tusik to Cape Chlanak

Kanaga Pass - From the count it appears that a male area exists in the Pass however the harvest area extended to Hive Rock and included a large portion of female area (Fig 6) The area is too broad as is shown in Table 5 gtvhere the number of pups counted for the whole area is relatively high The results of the harvest confirm the conclusions drawn from the count While the sex ratio for the entire area is roughly equal most of the males came from Kanaga Pass Some females without pups came from that general area but the majority of the females came from the area northeast of the Pass

The Tanaga harvest showed the male areas more distinctly for several reasons The total harvest was higher the entire count area was covered in the harvest and the hunting pressure was recorded more precisely Also as will be shown later sexual segregation is more pronounced in the spring

Cape Sudak - Most of the hunting pressure was concentrated on the tip of the Cape Some pressure was exerted on the area back to~mrd Pendant Point which is a female area as shmvn by the count The high percentage of males harvested from this area clearly confirms that the tip of the Cape is a male area

Tab

le

5

Com

pari

son

of

Pup

s C

ou

nte

d w

ith

S

ex R

atio

o

f H

arv

est

Jun

e

1968

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rea

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rvey

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cto

ber

19

68 H

arv

est

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ps

10

0

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ps

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0

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ult

s T

ota

l KA

NAGA

IS

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er

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nd

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d-S

ho

al P

oin

t 0

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45

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71

5

29

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a P

oin

t-K

anag

a B

ay

66

1

64

middot

18

5

81

5

20

5

79

5

Cap

e C

hla

nak

-Cap

e T

usi

k

51

1

93

2

40

7

60

3

53

6

47

Edd

y R

ock

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e R

ock

61

3

4

35

7

64

3

47

1

52

9

Jun

e

1968

A

rea

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rvey

H

ay

1970

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ves

t

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ps

10

0

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ps

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0

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ult

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l TA

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GA

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ep

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er ~ter

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F

M

F

Cap

e S

ud

ak-P

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ant

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int

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6

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nes

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in B

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ay

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uth

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igh

t 6

9

82

1

32

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51

8

49

Sexual Segregation - 4 - March~ 1971

Annoy RockHazard Point - The harvest did not confirm this area as a male area However the weather Vas marginal and this area was too rough to hunt Therefore the otters taken in the harvest ~vere from either side of the Point and almost none were taken from the portion that is suspected to be a male area Actually Kanaga Pass is narrow and shallmv Otters feed in all parts of the Pass when weather and tide rips permit This could be considered a single male a~ea that serves both islands

Cape Amagalik - Two harvest areas overlap this area Lash Bay-Inferno Reef and Harem Rock-Kulak Point The kill from both areas strongly reflects the presence of a male area Nost of the hunting pressure was concentrated on Inferno and this is probably the main male area

As shown above vle 1vere able to identify four major male areas from a relatively superficial pup count No male areas were located during the harvests that had not already been identified from the count ~mile more detailed observations including field identification of males would be desirable the pup counting technique appears to be an acceptable method

Identification of

Because male areas are usually on v1ell exposed points it is possible to pick likely areas from a chart and then confirm these areas by collecting animals This method avoids the expense time and danger involved in surface surveys of remote areas Actual collection of animals also provides the most concrete proof of the nature of an area

The follmving is a description of the sex composition of areas for which no skiff count data was available

Delarof Islands - No attempt 1vas made to predict the location of male areas in the Delarof Islands This is a difficult area to work in Ogliuga Skagul and a portion of Ulak Island v1ere hunted on Nay 9 1970 The hunting effort in the afternoon was concentrated around Tag Island and the rocks south of Skagul Island While the entire kill is lumped together most of the males bullJere taken in the afternoon The percentage of males taken for the day is higher than would be expected if only female areas were hunted Most likely there is a male area near Tag Island

Adak Island - Portions of Adak gtvere hunted in 196 7 and 1968 No attempt was made to identify male areas before or during either harvest The harvest figures indicate that no male areas have been hunted The Bay of Islands is a classical female area The results of extensive hunting in the Bay and around Eddy Island indicate that there is definitely no male area there The percentage of males taken between Mid Point and Blind Point vms higher than might normally be expected in a female area however the scarcity of subadult males indicates that no male area exists within this area A large congregation of otters is often seen near Finger Shoals This could be a male area but Ye have no information from there If this is a male area a few stray males from there could account for the slightly higher number of males in the kill

Sexual Segregation - 5 - March 1971

Amchitka Island - While much work has been done on Amchitka Island almost all of the effort has been concentrated on the southeastern end from Crmvn Reefer Point to a fegtv miles northvest of Rifle Range Point During the 1970 harvest an attempt was made to distribute the harvest over previously unstudied areas Two areas were suspected to be male areas These were the rocks off Chitka Point and either Bird Rock or Aleut Point Heather prevented us from hunting the area around Bird Rock and Aleut Point

The sex ratio of the kill does not indicate a male area near Chitka Point however heavy waves prevented hunting off the point Therefore a male area could be there but because no animals were killed there the kill figures would not reflect it

Composition of Female Areas

Kenyon (1969) found that 93 percent of the adult otters and 63 percent of the juvenile otters in female areas were females The May samples from the 1970 harvest agree with this The mean percent of adult females in female areas was 93 percent The mean percent of subadults not including pups was 73 percent Hmmiddot1ever in the September and October samples only 77 percent of the a-dults in female areas were female About 86 percent of the subadults were female

The seasonal difference in the sex ratio of adults is consistant for all samples These seasonal changes are most likely due to adult males moving into female areas to look for estrous females During late winter and spring breeding activity is at the lmmiddot1est point of the year and fev1er females are entering estrus In fall the number of estrous animals is at its peak and presumably more sexually mature males move into the female areas to breed It is interesting to note that there were about three times as many males in female areas in fall as in winter and spring Information from female reproductive tracts indicates that approximately three times as many females would be in estrus at any given time in fall It appears that the number of adult males in a female area is directly proportional to the number of estrous females From the data collected there appear to be 15 to 20 males for each female -vlith enlarged follicles at any given time hmmiddotrever hunter bias might influence this figure

Ages have been estimated for otters harvested in 1968 on the basis of cementum layering These age data indicate that virtually all of the males in discrete female areas are either pups (or very young subadults) or are 7 years old or older No males between the ages of 2 and 6 Here found This strongly implies some form of territorial behavior among actively breeding males Fighting among males has not been observed ofteci hmvever a mature male at the Tacoma Zoo became intolerant of a subadult male in the presence of females They frequently 11 fought 1 although neither seemed to attempt to injure the other Finally the young male had to be removed Under imiddotJild conditions the young male might have moved out of the area without repeated physical encounters

Kenyon (1969) described the breeding behavior of sea otters His description indicated that males patrolled an area and two to four males may pass a given point during a 3 or 4 hour period Vandevere (1970) observed some males

Sexual Segregation - 6 - March~ 1971

apparently defending a territory in California but from his observations t

it appeared that successful breeding males were more mobile Perhaps established breeding males are more tolerant of each other More likely they maintain a separation from each other which limits the number in an area In this way a number of males might be patrolling the sa~e area rather than each establishing a geographical territory This might be advantageous where female concentrations shift from place to place as weather conditions change

Why does the number of adult males in female areas decline when the number of estrous females declines The limiting factor may be competition for estrous females rather than territory size With fewer receptive females competition lvould be greater causing some animals to leave the area Another possible reason may be that males may have a seasonal fluctuation in fertility Lensink (1962) found that all adult males produced sperm at all seasons Limited studies since then support this however it is possible that most of these males were collected in or near female areas Also while a male may produce sperm at all seasons there may be less production at certain times of year

A contributing factor may be that males are capable of feeding in more exposed areas than pregnant females or females with pups With a reduced attraction to the female areas either middotbecause of fewer receptive females or a reduced fertility in the male the male may find it easier to obtain food away from the crowded female areas where competition for food is greatest

The fact that the number of males does appear proportional to the number of estrous females indicates that competition for these females may play an important part in regulating the number of males in female areas

The numbers of subadults in the samples is relatively low so fluctuations in sex ratio may be due to sampling errors However the greater number of subadult males found in female areas in winter and spring may be a true increase made possible by the reduction in the number of breeding males A subadul t male would have fewer encounters -vli th breeding males

There are areas within what would normally be considered female areas where subadult males tend to congregate particularly during the winter These are usually areas v7ith a ma-r-ginal food supply that are not heavily used by females Because there are fewmature females in these areas there are also very few breeding males The subadult males using these areas seem to be transients that find less pressure from breeding males there These areas are used less in summer probably because calrrer weather allows feeding farther off-shore reducing competition for food in the male areas

Examples of areas within female areas that are used by subadult males are Constantine Harbor on Amchitka and possibly Finger Bay on Adak

within Female

In the course of the harvest on Tanags in tIay of 1970 a large number of pregnant females tvith large fetuses were collected at one time Most of

Sexual Segregation - 7 - March 1971

these came from Tidgituk Island This raised the possibility that females may segregate according to reproductive condition Table 6 presents the numbers of females in each stage of the reproductive cycle by area on Tanaga Island More pregnant animals were collected in the South Bay area and within that area most of the females with fetuses weighing over 100 g were collected near Tidgiktuk Island This concentration was evident in the pup counts made in June 1968 when a very high percentage of pups was observed in the same area (Table 5) Host females ~vith large fetuses in May would pup by June

The data do not reveal any other areas of this type on Tanaga Harakov (1965) mentioned a bay on Hedny Island that tvas preferred by pregnant females and females with pups Obviously all females do not go to a specific area to pup but a female vi th a large fetus or small pup probably has more difficulty in exposed areas and seeks more protected areas This tendency can be seen by the casual observer Single animals are more likely to venture off-shore and remain in rougher waters In some areas such as Crown Reefer Point on Amchitka there is a male area near the tip of a point Males congregate directly off-shore However to either side of these male congregations there are females that are not accompanied by a pup and probab do not have large fetuses In this way we may find segregation in a single kelp bed off a point If we set a net near the outer margin of the kelp we will catch almost all males Ho~Vever if we set a net to the side of the bed and closer tmiddoto shore we will catch mostly single females Nets set in a nearby bay will catch more females with pups

Table 6 shoHs that an unusually high number of resorptions were found between Cape Sudak and THin Bays This is probably a reflection of poor physical condition of the otters in that area due to food Other data indicate a continuous decline in body condition from the east side of Adak to Kanaga Pass There is some evidence that condition improves west of Kanaga Pass The animals in the Kanaga Pass vicinity are probably in poorer condition than any of the other populations sampled Therefore the high incidence of resorptions in this area should not be considered an example of segregation within the population

ition of Male Areas

As mentioned earlier the harvest areas ivere broader than any male area Harvest areas containing male areas also contained portions of female areas Our identification of the location at rhich each animal vas collected is not precise enough to determine the exact composition of these rnale areas Kenyons (1969) data from winter and early spring harvests are more precise His data indicate that at that time of year 98 percent of the adults and 80 percent of the ju~veniles using male areas were males Of 128 males in male areas 100 gtvere adults and 28 were juveniles

Experience from netting in the summer and harvesting in fall indicates that the percentage of males remains very high all year While adult females may be very close to male areas it appears unlikely that a significant number regularly use these areas at any time year nile adult males enter female areas regularly and in predictable numbers for a specific purpose females probably ~nter male areas only occasionally in stray movements

le 6

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Sexual Segregation - 8 - March 1971

Juveniles of both sexes probably move more randomly than adults lihile definite sexual segregation exists among younger animals it is not as pronounced as in adults Because subadult males are tolerated less by breeding males they are more strictly confined to small areas than females of the same age This has been demonstrated in fall harvests where extensive hunting over a large area will turn up only an occasional subadult male Then ~v-hen the hunters move to a definite male area large numbers of subadult males are taken in a very short time and in a very small area

While Kenyons data indicate that 22 percent the males in male areas are subadults our experience indicates that this percentage may be quite a bit higher at least in fall Such a seasonal change in the age composition of males sounds reasonable We have demonstrated that adult males move into female areas in greater numbers in the fall perhaps tripling their numbers in these areas At the same time fewer subadult males are found in the female areas Presumably the adult males are coming from male areas and the subadults driven from the female areas move to the same male areas The result vmuld be a higher percentage of young males in male areas in fall than in spring

The total number of males of all ages in female areas doubles in fall About 13 percent of all animals taken in these areas in spring were ~ales while 25 percent ta-ken in fall were males This suggests that the total number of males in male areas declines in fall There are more adults leaving than subadults entering the area

Sex Ratio

With the sexes segregating and with the degree segregation changing seasonally and betmiddotween age groups it is extremely difficult to get a completely random harvest As a result we have not been able to determine the sex ratio of any population as a vhole Hith female areas being larger and in more protected areas where hunting is easier there is a strong tendency to take more females than males particularly in sp-ring ~Je have shown that through a concentrated effort the percentage of males harvested can be increased however even then the lowest percentage of females taken in recent harvests ivas 62 percent on Kanaga Island in October 1968 Similar results occur in capturing operations for transplants

On all islands studied there are fewer male areas thltm female areas The male areas are very small usually only a few hundred yards wide The male feeding areas off male areas are also very narrow although they may extend farther off-shore In short that portion of the total available sea otter habitat included in male areas is very smalL

All evidence indicates that there are more females in the population than males While the percentage of is probably greater than that indicated by most harvest statistics it appears that at least 60 perc2nt of the sea otters in the populations studied are females

There are probably several factors contributing to this uneven sex ratio First our reproductive data indicate that 56 percent of the pups are females at birth Secondly Kenyon (1969) has found a higher mortality rate among

Sexual Segregation - 9 - March 1971

newly weaned males There is also some evidence that males might not live as long as females These factors could easily account for the preponderance of females in the population

It is possible that the higher rate of mortality in juvenile males is in part due to the breeding behavior of adult males and the resulting sexual segregation Juvenile males tend to be restricted to male areas or areas of marginal feeding habitat where there is little competition from breeding males During the ltvinter the feeding activity of young males would tend to be confined to small areas close to shore in male areas Therefore they may be subjected to greater competition for food than females of the same age which have a broader area in ltvhich to feed Kenyon (1969) found that subadult males vere less hardy in captivity than subadult females Therefore we can not adequately evaluate the influence of segregation on mortality

The sex ratio probably varies from one population to the next Very little juvenile mortality occurs in expanding populations eliminating that source of sexual selection However there is some evidence that males are more numerous among the first animals to repopulate a new area While a high percentage of males might be found on a newly repopulated island the loss of these males tvould alter the sex ratio of the parent population even though juvenile mortality might not yet be a significant factor

If the nThuber of breeding males is regulated by the number of estrous females an even sex ratio might produce a surplus of sexually mature males This surplus would not be beneficial to the population and could be detrimental This would permit the situation of an unbalanced sex ratio to evolve

The segregation of sexes and ages and the composition of sea otter populations is complex Unless we can understand the situation we cannot fully evaluate the effects of mortality or habitat changes

REPRODUCTION IN THE FElIALE SEA OTTER

by Karl Schneider Narch 1971

A total of 1358 sea otter reproductive tracts collected between 1967 and 1970 have been examined Ovaries were sliced with a razor blade and examined macroscopically Uteri Here macroscopically examined both internally and Each tract vas classified as to its stage in the reproductive cycle

The follovJing criteria were used in classification

Nulliparou~- Horns of uterus small thin and smooth no corpora lutea or corpora albicantia although there is occasionally evidence of a past ovulation but no evidence of implantation

Nultiparous - Uterus thicker corpora lutea andor corpora albicantia present Includes primiparous animals (these are not readily separated from multiparous animals)

Anestrus - Largest follicle under 3 5 rnm no corpus luteum

Proestrus - Follicles over 35 mm

Es - Follicles approximately 10 rrm

Implan~elt_ Pregnant_ - Visible s~velling in uterine horn usually with fetal membranes embryo or fetus visible Corpus luteum usually over 10 mm

- Uterine horn sti11 noticeab enlarged fresh roughshy~=-=--=~

al scar newly formed corpus albicans with some luteal tissue remaining

The appearance of the uterus ~ras used to confirm the classification The thickness arrd of the horns thickness and consistency of the uterine walls coloration of the of the horns and condition of the rugae change with each stage In most cases it is possible to guess the condition of the tract by a superficial examination of its exterior This appearance was used only to confirm what was indicated by structures in the ovary hmvever

Table 1 smnmarizes the reproductive condition of the sexually mature females in the that are enough for comparison throughout the year In the following discussion reported by Sinha et (1966) and Kenyon (1969) are included 7here possible The January and Harch are from these studies

Tab

le

l Re

p iv

e co

nd

itio

n o

f se

xual

ly m

atur

e se

a o

tters

_

IF

etus

Wei

--------------+

__

_

ln~a

ctive

ov

arie

s 1

Act

ive ovari~

ht C

Jas

s

jmiddot

I P

ost

-I P

roes

tru

s U

nim

pl

lmpl

D

ates

L

ocat

ioQ

1A

nest

rus

Part

um

Res

orpt

ion

l +

E

stru

s P

reo

Pr

eQ

2 3

lJ

S

~

4-8

1970

T

anag

a 1

51

41

0

10

104

17

8 10

33

30

4 (1

68)

(1

35)

(3

3)

(3 3

) (2

89)

(34~

(5

6)

(27

) (3

3)

(11

4)(

10

9)

May

9

19

70

Del

arof

Is

18

14

1

4 13

27

10

0

3 6

8 77

4

) (1

82)

( l

3)

(5

(16

9)

( o

) (1

30

) (3

9)

(78

)(1

04

)

10-1

219

70 A

mch

ltka

I

34

18

3 8

27

48

5 6

2 23

12

13

8

( 0

(24

6)

(13

0)

(22

) (5

8)

(19

6)

(34

8)

(36

) (4

3)

(14

) (1

67)

(8

7)

~lune

23

middotmiddot ~middot~~middot~Amchitka

1

17

2 1

2 4

18

1 2

3 5

7 44

A

ugus

t 13

196

8 (3

86)

(4

5)

(23

) (4

5)

(90

) (4

09)

(2

3)

(45

) (6

8)

(11

3)(

15

8)

July

6-

~dgtAmchitka

1

17

1 0

4 14

10

0

1 1

2 6

46

Aug

ust

819

69

(36

9)

( 2

0 2

) (8

6)

(30

4)

(21

9)

(22

) (2

2)

(43

)(1

32

)

Sep

t 1

0-17

A

dak

72

5 0

16

31

40

6 6

8 9

11

164

1967

(4

39

) ( 3

1)

(98

) ( 1

89)

(

4)

(3 7

) (3

7)

(49

) (5

5)

(6

7)

Sep

t

25

-A

mch

itk

a l

55

6

0 18

19

17

4

0 0

0 3

115

OcL

6

19

67

(4

7~8)

(5

2)

(15

7)

(16

5)

(14

8)

(35

) (8

7)

(26

)

Oct

o 1

2-1

5

Kan

aga

I

58

6 1

13

31

31

4 4

7 11

5

140

1968

(4

13)

(4

3)

(o 7

) (9

3)

(22

2)

(22

2)

(29

) (2

9)

(50

) (7

8)

(36

)

Oct

18

-20

A

dak

l

46

6 0

1 24

13

2

3 0

4 4

100

1968

(L

16 0

) ( 6

0)

(11

0)

(24

0)

(13

0)

(20

) (3

0)

0)

(40

)

Num

bers

in

re

nth

esis

are

pe

rcen

t se

xu

ally

mat

ure

fem

ales

F

etus

wei

ght

clas

s 1

=

0-l

g

2 =

1-l

Og

3

= 1

0-lO

Og

4

= 1

00-lO

OO

g

5 =

Tra

nsp

lan

t m

ort

alit

ies

(all

o

ther

sam

ples

fr

om

har

ves

ts)

Reproduction - 2 - March 1971

There may be some problems associated with comparing samples taken from different islands and in different years but allowing for sampling errors the data for the most part fit a definite pattern indicating that the annual cycle remains fairly constant The one outstanding exception is the number of pregnant animals in the t1m summer samples In 1968 there were many more implanted pregnancies than unimplanted pregnancies and in 1969 the situation was reversed hmvever the total number pregnant was almost identical These are the t s being relatively small collected over a longer of time and consisting of transplant mortalities An average of the two samples fits the pattern established by the other samples Relatively large samples are necessary because all stages of reproduction are found at all times of the year The period from November to early January is not represented hmmiddotJever things are changing so rapidly during that period that any information from that time may be confusing unless a large sample was collected in a very short time

The information in Table 1 is shmvn graphically vJith Kenyons (1969) winter information in Figs 1 and 2 It has often been demonstrated that sea otters breed and pup at all times of the year A number of observers have noticed that roore pups are born in the spring aJd sumrrter than at other times and there have been conflicting reports about breeding times Kenyon (1969) was the first to demonstrate seasonal changes in pregnancy rates but only his January and Yarch sallples gtvere large enough to be at all reliable

Figs 1 and 2 demonstrate that there are cons le secsonal in the numbers of animals in each s reproductive cycle There is a definite period of increased bree activity and a definite period of increased pupping Hmmiddotlever ~ some anii1als are in each s at all times of the year and these periods of increased breeding and pupping do not have distinct boundaries

Each curve is influenced by two factors so it is difficult to isolate the magnitude of any single factoc For example the of implanted pregnancies vill increase ss blastocysts and decrease as births occur The birth rate may be increas but if the number of implantations increases at a greater rate the implanted curve will still rise The animal remains in the anestrus implanted pregnant~ and unimplanted pregnant categories for a relatively long time and there may be considerable carry-over from one sample to the next Because the animal a relatively short time in the proestrus-estrus and post partum categories there is little if any carry-middotover from one sample to the next and the changes betgtmiddotTeen are more likely to be absolute changes The mean fetus weight class prob does not mean much because it does not take the actual number of animals in each class into account ~hen there are many implanted pregnancies ard the mean lmiddotTeigh t class is lovJ there actually may be more Class 5 fetuses in the population than vhen there are fetv irr1planted pregnancies and the mean middotleight class is high Therefore the actual percentages of all sexually mature females middotlith Class 1 ald Class 5 fetuses have been plotted in 3 These curves represent the actual number of fetuses in the population Because tha sample sizes become quite small vhen the fetuses are divi(12d into classes and the surruner sample is believ2d

2

~ _s ~

~

c U)

~-1

r

~J

lt

c])

Reproduction - 3 - Narch 1971

to be biased tvo curves for each class are shmvn The solid curves approximate the data while the broken curves represent subjective adjustments to correct for sampling errors

cussion of the Annual

The number of animals entering estrus rises in the fall At this time the pregnancy rate is at its lowest point and a high percentage of anestrous (nonpregnant) animals are in the population Breeding activity increases and at this time the number of sexually mature males in 1female areas increases

The number of unimplanted pregnancies begins to rise sharply in October and November At the smne time the nlli~ber of anestrous animals drops as these animals enter estrus breed and become pregnant Thj_s is the peak breeding season

The birth rate is at its louest this period as is shovm by the lmv number of post-partum a1imals and the lovr number of large fetuses

By late November and December the pregnant rate is increas even though the birth rate is slightly This is due to ne7

blastocysts implanting as is shown the increase in Class 1 fetuses

By January the period of t breedLng activity has declined and the number of unimplanted animals ceaches a peak as the number of blastocysts implanting equals the number of The lanted pregnancy rate then begins to declLne as fevJer animals breed and more implant The greater rate of antation is reflected in a rise in the Class 1 fetuses as well as an overall increas in the er of lanted pregnancies However the rate of implantation is partially obscured by an increased birth rate removing animals from the total of lanted pregnancies This increase in the birth rate is shmm in the rise in post-partum females and a rise in Class 5 fetuses

This condition of a low rate of breeding high rate of implantation and increasing rate of birth lasts from February to Hay

At this point He come to the least reliable data The sumner are smaller gtJere collected over a period of seven weeks and were from animals that died as the result of capture and handling

Therefore tvhile the data shaH that the post-partum rate decreases sh indicating a reduction in the birth rate~ the number of Class 5 fetuses

a peak indicating a very high birth rate The true birth rate is probably some1middothere in betigtJeen Females 1-ith very young pups (hence are post-partum) are vlary and probably less likely to be caught in nets Females with fetuses appear to be more likely to from handling In several cases~ twisting of the reproductive tract by a fetus vJas the actual cause of death Anestrous females (including post partum) on the other hand are less likely to die

Reproduction - 4 - March 1971

There is no question that the birth rate remains high during June and July Kenyons (1969) sum11er sample is limited but shows a large number of Class 5 fetuses Again the sample is biased in favor of pregnampIt animals but not to large fetuses only His field counts as vJell as observations by most other observers indicate an increase in the nUIlber of dependent young througbout the surnmer The number of Lmted pregnancies declines throughout the suTimer tvhile the unimplanted pregnancy rate declines at a slmver rate and breeding remains fairly constant at a lm-v level This indicates that through June there are more implantations than conceptions hmvever the birth rate is substantially greater than the implantation rate By late July and August there are relatively fe1r implantations taking place as shmm by a leveling of the unimplanted pregnant rate and a lotr number of Class 1 fetuses The number of Class 5 fetuses declines as a result of births during June July and August

Therefore the season of greatest pupping may be quite broad running from April through August possibly with the peak occurring in late May or June Until a better summer sample is available tve cannot pin dmm the peak of this pupping season with certainty (See Fetal GrmmiddotJth Rate for further discussion on breeding and pupp peaks)

If the information were precise we could compare subtracting numbers of pups born and numbers of conceptions to deterrnine the actual percent of ferrales breeding implantLng ltmd pupping in each month Unfortunately the data are not Each curve is influenced by more than one factor and ue do not knmmiddotr for sure the magnitude of any of these factors Hmmiddotrever by comparing some of the major samples e may be able to get a rough idea of how much higher the birth rate is at one time of year than another

The combined Mty and the combined September and October samples provide the best comparison Both are large samples One occurs near the peak of pupp and a lmv point in breeding and the other is near the low point in pupping and the peak of b

The number of Height Class 5 fetuses~ post-partum femaless and proestrousshyestrous females come closest to represent a particular event The time spent in each category is comparatively short

There were approximately three times as many Class 5 fetuses in Hay thanmiddot in Septerqber-October There Here also about three times as wany postshypartum females in May In tember-October there were two and a half to three times as many proestrous-estrous animals as in Hay It ~middotwuld appear that the peaks of breeding and pupping are roughly three times as high as the lowest points

At the lmmiddot12st period of pupping 2 to 3 percent of the females have Class 5 fetuses and should pup middotJithin a month If vm assume the post-partum stage lasts L 5 to 2 months (see Recovery of the Uterus) it also appears that 2 to 3 percent pup in the ltwrest months of pupping

Reproduction - 5 - March 1971

Similarly about 3 percent of the females have enlarged follicles during the lmvest period of breeding He do not knmv how long this period lasts hovever

At the peak of pupping about 10 of the females have Class 5 fetuses and should pup in the next month About 15 are post partum indicating about 8 births per 100 females in the preceding month

These percentages are prob2bly slightly high because of hunter bias avoiding females with pups It appeatmiddots that at least 2 to 3 percent of the mature females breed and 2 or 3 percent have pups in any month of the year In the peak breedingmonths such as September and October perhaps 7 to 10 percent breed and similarly in peak pupping months sud1 as Nay 7 to 10 percent of the mature females pup

Gestation Period

Barabash-Nikiforov (1947) listed the gestation period of 8 to 9 months This apparently was based on the case where a fentele mated in captivity as reported by i-Ialkovich (19 37) According to Barabash-Nikiforov a pup was born 8 months later although it died While it was fully formed~ it may have been premature

A number of births have occurred in captive animiddotmals held by the of Fish and Game in recent years In all cases the pu-ps ~-ere sc-middot1all usually less tha1 1600 g but still than the smallest pups found in the -rild All died after birth

The gestation period of 9 months has been repeated in the literature but apparently these statemsnts are based on Barabash~Nikiforol (19lf7)

Lensink (1962) also estimated a gestation period of 8-9 months based on field observations He felt that the peak of breeding was in August or September and the of pupping ~Alas in l-1arch or April

It has been demonstrated that the gestation of sea otters a relatively long period of tation (Sinha 1966 Kenyon 1969 and present study) Therefore any attempt to estimate the length of gestation by examinatioD of tracts must take both the unimp1anted and Janted periods into accour1t

Kenyon (1969) presented an estimate of the implanted period by assuming that the cube roots of fetus weights fall in a straight line after the embryo is established (Hugget and Hiddas 1951) and by that the European otter and Aluerican river otter would have similar fetal growth rates By plotting tenn fetus on a line established by the European otter he estimated an gestation of about 120 days not including the period for establishment of the ewbryo

Kenyon (1969) then an estimate made by Dr D G Chapman using the method of Hugget and ~-Jiddas (1951) v7here a regression line is fitted

Reproduction - 6 - March 1971

to the cube roots of the raeail of the mean weights of fetuses in each of five vreight clas3es plotted t the mean time of year

From the regression coefficient ob he estimated an implanted period of 154 days By taking the mean of the pregnant animals that are unimplanted pregnant estimated the tmimplanted period as about 75 months 1Ulmv-ing a half month for establishment the embryo he estimates a total gestation of 12-13 months

The data used by Chapman included a January-February sample of 26 a March-April sample of 49 and a Nay-August of only 9 The surtmler sample is more biased than the others and quite small yet the estimate relies heavily on this sa~ple when fetus size is the greatest

Table 2 presents a neTw- estimate of the length of the implanted period using the method used by Chapman but including data from the present study in addition to Chapmans data Table 3 presents a similar estimate of the length of the unimplanted period

The following is a comparison of the two estimates

Unimplanted Period 230 days 66 days Establishment of Embryo 15 15 days Implanted Period 154_~~Yrgt~ _73 d~s

Total Gestation Period 399 d2ys 154 days or about 13 months 5 months

Obviously there is some problem tiith the the technique or both A similar calculation mcde before the Hay sample had been collected produced an estimate of 109 for the implanted period and about 8 months for the total ation period

This method of calculation assur1es that there are definite peaks in breeding and pupping If breeding and pupping occurred evenly throughout the year the average fetus size 1vould remain constant thruughout the year and the technique would not tvork An ideal situation v10uld be when all individuals breed and pup 1vi thin very short periods Sea otters do have but are not well defined Very large s v-ould be required in such a case because there are ahvays fetuses of all sizes and the periods of maximum breeding are broad

Obviously Chapmans sample vas inadequate llthile the number of fetuses available for the latest estimate seems large and are well distributed throughout the year the a_ctual nu~ober in any cne fetus vJeight class at any one time of year is small A look at Table 1 shows that there is little consistency in many of the classes (Fig 3 shmvs that Class 1 and 5 fetuses do fit a pattern)

Table 2 Estimated Length of Period

Percent of Fetuses in each Height Class

Time of Collection After January

1 month 31 15 15 38 0

3 months 18 12 22 35 12

5 months 18 8 8 36 30

7 months 3 8 16 29 45

9 months 20 7 10 40 23

10 months

14 18 12 33 23

---~--

r1ean Time in Nonths After January 50 57 53 58 70

Cube Root of Umveighted Mean 063 17 36 7lf lllf

Specific Grmth Velocity 0169day EstirGated Implanted Period = 73 days

Table 3 Length of Unimplanted Period

Months After Unimplanted Unimplanted Percent January Pregnant Pregnant Unimplanted

1 month 35 26 58

3 months 49 49 so

5 months 128 179 42

7 months 22 37 37

9 months 50 57 47

10 months 55 44 56

Unweighted

48

Reproduction - 7 - March 1971

All of the estimates very heavily on su111mer samples 1vhen the fetuses are largest These sauples are knmm to be biased probably in favor of larger fetuses There are more fetuses at this time of year but it is exaggerated in these SCilllples

The latest estimate is not necess better than Chapmans even though it uses more and larger samples The velocity is twice as high as that calculated for fur seal and a 5 month gestation period is not consistent with other information available

This technique for estimating the lanted gestation period is not capable of providing a reliable estimate with the available data Therefore the 12 to 13 month estimate and any calculations based on it are meaningless

Much of the discrepancy lies in the estimate of the unimplanted period The present estimate that 48 percent of all pregnancies throughout the year are unimplanted based on an even distribution of relatively large samples and is probably more accurate than Chaprnan 1 s estimate of 60 percent

There are several other ways of guessing at the length of gestation Kenyons (1969) estillate of an lanted period of 4 months using the cube roots of term fetus weightscannot be relied upon entirely but it may be closer than the other estimates If gtmiddotJe estimate the uninplanted period from this using the 48 percent ted factor and allow for the establishment of the embryo we get a total gestation of about 85 months

Lensinks (1962) method of detennining the and peak pupping periods and taking the time between the peaks as the total gestation period has some merit TJnile Lens ink tried to determine these peaks by field observations we can determine them more accurately using the condition of reproductive tracts

Lensink felt that the peak of breeding middotlas in August or September The data in Fig 2 indicate that he may seen the beginning of the period of est breeding activity but that the probably in October or possibly November Lensink estimated period in larch or April Again the data indicate that he v1as s the beginning of the period of greatest pupping but the actual peak is probably in June or July At this time the number of Class 5 fetuses is greatest The nucJber of post-partum females probably and the number of implanted pregnant females is dropping from its If the gestation period of

11 11the average animal lasts from October or November to June or July we get a period of 8 or 9 months tThile Lensink 1 s was early 1 the duration agrees vith the present data

The peak of implantation when the greatest number of blastocysts are implanting can also be roughly estimated from 1 and 3 In February the number of unimplanted middot is dropping and the number of implanted pregnancies at this time th2 number of Class 1 fetuses probably reaches a

Reproduction - 8 - March 1971

Therefore it appears that the unimplanted period lasts 4 to 5 months and the implanted period from 4 to 5 months 1vi th a total gestation period of 8 to 9 months It should be recognized that these estimates are based on general trends and not distinct Therefore they are approximate However the relative length of the unimp and implanted p2riods agrees with the percentages noted previous and this estimate agrees vlith the one based on Kenyons (1969) comparison vdth other species of otters

If the gestation period ~Jere 12 months the percentage of pregnant animals would remain constant throughout the year If it vJere only slightly longer or slightly shorter there would be only a slight fluctuation unless there were a distinct breeding period Fig 4 indicates a substantial fluctuation indicating a gestation period substantially shorter or longer than 12 months

The peak of pregnancy occurs in February after the period of t breeding activi The luH point on the pregnancy curve occurs after those animals giving birth dur the period of t pupping have pupped That is the t point on the pregnancy curve occurs after the main breeding and the lmves ~ point is after the main pupping period These periods actually last for 4 or 5 months so these points are well after the peaks of breedtng and pupping

The time betbulleen the peak when the pregnant and the lmr point Hhen the fewest are the average gestation period This is about 85 months (Fig 4) The anestrus curve (Fig 1) shmvs the same thing betng a negative of the pregnancy curve

Hith three different methods we have estimated a gestation period of 8 to 9 months Hith the unimplanted period roughly equal to the implanted period This agreas bullmiddotlith the observed period in captivity (BarabashshyNikiforov 194 7) and with field estimates Any gestation period differirg greatly from this does not fit the data Conceivably one could apply a 20 to 21 month period to the SaTEpound information however if this 1vere the case females would have to breed vJhile accomp by a pup or be on a four year reproductive cycle This prospect stretches the imagination

There is a definite possibility that the gestatton period varies in sea otters as in land ot te~cs This most likely -middotwuld occur through variations in the unimplantec1 period The above discussion refers to the average gestation period Until better information to the contrary is obtained we must continue to assume that the average period is 8 to 9 months

Fetal Rate

the estimates of the gestation curve are corrects Kenyons (1969) Fig 4 should approximate the grouth curve of the fetus Fig 5 duplicates this and shmvs the actual Heights Fig 6 shous the same information more graphically It is possible to esttmgtte the of time in each v7eight classlt Fig 6 does not take the period for establishment of the embryo into account so Height Class 1 may be longer than shmm perhaps 15 days if we accept Chapcnans (Kenyon 1969) es e

--

3 0

~

shy

----- -shy --middotmiddot~middotmiddot

___ ___1__

_

(middot-)

_ ft -) J ~)___

~-

3

Reproduction - 9 - IYarch 1971

The short time span of Classes 2 and 3 explains the relatively small numbers and the lack of a consistent pattern found in those classes (Table 1)

Using this curve it should be possible to predict the time of birth and with less precision the time of Luplantation and conception of any particular fetus By plot the estimated birth dates of a large number of fetuses rve should be able to drav curves Ttlhich approximate the breeding implantation and pupping rates of the population throughout the year

Unfortunately there is overlap bet1veen samples collected at different times of year Some of the samples are too small to provide a comparison We vould have to give eight to each in order to combine them Therefore only the ttay September and October samples are used lfuere the September and October samples overlap t1lt10 separate points are used Fig 7 presents the estimated birth dates grouped by month Fig 8 separates the dates into half month groups February and Narch are not represented and April is an estimate based on post-partum females rather than fetus size (see Recovery of the Uterus after Parturition)

The peak in this curve may be exaggerated It has been shmvn previously that fetus size is less consistent bet-Jeen samples than the other categories (Table 1) because the sample size of each grouping is small Therefore the exact shape of the curve may not be correct If however we assume the approximate of the peak is correct we can construct a model of the based on our estimates of the gestation period besed on a fetal grmJth rate estimated from this estimated gestation Therefore our reasoning would be some~hat circular if -Je used thjs to prove the estimate Hmvever we can see how this model fits the rest of the data A good fit would tend to support the model and the estimates on which it is based

For this model He use the curve in 8 to represent births We assume a gestation period of 8 months -rith the unimplanted and implanted periods each lasting 4 months TheTefore v8 draw curves identical to the birth curve but moved backward 4 months for implantation of the blastocyst and 8 months for conception (Fig 9)

A comparison of the peaks in Fig 9 with the data in Figs 1 2 and 3 and the narrative b on those indicates a fairly close fit The data indicat that the peaks may ~ctually occur slightly later than indicated in 9 but in the timing and distances bet7een the peaks fit

The sharpness of the birth curve indicates that the breeding in1plantatimiddotm and pupping peaks mey be more distinct than indicated by the curves in Figs 1 and 2 It is possible to fit curves with more abrupt changes to the data Fig 10 shows the same data lith the curves drawn to more closely fit the model tfuether these curves are more accurate or not is open to debate The data on Hhich the birth curve is based is not that strong The numbers in each month are relatively snall If such distinct did occur it seems likely that field observers ~vould have seen the effects Nost field observations indicate broader peaks that are not gtvell defined

2

G

X

middot ~ -- (-- ) r~ ~ - - -- ~

yen~ r- ~--~

3

i__middot -~---

~

gt

- cshy

Reproduction - 10 - March 1971

~e of _Sexual Maturity

Tables 4 - 7 list the frequency of nmnbers of corpora albicantia and corpora lutea found in females of different areas It should be recognized that the ages may not al-1ays be exact Animals ATTichi tka to become sexually mature Hhen about 4 years old It appbull~ars that more animals may become mature earlier on Adak The information is too limited to dratmiddotJ any definite conclusions from this hmmiddotJever the food availabili ty is probably better at Adak and an earlier age of sexual 1naturity may be a response to better nutrition

Ovulation

From Tables 4 - 7 it can be seen that the maximum number of corpora albicantia for each age roughly the numb~r of years after sexual maturity that some animals ovulated once every year after maturity It can be assumed that many corpora albicantia are invisible macroscopically after several years particularly if they are not remnants of a corpus luteum of pregnancT Therefore it Ls possible that most females ovulate every year This poss ili ty is also supported by the high incidence of large follicles in lactating females (TabL~ 10)

faximum

Tables 4 ~ 7 indicate that at t some females continue to breed at a very old age The samples are teo small to determine 1rhether a t number become unproductive at any point

Ovulation

Many mustelids are induced ovulatoTs

On September 14 1967 a female vJas tsd Hhile copulating She appeared to have Oilnlated shortly before Because sea otters may copulate several times over a period of two or three (Kenyon 1969) ovulation may have been induced by a previous

ficance of

Delayed lon is a charact8ristic of the reproductive cycle of most mustelids Several theories on the ccdv of a delay have been advanced Most assume that spring is the most favorable time of year for survival of nemiddotJborn young but that either a wir1ter breeding season is not favorable or that the presence of a Ttlale vhich occurs only during the

season is for survival of the young of the previous pregnancy (1963) discusse the evolution of delayed implantation in mustelids and points out that there are exceptions even in arctic areas middotJhere time of birth be consLdered more may be born at any tin1e Scheduling the and made possible by delayed mey be advantageous but is not necessary

----

a - o lt bull y bull bull l w laquo bull _ ~ ~

Table 4 Frequency of occurance of Numbers of Corpora Albicantia Amchitkll - June-August 1968

N U M B E R 0 F C 0 R P 0 R A A L 8 I C A N T A

I

~I -AGE -~ 1 I 1084 6 92 5 7(Years) 1 middot shy

- 0-l - ~~ ~- middot~middot- middotmiddotmiddotmiddotmiddot

1-2

2

3

4 1 I

2 (1)

6

middot5

1 (1)1

2 (l) 17

1 ( 1 )

2 ( 1)

18

l (1) I ( 1 ) 9

10 1 ( 1) L ~j( 2 1 ( 1 )

2 ( l) 1

12

11

1 (1)

I I (I)

1

2 ( 1)

13

1 ( 1 )

2

114

115 I116

17

I 18

I I

I

119 I

Numbers inside parentheses =Number of individuals with corpus luteum

11

Table s Frequency of oc~urance of Numbers of Corpora A1bicantfa _Kanaga - October 1968

N U M B E R 0 F C 0 R P 0 R A Al B CANT IA - I I I

AGE 84 I I

5 I l 6 7z 3(Years) 1

l rbull ~middot--

--- shy

0-l

J l-2

~ 2

3

4 2 ( 1)

5 3

Ibullbull 6 6 (3)

4 (4)7

8 4

3 9 2 ( 1 )

j iI 10

I 1 (1)11

12

13

14

1 ( 1)

16

15

17

18

Numbers irisi

-

J

(Plus 2 with corpus 1 uteum but no corpora a 1 b i cant i a)4(1)

1

1 (1)

5 (1)

10(5)

3

3 1

3 (2)

2 ( 1) 3 (3)

1 ( 1)

2

1 ( 1 ) 5 ( 1)

1 ( 1 ) 3 ( 1)

1

3

2 ( 1 )

1 (l)

1 (1)4 ( 1)

2) 1 (1) 3 (3) 4 (1) 12 ( 1 ( 1 )

1 ( 1)

1

2

1 ( 1)

1 ( 1)

1

I I parentheses = Number of individuals with corpus luteum

11

Table 6 Frequency of occurance of Numbers of Corpora Albicantia Mid Pt-~Blind Pt~ Adak- October 1968

N U ~1 B E R 0 F c 0 R P 0 R A A L B I C A NT I A

-1AGE

~

9 1084 75 6Years) 1 2 3 - shy ----- --middot shy~--

- _ Q~L

1-2

(Plus one with_corpus luteum but no corpus a1bicans)2 1 l I I I I I ) (One with corpus luteum but no corpus albicans 3

4 2 (1)

l5

16 1 (1)

1 1)7

8

2 (1)9

10

111

12

I13

14

15 1 (1) 16

17

18

1 ( 1)

1 (l)

1

1

1

2

1

1

1

1(1 )

1

1 (1)

1 (1)

1

I

1

1

1

1

-I

Numbers Inside rentheses = Number of individuals with corpus luteum

1

Table 7- Frequency of occurance of Numbers of Corpora A1bicantia Three Arm Bay - Bay of Islands Adak - October 1968

AGE (Years)

Q-1

t-2

2

3

4

s 6

7

8

9

10

11

12

13

14

15

16

17

18

Numbers

N U ~ B E R 0 F C 0 R P 0 R A A l B C A N T I A

~L 3 4 slE - 1middotshy

7 1~8 __J~l_11 shy

1-

One -Ji th corpus 1uteum but no corpora _a 1bicantia I I I I I I I

One vJi th corpus luteum but no corpora al bicantia

12 I I (Plus one with corpumiddots luteum but no corpora albicantia)1 (1) 3

1 (1) 2 (1)

1 2 (2) l 1

1 (1) 4 (1)

2 (1)

2 1 )

1 ( 1 ) 1

2 (1)

2 (2)

l22 3

3 ( 1)2 21 ( 1)

2 ( 1)

2 ( 1)

l ( 1)1

1 1

1 I ll

I

I I

inside parentheses Nurnber of individuals with corpus luteurn

Reproduction - 11 - March 1971

The sea otter lives in an area of relatively uniform temperatures However storms tend to be more frequent and violent in fall and winter Survival may be better in the late spring and sunrmer Therefore there may be some advantage to birth in the late spring However there does not appear to be a great deal of advantage to a breeding season at any particular time of year

Wright (1963) pointed out that most musteHds lant after daylight begins to increase usually around February This is the period when sea otters implant at the t rate If the time of implantation is influenced by daylight the period when the greatest number of births occur could be shorter than the equivalent breeding period The data do not show this although the possibility cannot be ruled out

The fact that substantial numbers of sea otters are breeding implanting and pupping at all times of the year indicates that hile there may be some advantage to certain events taking place at a particular time of year that advantage is not great enough for a distinct breeding season to evolve

Sea otters exhibit certain characteristics that are comnon to most mustelid breeding cycles but these chfracteris tics are not as well developed as in most es

It is interesting to note that Hright (1963) correlated the molt of Mus with the implanted period TI1e molt began around the

ion and ended around parturi tioD

Sea otters molt all year and some are implanted pregnant at all times At this time e cannot say uhether any between molt and pregnancy exists

Fetal

Kenyon (1969) found that the number of caudally presented fetuses roughly equalled the nunber of cephali presented fetuses He suggests that this indicates a lack of tatim1 for birth occurring in water and infers that sea otters must give birth on land

Fetuses of all 1veights in the present study also appear to be randomly oriented however 97 Class 5 fetuses (1000 g and over) showed 60 percent cephalic ion while only 40 percent shovJed caudal presentation

The orientation of large fetuses should be a better indicator of orientation at birth Smaller fetuses may position Therefore it appears that more sea otters are born head first than tail first

If you accept Kenyons premise that caudal presentation is necessary or at least beneficial to birth in water this more recent information supports the supposition that birth occurs on land The little information available indicates that birth does occur on lando but there have been unconfirmed reports of birth in water

Reproduction - 12 - tltIarch 1971

At least some sirenians are born head first indicating that caudal presentation is not necessary North of Unimak Island a large population of sea otters lives off-shore There is little evidence that any numbers come ashore It seems likely that many of these animals are born in the water

Of 305 implant pregnancies 138 fetuses (45 percent) had implanted in the left horn and 167 (55 percent) in the right horn Most of the difference was in the 1970 Amchitka sample ~fithout this sarnple 48 percent implanted in the left horn and 52 percent in the right horn Kenyon (1969) found an equal number in each horn If a real difference exists it is probably exaggerated by the 1970 A~1chitka sample

Out of 305 implanted pregnencies the point of implantation was on the same side of the reproductive tract as the corpus luteum in all but three instances In one case the corpus lutaum was in the left ovary and the fetus in the right porn In the second case the corpus luteum in the right ovary but the fetus was in the left horn In the third case there was a corpus luteurn in each ovary and tHo fetuses (of different sexes) in the left horn

It appears that blastocysts rarely cross from one horn to the other

Because there is usually a long period of time parturition and the next est_rus is little if any inhibitory effect on ovulation caused by the corpus of the previous pregnancy The ovary producing the largest follicle appears to be random and there does not appear to be the tendency for a pregnancy to be from the opposite ovary from the previous pregnancy In many cases there are many more corpora albicantia in one ovary than in the other

When a loses a pup shortly after birth and enters estrus before the uterus has conpletely recovered and before the corpus luteum has completely degenerated there may be some inhibitory effect Of the ll+ such cases identified six had large follicles in the same ovary as the new corpus albicans and in the opposite ovary It appears that any inhibitory effect by the corpus luteum only lasts for a short time

A few cases bullv-ere found vJhere one ovary was not developed or othenvise not capable of producing ova Because one ovary may support repeated pregnancies such animals may be as productive as those ~vith both ovaries active

Kenyon (1969) reported that most implantations occur within the central third of the uterine horn This held true for the animals in the present study they may implant In one case with a near term fetus the placenta covered the at the base of the horn blocking the exit of the fetus

Reproductive - 13 - March 1971

Birth iltleigh t

Barabash-Nikiforov (1947) listed the vJeight of a ne-v1born sea otter as 2000 g Kenyon (1969) presents the best quantitative data published to this time He found a maximum fetus weight of 1869 g 1-Thi1e he found pups as small as 1020 g those under 3 lb (14 kg) had died shortly after birth He estimated that successful birth weights range from 3 to 5 1b dr 1 4 to 2 3 and for purposes of calculation assumes an average birth weight of 1850 g to 1900 g

The weights of Weight Class 5 fetuses the smallest pups found living in the wild and three pups born in captivity (all died within 24 hours) are presented in Fig 11 These data tend to support Kenyons (1969) estimate of the ~veight at birth Few living pups middotHeighing less than 1350 g or about 3 lb and fevJ fetuses over 2000 g ( 4 5 lb) are found Kenyons upper estimate appears to be slightly high although there are extreme cases in both directions One pup weighed only 2 lb (900 and one fetus could not be weighed accurately on available equipment but weighed over 2500 g

One female with a 4 lb (1816 g) pup still had the placenta attached to the uterine wall and must have given birth shortly before being collected

From 11 it appears that most pups are born when they weigh in the neighborhood of 1800 to 1900 g in most of the populations sanpled Hmvever the and Delarof s indicate a bith Jei of between 1600 and 1700 g The Delarapound sample is srEall but the Tanaga sample the largest containing over a third of the large fetuses collected and was collected at a time of year when the birth rate is very high

This may be a reflection of the phys condition of the adult females and indirectly a reflection of food availab ty There are other factors indicating that the population is declining because of overpopulation

This lov1er birth yeight raises the question of middotwhether the pups are born earlier or whether are in poorer condition when born In an effort to gain some insight into this the fetal of Class 4 and 5 fetuses from Tanaga were plotted against their total lengths (Fig 12) middot Fetuses from other eastern Adak which appears to be a very healthy population were checked this curve fonaed by the Tanaga fetuses In all cases they fell Jell Jithin the limits of the Tanaga data indicating that there is no difference in the of fetuses of a given length It seems unlikely that a reduction in the rate of weight gain vould be completely proportional to a reduction of linear growth It appears that the rate of growth of fetuses is relatively constant but that females in poor physical condition may not be able to support as large a fetus as those in good condition so parturition occurs earlier

This may have sone effect on information on this

Fig 12 demonstrates some other interesting points There is no difference in the ratio and the maximum fetus size of males and females

~

(

~

-s 2 0 C

U)

0

--

ez ()

~) I) middot _t t- (

j -~middot -~

Reproduction - 14 - March 1971

This infeTs although dolt2s not prove an identical grmgtlth rate afld birth weight for males and females The circles on Fig 12 show the ~eight-length ratio of pups These fall below the curve formed by fetus weight-length ratios indicating that pups of a given length are some-Jhat heavier than fetuses of the sarne length This indicates an increase in weight immediately after parturition

Sex Ratio at Birth

Kenyon (1969) found that of 58 fetuses for which sex could be determined~ 45 percent vJere male and percent were female He assumed an equal sex ratio until a larger sample could be obtained

In the present study 111 fetuses were male and 144 ~vere female When this sample is combined with Kenyonts the sex ratio of 313 fetuses is 44 percent male and 56 percent female

Sex Ratio of the

In the process of harvesting capturing animals for transplants and delineating male and femalen areas it has become apparent that there are more females in the population than males Our general ion is that perhaps 60 to 65 percent of the sea otters are females Kenyon (1969) demonstrated that more juvenile males than females are found dead on the beaches of Amchitka lhe age structure of harvested in 1968 indicates that males may not live as long as fetnales

The lower birth rate higher juvenile mortality rate and slightly shorter life expectancy of males could account for the unbalanced sex ratio in the population

The sex ratio may vary from one island to another Presumably there -vmuld be more females in a population that has reached the carrying capacity of the habitat where juvenile mortality is higher There is some indication that males are the first to expand into new areas of unoccupied habitat Therefore in recently populated anas one might find more males However this situation vould be

Lit

The normal litter size of the sea otter is one This has been recorded by almost every observer since Steller Barabash-Nikiforov (194 7) reported twins in utero and mentioned other reports of twin fetuses from the early days of SnoF (1910) recorded a set of neHborn tdns In more recent studies~ both Sinha et al (1966) and Kenyon (1969) found reproductive tracts with two corpora lutee but in all cases only one fetus las present In thousands of hours of observation by many observers no tbullvin pups have been reported

In the present study 24 instances of multiple ovulations were recorded Five of these resulted in hv-in and one in triplet fetuses The information is summarized in Table 8

Table 8 Multiple Corpora Lutea Fetuses

Total Pregnant Females 565

Implanted Pregnancies 316

Unimplanted with 2 CL (corpora lutea) 9

Implanted Pregnancies with ) L CL and 1 Fetus 8

Implanted Pregnancies Vlith 3 CL and 1 Fetus 1

Implanted Pregnancies gtvith 2 CL and 2 Fetuses 5

Implanted Pregnancies middotwith 3 CL and gt Fetuses 1

Total Nultiple Ovulations

I

Percent Nultiple Ovulations 42

Percent of Pregnancies tvith Nultiple Fetuses 19

Corpora Lutea per Pregnancy 105

Fetuses per 102

Reproduction - 15 - lvlarch 1971

From these data it appears that in over 4 percent of the estrus cycles more than one ovulation takes place Of these about half result in the development of more than one fetus In most cases the were of a relatively large size and probably would have been born normally

All multiple fetuses appeared to be the result separate ovulations All had separate placentas and in some cases were different sexes Four tracts with twins had both fetuses in the same horn one had one in each horn and the tract with triplets had one fetus in one horn and t~vo

in the other Another tract had skeletal remains of triplets in one horn that were being resorbed Another tract appeared to have had five fetuses in one horn but they were almost completely resorbed One might infer that physically there is not enough room in one horn to accomn10date more than two fetuses for any length of time

With the exception of the newborn tvins reported by Snow (1910) there are no docu_mented records of female sea otters with tvin pups There are occasional unconfirmed reports of twin pups but as Kenyon (1969) points out a female may tolerate a pup in addition to her ovm but it is unlikely that a female could care for more than one pup It is unlikely that more than one pup survi~es In any case orily 2 percent of the pregnancies produce more than one pup 1middot1ultiple births cannot be a significant factor in the productivity of sea otter populations

The presence of a corpus luteum without gross evidence of implantation t-7as assumed to be an unimplanted pregnancy No attempt ~vas made to locate unimplanted blastocysts As a result e have little information concerning the survival of the blastocyst Of 206 reproductive tracts classified as nulliparous on the basis of the appearance of the uterus 12 had one corpus albicens and one had two Some of these may have been large attritic follicles Others may have ovulated and even conceived but implantation did not occur All of these cases represent the failure of the antmals first estrus

A total of 16 resorptions or possible abortions were identified (Table 1) The follmJing is a brief descdption of these ~7i th possible causes of some

Des

Resorption of blastocyst or ovum (corpus luteum 4 degenerating no evidence of ion)

Failure at time of (possibly because 1 of growmiddotth inside horn adjacent to implantation site)

Resorbed or aborted fetus (implantation at end of horn) 3

Resorbed fetus (umbilicus tvTisted tightly) 1

Resorbed fetuses (three or more fetuses in one horn) 2

Resorbed fetus (cause unknmm) 5

Reproduction - 16 - March 1971

Six of these resorptions may have been caused by a lack of room for growth of the placenta because of crowding from other placentas a growth or the end of the uterine horn These six and the one vith the twisted umbilicus are probably the result of random events or accidents

The numbers of resorptions are too small to accurately measure the frequency of such occurrences although the relatively large number in the Tanaga sample (Table 1) may be a reflection of the physical condition of the animals Part of this number is probably due to the fact that there are more pregnant animals in the sample In general the samples with the most pregnancies also conta~~ned the most resorptions Half of the Tanaga resorptions can be attributed to random events but five vere still due to unknown causes Eight of the 10 resorptions fo1md on Tanaga came from a 20-mile stretch of shore about 25 percent of the harvest area

While concrete conclusions cannot be drawn from these data they do raise the possibiliy that under certain conditions 5 percent of the pregnancies may fail

near Parturition

Collecting methods were such that the presence of a pup with a female was often overlooked Therefore the lack of a pup with a post-partum or lactating female did not mean that the pup had died However 14 females shmving of recent pregnancy were estrus presumably having lost their pup shortly before or shortly parturition

Again the magnitude of this mortality cannot be determined but it appears to be of the same order as the in utero mortality

after

Reproductive tracts were subjectively classified as post-partum or anestrous on the basis of degeneration of the corpus luteum (or appearance of the corpus albicans) the size of the horn of pregnancy and appearance of the placent scar In an effort to get an idea of how long it takes for the tract to return to normal the size of pups accompanying females in each category is listed in Table 9

It appears that females pass the subjective boundary from post-middotpartum to anestrus Hhen the pup about 10 lb and is around 75 to 80 em long

We do not know a great deal about the early growth of a sea otter pup but judging from the cur1e e9tablished based on cementum deposition of older animals and from tooth eruption a 10 lb pup is probably in its second month of life

Table 9 Pups Accompanying Pos artum Females

Sex llleight

M 2 lb F 3 F 4 M 5 F 55 F 625 F 775 M 10 F 11

Total Length

47 em 52 56 60 65 65 70 81 (borderline post partum-anestrous) 82 (late post-partum)

Pups Accompanying Anestrous Females

F 10 lb 78 em M 11 79 M 12 8l F 13 83 M 13 92 H 16 83 M 17 91 1 19 87 M 19 90 F 21 107 F 22 96 1 22 101 M 2Llmiddot 97 1 26 100 F 28 103

Reproduction - 17 - Narch 1971

Frequency of Breeding

Steller (1751) reported female sea otters with ne~vborn pups also accompanied by another pup that seemed to be no more than a year old Nurie (1940) recorded an instance of copulation by a female accompanied by a pup and Jones (1952) caught a young pup simultaneously gtvith a copulating pair Lensink (1962) felt that few females breed until the pup is a year old but mentions females with small pups also accompanied by large pups

I believe that some of thes2 observations may be misinterpreations of the situation Females will occasionally leave small pups for a time often near other animals The pups Lensinkmentions would be 2 years old and probably veigh 30-35 lb Sea otters are gregarious and subadults may remain near other animals appearing to be accompanying their mother I suggest that most of Stellers and Lens inks large pups ere such cases

Females gtvi th pups probably do breed occasionally but the most recent evidence indicates that this rarely occurs Kenyon (1969) records no instances of females accompanied by pups copulating and states that he found no females- knm7n to be accompanied by a pup that were This holds true for the animals collected in the present study However the nature of the harvesting operation is such that many females accompanied by pups were not recorded as such Therefore we are forced to look at lactating and assume that they were accompanied by a pup at the time of collection or shortly before le 10 sm1marizes the reproductive condition of the lactat females In the 1967 and 1968 samples some animals vith enlarged marmary middotmre listed as lactating As a result some females with near tenrr fetuses Here included Because this condition is considered to be associated with the unborn pup rather than a previous pup these animals were with the anestrous animals in le 10 In 1969 and 1970 the presence of milk was used as the sole criterion and no term animals we-rmiddote in the sample

Of 246 lactating females only one had an ted fetus (excluding the near term animals from 1967-68) Several others had small corpora lutea indicating that they recently become or possibly had large

vhich had luteinized but tvere not actually pregnant The remainder had follicles

This information that accompanied by a pup rnay enter proestrus and even ovulate but that they only rarely mate That they enter estrus is subs iated by the f2ct that the number of corpora albicantia in many tracts equals the age of the amplimal minus the three years prior to nBturity (see les 4-7) This indicates that large follicles tend to develop year after sexual nl8turi ty is reached whether the female is accompanied by a pup or notlt

Some of those lactating females with s1all corpora lutea may have recently weaned or othenvise lost a pup and are already pregnant again Malkovich (1937) took a pup least 3 months old) away in cap She mated 12 days later here she had the male It appears that a female enters estrus shortly after Heaning

Table 10 Reproductive Condition of Lactating Females

Location

Bay of Is Adak I Sept 1967 9 1 100

Allchitka I Sept-Oct 1967 17 2 105

Mid Pt - Blind Pt Oct 1968 19 3 136 Adak I

Bay of Is Adak I Oct 1968 22 4 154 3

Kanaga I Oct 1968 32 s- 135

Al1chitka I July-Aug 1969 4 2 333

iTanaga I May 1970 56 10 152

Delarof Is May 1970 18 3 143

Amchitka I May 1970 36 3 77

---~--middot--middot----~-~----~----middot~-middot--~----~----- -------~-~ middotmiddot--middot-shy

TOTAL 213 33 134

Anestrous or pregnant with term fetus

]_ Large follicles or corpus luteum present

3 Includes one implanted pregnancy with small fetus

Reproduction - 18 - March 1971

The question of hml long it takes for a female to enter estrus again after losing a small pup arises Eleven tracts of the 1970 sample and ttvo of the 1968 Kanaga Sampnple shmJed signs of being pregnant recently) but were in proestrus These animals had an enlarged horn usually middotlith a slightly pigmented area but no actual placental scar a11d a recently fonned corpus albicans However they also had enlarged follicles and the tvalls of the uterus were typical of proes trous animals Another animal was similar but appeared to have already ovulated and a corpus luteum was present In these cases the female has lost a pup either through abortion or vithin the first veeks after parturition and has started into another estrus cycle Three or four of these had follicles vhich may have started to become attritic This may be because the uterus had not recovered sufficiently from the last pregnancy

The implication of this information is that with a normal pregnancy and successful rearing of a pup the female becomes pregnac1t only after the pup has been weaned This n~y be a year after parturition However if the pregnancy fails or if a pup dies even at birth the female may become pregnant again in a few weeks rather than _waiting the full year

Kenyon (1969) suggests that the period middotbetween and the next estrus may long because he found animals that tvere not lactating but had a faint placental scar and an indistinct but recent corpus albicans These animals ltJere cLtssi as anestrus in the present study They are undoubtedly bet1veen veaning of their last pup and the next estrus The number of suh animals is t durirg Sspte12~ cnd October At this time the rate of estrus increases a1d the number of anestrous animals decreases sharply the next feJ months Therefore many of these anestrous anirrals become pregnant 1lithin a month or two Kenyon (1969) suggests that if the female keeps the pup for about a year that the period between births may be smreltmiddotJhat greater than two years He is assuming a 12-13 month gestation period However the present study shows that the estimate of a 12-13 month tation period was based on an inadequate sample Estimates in the present study put this period at closer to 8 or 9 months Presumably the anestrous nonlactating females are in the 3 to 4 month period

This does not change Kenyons (1969) point that the and the next estrus may long It indicates that in a normal cycle it may be months Hmvever as it vvas shmvn above~ it is possible for a female to enter estrus very shortly after losing a pup

We have demonstrated that sea otters breed at all times of year and it is possible for a female to become pregnampJt again after losing a pup sooner than she vould have if the pup had survived HmmiddotJever we have also demonstrated that annual of b and pupping occur and that these are similar in different years and in different populations The assumption can be made that the average length of the entire reproductive cycle is some multiple of a year

Kenyon (1969) assumes that tbe pup remains with its mother for about a year ~-Ieights and measurements cementum deposition skull changes~

Reproduction - 19 - March 1971

observations in the wild and in captivity all support this assumption although it cannot be said with certainty that it averages 12 months It could be s tly ITDre or less

Assuming about 12 months for rearing the pup 8 to 9 months gestation plus a fev months rest bet~veen and the next estrus the average reproductive cycle takes two years

This is supported by Fig 4 which shows that slightly over half the mature females are pregnant in a year Kenyon (1969) pointed out that his January and Harch samples gtvere biased against females with pups a1d therefore in favor of pregnant females because of selective hunting This bias applies to the fall sa~ples as well The summer samples were biased similarly because pregnant females appeared to be less resistant to handling during capture Therefore the percent of pregnant females is shown to be higher than lt actually Has -rhile no correction factor can be applied to all samples the evidence indicates that the percent of pregnaJt females is usually around 10-middot15 lmrer than in the sample For example the 1970 Alnchitka sltLrnple t-ras less biased and was about 9 percent lower than the Tanaga sample vrhich is l)nm-rn to be more biased Also on a June skiff survey 7 percent of the animals counted had pups that were readily identified If ~re assune that this is the nunber of animals avoided it can be calculated that 15 of the sexually mature females all of vhich are t middotTen~ avoided Actually this will vary with the hunter weather time of year etc Hmvever it indicates the order of magnitude of the bias

Using this as a rough correctton for the data in 4 we find that about half or slightly fe-Jer of the sexually mature females are pregnant each year or as indicated ly the average female has a pup every two years

Reproduction - 20 - March 7 1971

Conclusions

1 While sea otters may breed or pup at any time of year there are seasonal fluctuations in the illliTlbers of females in each reproductive stage The pattern of these fluctuations remains the same from year to year and population to population

2 Breeding activity is highest in September October and November

3 More implantations occur during January February and March than at other times of year

4 The birth rate is highest during April Y~y and June

5 Tlvo and a half to three times as many females breed during the peak breeding months as during the months of lowest breeding activity

6 Similarly two and a half to three times as many females pup during peak pupping months

7 During the period of lovest b activity seually mature females breed at a rate of 2 to 3 percent per month During the peak of breeding 7 to 10 percent per month breed

8 During the period of lov1est pupping activity 2 or 3 percent per month pup and during the peak of pupping 7 to 10 percent per month pup

9 The gestation period about 8 to 9 months on the average About half of this time is ~~ ted period

10 Female sea otters become sexually mature ihen 3 to 4 years old

11 Females may ovulate on an average of once a year after sexual maturity is reached

12 Females continue to breed at Cn old age

13 Sea otters retain certain characteristics common to the reproductive cycles of many other mustelids however exceptions are common Scheduling of the reproductive cycle during the year may have some advantages or it may be a trait that has not been entirely lost through evolution

14 Sixty percent of the near term fetuses are cephalicly oriented and 40 percent are candllly oriented

15 Sea otters probably birth both on land and in the water

16 Blastocysts rarely cross from one uterine horn to the other

17 Consecutive pregnanctes may occur in the same horn The side of pregnancy appears random

Reproduction - 21 - March 1971

18 Implantation usually occurs in the central third of the horn but it may occur any~rhere in the horn

19 Birth weights may range from 900 g to over 2500 g however most pups weigh 1800 g to 1900 g at birth

20 Females from areas gtvhere the population is declining because of food shortages may pup earlier and as a result the pups 1 birth weight may average 200 g lmver than nonnal

21 Male and female fetuses gr01middot1 at the same rate ampJd are the same size at birth

22 Body condition of the female has little effect on the growth rate of the fetus

23 There is a rapid ~Jeight gain shortly after parturition

24 The sex ratio at birth is 44 percent male and 56 percent female

25 The normal litter size is one hmmiddotever in 2 percent of the implanted pregnancies two or fetuses survive to near term

26 Multiple ovulations occur in 4 percent of the estrus cycles

27 es do not appeErc- to be able to more than two fetuses in a single horn

28 Up to 5 percent of the p may fail dne to in after implantation

29 An unknown but probably significant number of first pregnancies fail before implantation

30 The uterus is usually considered recovered from a pregnancy when the pup vleighs about 10 lb and is probably in its second month of life

31 Sexually matu1middote females normally have one pup every two years

32 Females vrith a pup may ovulate but rarely mate

33 Slightly less than half of the sexually mature females are in any given year

34 A female may enter estrus within a few weeks of losing a pup Consequently she may become agaln gtvi thout waiting the same length of time as if the pup hacl survived

35 Hhen a pup is weaned normally there is a longer period perhaps 3 or 4 months between weaning and the next estrus

Reproduction - 22 - March 1971

CITED

Barabash-Nikiforov I I 1947 The sea otter (Kalan) Soviet Ministrov RSFSR Translated from Russian by Dr A Birron and Z S Cole Israel Program for Scientific Translation 1962 227 pp

Hugget A St G and W F Widdas 1951 The relationship betlveen marmnalian foetal -reight and conception age Jour Physiology 144306-317

Kenyon K W 1969 The sea otter in the eastern Pacific Ocean USFWS North American Fauna No 68

Malkovich T A 1937 The sea otter in captivity Priroda No 381-87 Translated by J T Naximovitch 1966 Fisheries Research Board of Canada Nanaimo BC Translation Series 657

Sinha A A C H Conavay and K t-J Kenyon 1966 Reproduction in the female sea otter J Hildl Ngrrit 39(1)121-130

Snow H J 1910 In forbidden seas Edward Arnold London 303 pp

Wright P L 1963 Variations in reproductive cycles in North Arrierican mustelids In Enders A C ed Delayed tation Univ of Chicago Press 318 pp

  • SEA OTTER STUDIES - 1970
    • Surveys
    • Harvests and Transplants
      • Sexual Segregation in Sea Otter Populations
      • Reproduction in the Female
        • AERIAL COUNT OF SEA OTTERS- SOUTHSIDE OF THE ALASKA PENINSULA
          • Description of Conditions During March 1970 Aerial Count of Sea Otters
          • Discussion of Results
          • Sanak Island-Sandman Reefs
          • Shumagin Islands
          • Sutwick Area
          • Augustine Island-Cape Douglas
            • SUMMARY OF SEA OTTER SURVEYS
            • Amagat 1 to False Pass Ikatan Peninisula Cape Lazaref
            • Sanak Is
            • Deer 1
            • Pavlof Is
            • Northern Shumagin Is
            • Cold Bay 1
            • Kupreanof Peninsula
            • Mitrofania 1 to Kupreanof Pennisula
Page 3: Sea otter studies - 1970 - Alaska

AERIAL COUNT OF SEA OTTERS- SOUTHSIDE OFTHE ALASKA PENINSULA

March 1970

Between March 16 and March 27 1970 an aerial count of sea otters was rnade along the south side of the Alaska Peninsula from Shaw Island to Cape Lazaref on Unlmak Island Sanak Island the Sandman Reefs the Pavlof Islands northern Shumagin Islands and Sutwick Island were included A BLM Aero Commander N6392U was flown by Cal Ward at an altitude averaging 150 feet and an airspeed bullgtf 100 mphmiddot Observers were Karl Schneider next to the pilot and Jim Faro behind the pilot This is the same technique aircraft pilot and observers as were used in the 1969 Aleutian count The count on March 27 was made in a Gruman Goose 1r1ith Robert Wolfe substituting for Faro Pups with their mothers were not counted

Because of conditions of visibility and distribution of animals all of the counts made are considered to be low

In general the variability in factors influencing this type of count is so great that the results are not rei iable for population estimates or for determining short-term fluctuations in dense populations However aerial counts should be useful to determine general distribution and abundance and to follow large changes in population size Such changes are occurring where relatively dense populations are expanding into habitat that is either unshyoccupied or sparsely occupied Aerial counts are also the quickest way to familiarize new personnel with large areas of habitat

The following is a 1ist of definitions of terms used to describe counting conditions Application of these terms is completely subjective and based on the writers experience

Excellent- Surface of water relatively calm Usually overcast with 1ittle surface glare Single animals easy to spot at a distance

Very good - Surface may be slightly choppy but not enough confusing reflection to prevent spotting of animals off shore

Good - Off shore animals may be difficult to spot but single animals near shore and in kelp beds and small pods everywhere are readily spotted

Fair -Usually choppy or surface glare Single animals ~in kelp beds or in the lee of rocks or shore and most groups of animals relatively easy to see Other single animals and some pods in deep bays in the shadow of cliffs or off shore may be missed

Poor -Single animals difficult to spot and many pods may be missed but conditions still good enough to get a rough idea of the distribution of animals

The conditions encountered for each area are listed below

Description of Conditions During March 1970 Aerial Count of Sea Otters

March 20 Aniakchak Bay to Chignik Lagoon (1022 AM- 1240 PM) Conditions were generally fair with a heavy chop The south side of Sutwick Island and the area between the island and mainland were poor with heavy surf Visibility was very good inside Kujulik Bay however many animals were scattered throughout the area On two passes ~cross the bay 12 and 27 were seen therefore many were probably missed Similar scattering occured in Chignik Bay The overall count for the area is probably low

Pavlof Bay to Cold Bay (130PM-200PM) Conditions poor with heavy chop and squalls The count was superficial however a second superfi cia 1 count of this a rea was made on March 21 and only one otter was seen in Cold Bay

Cold Bay to Caee Lazaref (310PM -420PM) Conditions very poor fair in a few spots Heavy chop dense kelp

March 21 Northern Shumagin Islands (943AM- 1137 AM) Poo~ with chop on windward sides Fai_r on leeward sides (westerly wind) however air turbulence forced us away from high cliffs Heavy surf created very poor conditions on the south side of Unga

Pavlof Islands (1145 AM- 125ff PM) Conditions similar to those in the Shumagins

Chignik Lagoon to Pavlof Bay (305 PM- 535 PM) Conditions varying from fair to poor with chop and surface glare Count concentrated on points and better looking areas Most deep bays not completely surveyed

March 22 Sanak Islands (849AM- 1020 AM) Conditions fair to poor with chop on south side and surface glare on north side Animals scattered and very difficult to see Many otters on rocks Count is probably very low

Sandman Reefs ( 1025 AM~ 1115 AM) Conditions similar to Sanak With the exception of one pod of 450+ the animals were widely scattered and very difficult to pick out Only those very close to the aircraft could be seen Bad squalls moved in and the wind increased preventing a complete count of Deer Island and the eastern half of the reefs

March 23 Wide Bay to Aniakchak Bay (1123 AM- 140PM) Conditions generally good at first becoming fair after Cape Providence with a light chop and some surface glare Otters in Amber and Aniakchak Bays were widely scattered and many may have been missed

Port Heiden to Naknek_ Superficial counts were made from Ugashik Bay to Naknek on March 19 and from Port Heiden to Egegik on March 23 These were made under poor conditions by flying outside the surf line The water was choppy and full of silt No sea otters were seen

11arch 27 Shaw Island to Puale Bay (1030 AM - 1205 PM) Conditions very good to excellent High overcast and very little wind down to Katmai Bay After this the conditions deteriorated rapidly to fair and the count had to be stopped because of a combination of bad weather and lack of fuel As shown on the charts the count was concentrated on the points and islands however the good visibility allowed us to see well into the bays We feel certain no concentrations were missed

The numbers of sea otters counted are presented in the Table Specific locations and numbers are shown on the charts Where coverage of an area was not complete the approximate path of the aircraft is shown

Discussion of Results

Surveys along the Alaska Peninsula have been fragmentary and most of those made were done under less than ideal conditionsmiddot There are a number of reasons for this Weather tends to be poor along the south shore where squall5 form along the mountains Areas where an aircraft can refuel are north of the mountains and there are relatively few passes therefore range of the aircraft is a problem Also there is much shallow water off shore in the area from the Shumagin Islands to Sanak Island Coverage of this area is difficult and many animals are undoubtably missed

The present survey covered most of the area however conditions were such that most of the counts are probably quite low Despite these problems and a lack of continuity we have a fairly good picture of the recovery of sea otter populations in this area

Reports from various individuals are available from the 1930s and 1940s 11ajor surveys by the Fish and Wildlife Service in 1951 (Jones) 1957 (Lensink) 1962 and 1965 (Kenyon and Spencer) covered at least portions of the area In 1969 the southern Shumagin Islands were counted by the Alaska Department of Fish and Game and the present survey covered most of the rest of the area

Basically there are four distinct population nuclei in the area These centered around the Sanak Island-Sandman Reef area the Shumagin Island area the Sutwick Island area and the Augustine Island-Cape Douglas area The following discussion is based on information from reports by Lensink and Kenyon and from Alaska Department of Fish and Game surveys middot

~anak Island-Sandman Reefs

Small numbers of sea otters have been reported at Sanak Island since 1922 No reports came from the Sandman Reefs until 1942 In 1948 27 were sighted at

----Cherni Island The 1951 aerial survey showed 65 around Sanak and 97 in the Sandman Reefs In 1957 251 were seen around Sanak and 508 around the Sandman Reefs In addition two were seen in the Pavlof Islands however few were on the mainland In 1962 548 were counted in the Sanak area and 638 in the Sandman Reefs None were reported from the mainland or Pavlof lsland~however

much of the increase was in the northern area around Deer Island The 1962 survey was probably the best being made under excel lent conditions

AERIAL COUNT OF SEA OTTERS MARCH 1970

Partial or Location Count Date Camp 1ete Count Visibility

Cape Lazaref - Cold lsecty_

Cape Lazaref 3 320 Complete Poor

lkatan Peninsula 71 320 Complete Poor

False Pass - Amagat I 66 320 Camp 1ete Poor

Cold Bay 321 Partial Poor

TOTAL 141

Sanak Islands 239 322 Camp 1ete Fair to Poor

Sandman Reefs

Clubbing Rocks 12 322 Complete Fair to Poor

Cherni I amp Rocks 495+ 322 Complete fair to Poor

Goose I 7 322 Complete Fair to Poor

Hay I 18 322 Camp lete Fair to Poor

Hunt I 2 322 Complete Fair to Poor

Deer I 322 Partial Fair to Poor~

TOTAL 568+

Pavlof Islands

Inner 11iasik 2 321 Camp 1ete Fair to Poor

Outer lliasik 16 321 Camp 1ete Fair to Poor

Goloi 2 321 Complete Fair to Poor

Dolgoi 67 321 Complete Fair to Poor

Poperechnoi 29 321 Complete Fair to Poor

Ukolnoi 2 321 Complete Fair to Poor

yenWosnesenski 4 321 Complete Fair to Poor

TOTAL 122

Partial or Location Count Date Comp 1ete Count vis ib j 1i ty

Northern Shumagin Islands

Unga 184 321 Complete Fair to Poor

Popof 52 321 Complete Fair to Poor

Korovin 46 321 Complete Fair to Poor

Karpa __2t 321 Complete Fair to Poor

TOTAL 286

Cold ~ to Beaver ~ 0 320-21 Partial Poor

Beaver Bay to Kupreanof Pen

Beaver Bay 2 321 Partial Fair to Poor

Cape A 1 iaks in 4 321 Partial Fair to Poor

Guillemot I 16 321 Partial Fair to Poor

__1Kupreanof Peninsula 321 Partial Fair to Poor

TOTAL 23

Kupreanof Pen to Castle Cape 0 321 Partial Fair to Poor

AntJrrCastle Cape to-1-Ji e ~

Castle Cape-Castle Bay 16 321 Partial Fair to Poor

Chignik Bay 118 320 Complete Fair

Nakchamik I 5 320 Complete Fair

Hook Bay 13 320 Complete Fair

Cape Kum 1i un 88 320 Complete Fair

Kujul ik Bay 1199 320 Complete Very Good

Univikshak I 62 320 Complete Fair

Cape Kuml ik 54 320 Complete Fair to Poor

Sutwick I 14 320 Complete Fair to Poor

Cape Agutka 323 Complete Fair~

TOTAL 1766

Partial or Location Count Date Comp 1ete Count vis i b i 1i ty

Cape Kunmik 13 323 Complete Fair

Naka 1i kok Bay amp offshore islands 16 323 Complete Fair

Chiginagak Bay 5 323 Complete Fair

Agripina amp lmuya Bay 105 323 Complete Good

Wide Bay to Puale Bay N 0 T S U R V E Y E D

Puale Bay to c Kubugakl i __]_ 327 Partial Fair

TOTAL 146

Kashvik Bay to L Chiniak 0 327 Partial Very Good to Excellent

Cape Chiniak to Shaw _I

Shakun Is amp Rocks 71 327 Partial Very Good to Exce 11 ent

Kiukpal ik J to Shaw 1 0 327 Partial Very Good to Excellent

TOTAL 71

In the present survey which was made under relatively poor conditions 239 were seen in the Sanak area and 568+ in the Sandman Reefs The latter count Was incomp 1ete and conditions were such that the number of otters was more a function of time spent counting rather than area covered Obviously many were missed As a result not much can be said about total numbers The main change evident is that 141 were seen along the mainland and eastern portion of Unimak Island and 122 were seen in the Pavlof Islands This indicates that a fairly extensive movement northward and along the Peninsula is occuring

A remnant population probably remained along the south side of Sanak Island in the early 1900bulls By the late l940 1 s they began spreading into the Sandman Reefs This northward expansion has continued to the present time There is still unoccupied or sparsely populated habitat along the mainland shore and in the Pavlof Islands There should be continued expansion of this population for a number of years although the numbers around Sanak Island and the western Sandman Reefs may have already reached a peak

With the arrival of substantial numbers in the Pavlof Islands this population is probably on the verge of mixing with the Shumagin Island populshyation No doubt some individuals have moved back and forth in the past but the two populations are almost continuous at the present time

As in past surveys few otters were seen around the north side of Sanak Island and Caton Island This is probably poor habitat for otters The main population is around the south and west sides The higher count on the west end is probably more a result of intensive searching rather than a higher population

Shumagin Is 1 ands

This has been considered the largest of the four populations in the Alaska Peninsula area The most recent counts have not been adequate enough to show any major increase in numbers in the last decade however major changes in distribution have occured which are very similar to the pattern mentioned in the Sanak-Sandman population

There were occasional reports of sea otters in the Shumagins in the 1930s By 1947 Victor Scheffer estimated that 500 1 ived around Simeonof Island In 1953 Hooper counted 633 around Simeonof and Little Koniuj i Island The 1957 survey showed 1829 Most of these were- in the southern islands Five individuals were scattered in the northern islands and one was on the mainland shore near Elephant Point The 1962 survey totaled only 1352 The animals were scattered well off shore and the count was low However the count on Nagai increased from 149 to 338 indicating a continued northward expansion

In 1969 1510 were counted in the southern islands under relatively poor conditions An additional 286 were counted in the northern islands in the present survey and 23 were seen along the mainland north of the Shumagins Again survey conditions were not good

There is a very clear expansion of the population from a center near Simeonof Substantial numbers now occur on all the islands and repopulation of the mainland is occuring The population in the northern islands should continue to increase and most of the habitat on the mainland is not occupied

There is no evidence of an increase in total numbers since 1957middot However the last two surveys have been less than ideal and the large shallow off shore areas have not been covered adequately Considering the survey conditions and the obvious extentions of range it is almost certain that the populations from San-ak Island to the Shumagins have continued to increase In general it appears that the southern islands and reefs have become fully populated and the northern areas are just developing significant populations The entire a rea may be comp 1ete 1 y repopu 1 a ted in the next 10 years This wi 11 depend largely on the status and potential of the off-shore areas

Sutwi ck Area

Reports of sea otters near Sutwick Island are available since 1936 This population was far removed from other populations and is probably a remnant left after hunting ceased In 1951 Jones counted 388 between Cape Kuml iun and Cape Kunmik Most were near Sutwick Island In 1957 889 were counted Nineteen of these were between Cape Kunmik and Cape Providence indicating some northeastward expansion In 1962 949 were seen with individuals straying as far as Cape lgvak and one between there and Cape Kuliak In 1965 a stray otter was seen at Kinak Bay A major shift in the population from Sutwick into Kujulik Bay occured This may be the result of time of year and weather

In the present survey 1766 were counted between Castle Cape and Cape ~~~gtf~aip the majority were in Kujulik Bay Large numbers have moved

~~ranging as far as Cape Kubugakl i The area from Wide Bay to Puale Bay was not surveyed because of weather

The total count for the population was 1912 even though conditions were less than ideal throughout the bcunt and we feel many were missed It is possible that the increase in numbers is entirely due to reproduction assuming a 10 percent annual increase However it is difficult to compare surveys

There is still room for expansion in both directions from this population The greater movement to the northeast is probably the result of better habitat The population around Kujulik Bay is very dense and a large movement of animals may occur if competition for food becomes serious Otherwise we should expect to see continued steady expansion into adjacent areas for a number of years

to come

Augustine Island-Cape Douqlas

For some time a population has existed around the Augustine Island area at the mouth of Cook In 1 et In 1948 approxImate I y 50 sea otters vvere reported from Augustine Island Reports of sightings have been made from Shaw Island to Tuxedni Bay In 1957 Spencer counted 40 at Augustine and one at Shaw Island Lensink counted 52 on Augustine in 1959 but he considered it a poor count In 1965 Kenyon counted 18 on Augustine and 101 in the Shaw Island-Cape Douglas area In March of 1969 Loren Flag saw 62 around Augustine and in May 1969 Jim Faro counted 130 another observer saw about 30 some of which were probably not tallied by Faro

On the present survey Augustine Island could not be surveyed because of weather No otters were seen around Cape Douglas but 71 were seen in Shakun Islands and rocks near the abandoned village of Kaguyak These were probably from the Cape Douglas group

It appears that the animals move about in the area To date no complete survey of the area has been made at one time Until this is done 1ittle can be said about the population other than that several hundred probably occur there

The following table is a summary of sightings and counts made by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game These data were collected by different individuals using different types of aircraft under varying conditions of weather A strict comparison of numbers should not be made from this table

SUM

MAR

Y OF

SE

A OT

TER

SURV

EYS

(Com

pile

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tio

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5 R

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re

1951

19

51

1957

19

59

1962

19

65

1969

19

70

AU

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DOUG

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ne

Isla

nd

A

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(1

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139

197 14

1 2

53

101 62

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19

59

1962

19

65

1969

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70

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75

~-

MARINE MAMMALS SURVEY - BRISrOL BAY TO AKUTAN ISLAND June 2 1970

OBSERVER J Vania J Faro

PILOT George Tibbets Jr

AIRCRAFr Piper Azetex

middotWEATdER Partly cloudy throughout flight wind 10 miles per hour or less visibility generally excellent One bad area and that was at Akutan Island Air was turbulant there and we were unable to go completely around the island because of fog in Akutan Pass Water had slight ripple throughout flight and ocassionally there was glare but overall conditions for sightinmiddot=r animals was good

Departed King Salmon 1115 AM Returned to King Salmon 745 PM

FLIGHT PA~1 middotAfter leaving King Salmon we went over to the north side of the Alaska Peninsula and proceeded down the coast flying at 300 to 600 feet of altitude Generally we flew aboumiddott one mile off the beach At Port Heiden Moller and Cinder River we checked the outer sand bars for seals and some of the inner bars where I knew seal hauled out

We did not cover Izembeck Laroon very well Instead we flew offshore and checked Amak Island We then fueled up at Cold Bay Leaving Cold Bay we continued dmvn the north side of the peninsula (sea otter sighted) down to Cape Sharichef and crossed over to Akutan Island flying along the south side of it We were able to get around Cape Morgan and fly doNn the beach a few miles but then had to turn back becausmiddote of fo9

The flight path was then eastward to the north side of Tigalda Island At Ugamak Island we circled it completely flying at about 300 to 500 feet We completed the survey at this point and returnt~d to King Salmon generally following the route we had taken on the flight down

SEA OTTER SIGHTINGS

Seven pods of sea otter were seen south of Amak Island These were photographed and later counted from middotthe photographs Beshycause of the density in the larger pods and the fact that some animals in the smaller pods were diving when the pictures were taken some individuals may have been missed

The number counted in each pod is as follows

1071 520 357 68 36 75 30

TOTAL 2157

Several hours later the pods had drifted four or five-miles northeastward (see map) bull In addition 1 one sea otter was seen at Amak Island and sixteen near Tigalda Island

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49

)

~middotmiddot

middot 5

(

47

middot~~_

middot

38

5

6

535

3

middot

48

45

39

61

61

ti

de r

ip

58

56

54

54

56

n 1

1

73

An

41

bull6

gt ~51

~lf

gt1lt gt1lt

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6

~~ ltshy lt) lts

6

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shy

SEA OTTER COUNT BARREN ISLANDS and SHUYAK - AFOGNAK ISLANDS

June 9 1970

On June 9 1970 an aerial survey of sea otter was flown in the Barren Islands and in the AfognakShuyak Islands area A Gruman Goose was used with Schneider serving as the only observer Offshore coverage was poor Conditions of visibility were as follows

Barren Islands- 1140 AM to 1225 PM- Water calm moderate surface glare Conditions verygood Count complete

Ban Island to Pernosa Bay- 1255 to 135PM- Conditions similar to Barren Islands but surface glare worse in spots Conditions generally good

Marmot Island to Latax Rocks - 255 to 345PM- Increased ripples on surface Glare Bad Conditions fair Count around Sea Otter Island complete although some otter may have been scattered offshore Remainder of count very superficial

Area

BARREN ISLANDS East Amatuli Island West Amatuli Island Sugarloaf Island Ushagat Island Rocks south of Ushacat Sud Island Nord Island

AFOGNAK-SHUYAK ISLANDS Ban Island-Shuyak Strait Pernosa Bay Marmot Island Sea Otter Island area East side of Shuyak Jest side of Shuyak Latax Rocks-Dark Island

TOTAL

TOTAL

Number of Sea Otters

0 2 0

76 200

29 0

307

18 6 3

121 0

33 26

207

Complete or Visibility Partial Count

Very good Complete ll If II

It II

II

II IJ

Good Complete II

Fair Partial If Complete II Partial II II

II

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~ )Ilt

lt6gt iltj- cgtcgt Jgt

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9

I

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J

64

7

4

49middot

675

6

57

55

-

7

4

50

5

6

61(

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55

5

467

61

62

5

3

53

49

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5

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6

4

66

48

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-

52

62

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5

48

2

4

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46

4

3

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48

49

)

46

9~

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7

4-2

47

41

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63

5

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62

6

3

42

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4

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6

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---

)

middotshy

59

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3~-6

74

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_

20

6

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1 5

5

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34

3

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48

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3

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93

52

26

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66

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43

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6

8

60

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middot~

9

3

74

12

(+

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3

30

5

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46

)

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deg

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36

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3

40

3

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sh j

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547

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30

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I

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52

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11

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26

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3

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t-

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-18

29

3

8

4

3

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73

97

7

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19

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0

13

83

31

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0

(

46

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31

38

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29

33

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rl

24

17

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3

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26

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ll

1~~ lt 5

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37

2

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3

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9

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17

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6

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r

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77

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53

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4

3

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+

13

46

( i

49

28

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)

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+

28

tide

hp

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27

26

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3

0

36

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l-52

bull

I

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f

13

39

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6

35

25

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1

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54

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2

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36

6

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27

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3

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37~~~

32

66

72

(~I

-

1 2

2

3~

3

0

149

51

SEA OTTER COUNTS - KENAI June J970 to January 1971

Carl Divinyi flew aerial surveys of seals along the outside of the

Kenai Peninsula on a monthly basis He recorded all sea otters sighted

on these surveys The surveys did not cover the entire coast line and

the type of aircraft weather conditions and observers varied from one

survey to another Therefore the seperate counts cannot be compared

However when viewed in total they indicate the areas used by sea otters

and the relative abundance of otters in each area

The attached table presents the counts by broad areas

--

SEA

OTT

ERS

CO

UN

TED

ON

A

ER

IAL

SUR

VEYS

O

F K

EN

AI

PEN

I NS

ULP

Ju

ne

1

97

0-

Jan

uar

y

1971

8i5

8

JUN

E 5

amp 9

JU

LY

15-2

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AERIAL SURVEY Prince William Sound

April 1970

Between April 15 and 18 1970 Carl Divinyi and Julius Reynolds fletr an aerial survey of most of Prince William Sound using a Cessna 185 The survey tras primarily to locate seals however sea otter were counted The following is a summary of the survey

I departed Anchorage on April 14 via Alaska Airlines and arrived in Cordova at 600 AM tJeather conditions prevented flying so Julius and I drove the local road network looking for anything in the way of wildlife

April 15 vleather was a little better than yesterday so we decided to try surveying along the east side of the Sound up to Valdez Arm We started the survey at Bomb Point and flew the coast and offshore islands up to Galena Bay

Observations Seal - Scattered small groups with the majority of the animals being in Port Fidalgo There were no noticeable concentrations (50 on up) of seals

Sea Otter - A total of 105 otter were counted with the majority (60) being located in St Matthews Bay The remainder of the animals were concentrated between Red Head and Knmvles Head outside of Port Gravina

Sea Lion- Small scattered groups in Port Fidalgo (60-80 total)

April 16 Weather inclement - no flying

April 17 We flew Hawkins Hinchinbrook Montague and Green Islands Very few seal- many otter and two large concentrations of sea lions (See maps for numbers and locations of seals sea otter and sea lion) We also counted 5 otter around Kayak Island

April 18 Flew those islands from Montague Strait to Icy Bay and from Port Bainbridge to Knight Island Passage (See maps for numbers and locations of seals sea otter and sea lions)

Julius Reynolds continued the survey along the mainland from Knight Island Passage to Esther Island and around Knight Ingot and Eleanor Islands

The area from Esther Passage to Valdez Arm and Culross Perry Lone Naked Peak and Storey Islands were not surveyed

SEA OTTER COUNTED Prince William Sound

April 1970

Number of Area Sea Otter Date

PORT BAINBRIDGE - ICY BAY Bainbridge Island 30 418 Evans Island 14 418 Elrington Island 4 418 Latouche Island 46 418 Whale Bay - Icy Bay 39 418

CHENEGA ISLk~D - MAIN BAY Chenega Island and Dangerous Passage 33 52 Crafton Island 2 52

PORT NELLIE JUAN - ESTHER ISLAND Blackstone Bay 1 52 Culross Island NS

ESTHER PASSAGE - VALDEZ ARM NS

GALENA BAY FISH BAY 103 415

FISH BAY - GRAVINA POINT 1 415

GRAVINA POINT - CORDOVA 1 415

HAWKINS ISLfu~D 1 4l7

HINCHINBROOK ISLfu~ 99 417

EGG ISLAND 2 417

MONTAGUE ISLAND 259 417

GREEN AND LITTLE GREEN ISLANDS 102 4J7

KNIGHT ISLAND 136 52

INGOT ISLAND 6 52

ELEANOR ISLAND 3 52

SMITH ISLAND 4 52

SEAL ISLAND 0 52

APPLEGATE ROCK 1 52

PERRY ISLAND NS

(continued) SEA OTTER COUNTED Prince William Sound

April 1970

Number of Area Sea Otter Date-

LONE ISLAND NS

NAKED ISLAND NS

PEAK ISLAND NS

STOREY ISLAND NS

KAYAK - WINGHAM ISLAND 5 417

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PRE-TRANSPLANT SURVEY OF SEA OTTERS Green 1 andMontague 1 -July 171970

On July 17 1970 an aerial survey of sea otters was made in the area

around Green Island and the adjacent coast of Montague Island The purpose

of the survey was to locate concentrations of animals prior to a capturing

operation which began the next day A Dornier twin-engine arrcraft was used

with Schneider Reynolds and Somerville acting as observers The sky was

overcast almost no wind and the sea calm However heavy rain showers intershy

fered with forward visibility and the counting conditions were only fair on

the average Conditions vmiddotere especially poor in Port Chalmers and Zaikof Bay

The count began at 210pm and ended at 345 pm

The numbers and locations of otters are shown on the map In addition

16 sea otters were seen on a superficial look at Constantine Harbor on Hinchshy

in brook Island and a pod of 35 were seen three miles east of Johnstone Pt

Most of the area included in the survey was traveled repeatedly in a

skiff over the next four days Durlng this time the weather became calm and

clear for the first time in several days and the bulk of the sea otters shifted

from the coast of Montague out into the shallow areas between there and Green 1

and to Green and Channel Islands It was obvious that many otters had been missed

Fn the aerial sur~ey and that the population in the area is quite dense A

total of 48 sea otters were captured in three days of netting

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SEA OTTER SURVEY Salisbury Sound to Cape Spencer

August 23-25~ 1970

On August 23 1970 an aerial search for sea otters was made from Salisbury Sound to Torch Bay in the area north of Sitka A Helie Courier aircraft was used with Karl Schneider Alan Courtright and

middotWalt Cunningham serving as observers

The water on the outside coast was too rough for good visibility however it was calmer in the lee of rocks and islands and visibility was fair to good

On the initial survey only two otters were seen These were both in Surge Bay on Yakobi Island While returning after completing the count we located approximately 25 in the same area we saw the first two About 15 of these including at least one pup were in a single pod This illustrates how relatively large numbers of otters can be missed from the air The fact that none were seen in other areas does not mean that established populations do not exist elsewhere

On August 24 and 25 a search of the release area north of Khaz Bay was made by the skiff While flying into Kla) Bay for this seach two otters were seen near the old mine of Chichagof Two miners who have been working there reported that two otters have been there off and on since May

Rough weather and outboard problems restricted tl the search area~

and the effectiveness of the search in that area The following sightings were made

82470 I Adult West of Granite Islands 1 Adult South of Granite Islands

11 0A~dgtzl~4JVflup) middot1 In rocks SE of ranite Islands 82570 I Adult -n Southwest of Gran~te Islands

These sightings represent from four to seven animals

Reports over the last year indicate that sea otters regularly move in and out of Kla~ Bay and occasionally as far in as Sisters Lake bull

The fact that the count in the Khaz Bay area Has less than that made in 1969 does not mean much The animals appeared to be scattered during the present count the search did not include all of the nearby sea otter habitat and survey conditions were less than ideal

The population in Surge Bay appears to be separate and is probably well established Two otters were reported in the same location in January 1966 after only 23 otters had been released near Khaz Bay In 1969 two were seen in the same spot from the ait

While no estimate of the population of sea otters in the area can be made from the available information is evident that sea etters can be found along the entire west coast of Chichagof Island and that concentrations occur near Black Island and the Granite Islands north of Khaz Bay and in Surge Bay on Yakobi Island

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Harvest

In May of 1970 a sea otter harvest was conducted on Tanaga Island the middotDelarof Islands and Amchitka Island The numbers to come from each island

and the distribution of the kill around each island were planned in advance Conservative estimates of the population tvere made and the total harvest numbers were based on these estimates The harvest level for Tanaga and the Delarof Islands was set at 15 percent assuming that the next harvest in these areas would be three years later so the rate of harvest would be 5 percent per year Amchitkas harvest level was set at 10 percent assuming an annual harvest and a harvest rate of 10 percent per year

The harvest plan was based on information collected on a June 1968 skiff survey for Tanaga on the 1965 USFWS aerial survey and the 1969 ADFampG aerial

survey for the Delarofs and on 1968 helicopter survey by the USFWS for Amchitka There is good evidence that all the population estimates tvere low especially for the Delarofs

Hunting was done from avo skiffs with two men each Both men shot when possible A 117-foot vessel was used for skinning and living quarters Table 1 shows the schedule of the hunt with the daily harvest numbers

Table 2 presents the projected and actual harvest numbers by area Table 3 presents the numbers killed pelts saved specimen and pelt numbers used and the sex ratio of the kill for each area Figures 1 2 and 3 show the actual numbers and locations in more detail tveather was the main factor causing deviations from the original harvest plan Possible male areas were identified on the 1968 skiff survey on Tanaga An attempt was made to concentrate more pressure on these areas to achieve a more balanced sex ratio This effort is reflected in the sex ratio of the kill A high degree of sexual segregation occurs at this time of year and without specific pressure on male areas we could expect 85 percent females to be taken as occurred on Amchitka With the effort the female percentage was reduced to 65 percent on Tanaga In the Delarofs a male area which had not been identified previously was hit and the effect was much the same

Transplants

Two transplant operations took place in 1970 The first was done in the same manner as the 1968 and 1969 transplants A total of 86 otters tvere captured at Amchitka One aircraft load was shipped out Twenty-nine were released in Oregon and 30 in vashington The animals were held in floating pens at the release sites for several days to recover from the trip Survival appeared to be excellent

The second operation took place in Prince William Sound The Canadian research vessel G B Reed came to the Green Island-Port Chalmers area with tanks built on deck Forty-six sea otters were captured Approximately 44 were alive when the vessel left Prince William Sound however only 14 survived to be released off Vancouver Island

Table 1

April 20

middotMay 1

May 2

May 3

May 4

May 5

May 6

May 7

May 8

May 9

May 10

May 11

May 12

May 13

Schedule of the 1970 harvest

1030 pm depart Homer

Midnight arrive Adak MV Active

Refuel and prepare boat

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Complete skinning pack hides etc in pm

Delarofs

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88

131

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Table 2 Projected and actual harvest numbers May 1970

Projected Actual Tana~a Island Harvest Harvest

Bumpy Point - Hot Springs Bay 200 191

Hot Springs Bay - Twin Bays 50 50

Twin Bays - Cape Sasmik 50 ) ) 199

Cape Sasmik - Tower 120 )

Tower Tanaga Bay 180 166

TOTAL 600 606

Delarof Islands

Gareloi 20-25 0

Kavalga Ogliuga and Skagul 75-100 125

Ulak and Amatignak 35-55 19

TOTAL 150 144

Amchitka Island saved for

USFWS Counting Units 1 amp 2 100 transplant

3 30 82

4 50 36

5 120 87

TOTAL 300 205

Table 3 Details of sea otters harvested in May 1970

Tanaga Island

Total kill 606

Number of pelts saved 598

Number of small pup pelts discarded 8

Specimen numbers SO 70-4 to SO 70-609 Pelt numbers 3198-3769 and 3771-3796

(Seal 3770 void - damaged)

Approximate sex ratio - 220 males 386 females or approximately 64 percent females

Delarof Islands

Total kill 144

Number of pelts saved 138

Number of pups discarded 6

Specimen numbers SO 70-610 to SO 70-753 Pelt numbers 3797 to 3934

Sex ratio- 44 males 100 females or approximately 69 percent females

Amchitka Island

Total kill 205

Number of pelts saved 194

Number of pups discarded 11

Specimen numbers SO 70-754 to SO 70-958 Pelt numbers 3935 to 4128

Sex ratio- 27 males 178 females or approximately 867 percent females

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A combination of bad weather a lack of time to let the otters recover from their capture and problems with the holding facilities probably caused the high rate of mortality

Summary of Harvests and Transplants

Table 4 summarizes the sea otters removed from Alaskan populations in the middot last 20 years An attempt has been made to remove 300 otters from Amchitka each year since 196 7 Originally this number was set as 10 percent of the population which was conservatively estimated to toal 3000 Recent USFtfS helicopter counts of up to 3927 show that this estimate definitely tvas low In the past the removal of animals was from the southeas tern half of the island The 1970 harvest was purposely concentrated toward the northwestern end in order to spread out the pressure and to learn something about the population in that area

The numbers of animals released in all transplant attempts by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game are presented in Table 5

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30

SEXUAL SEGREGATION IN SEA OTTER POPULATIONS

by Karl Schneider March 1971

Workers have recognized for some time that sea otters tend to segregate by sex Lensink (1962) described this segregation around the southeastern end of Amchitka Island and identified three male areasn and three female areas He speculated that females used areas of more favorable habitat and that younger males were excluded by territorial males scattered throughout the female areas The implication is that male areas contain younger or at least nonterritorial males

Marakov (1965) mentions sexual segregation around Medny Island in the USSRs Commander Islands

Kenyon (1969) gives a more complete description of the sexual segregation around the southeastern end of Amchitka and supports it with quantitative data from harvested animals

Both Lens ink (1962) and Kenyon (1969) w_ere primarily describing hauling grounds used by different sexes While Kenyons harvested animals included otters near shore neither study provided much information on the use of off-shore areas

A knmmiddotlledge of the degree of sexual segregation and the location of male and female areas is important to the management of sea otter populations Harvests must be regulated to avoid putting too much pressure on one segment of the population Then capturing animals for transplants it may be necessary to set nets in specific areas to obtain the desired sex ratio case of a localized kill of otters such as when oil spills occur it is important to know the distribution of sexes to evaluate the importance of kill to the population

In

the

By the same token much can be learned about the distribution of sexes through harvest and capture operations The area from which each sea otter was taken during the harvests of 1967 1968 and 1970 was recorded Because a single hunting party might kill 30 otters over up to 10 miles of coast in a few hours it wasnt possible to record the precise locations Therefore the coast was broken up into areas and the numbers killed in each area ltJere totaled

Figs 1 - 5 show the locations of the areas letters correspond to those in Tables 1 - 4 which present the sex and age composition of animals harvested in each area The ages of the 1968 animals 1vere based on cementum deposition and reproductive condition The other samples were broken down using body size In general all males under 25 lb females under 20 lb and both sexes shorter than 100 em were considered dependent pups Females under 35 lb and 120 em and males under lb and 130 em were considered subadults These criteria probably underestimate the age of some animals Cementum deposition information for the 1967 and 1970 samples is not available yet

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Sexual Segregation - 2 - March 7 1971

Information from the 1967 Amchitka harvest and the 1968 1969 and 1970 Amchitka transplant capturing operations is not comparable and is not presented here However knmv-ledge gained from these operations has contributed to our understanding of sea otter distribution and this knowledge will be used in the following discussion In general this data confirms Kenyons (1969) observations for the Amchitka area although these summer and fall collections indicate a higher percentage of males in female areas than Kenyon reported from his winter and spring samples

Kenyon (1969) states that female areas are more numerous and less discrete than male areas He identified three male hauling grounds and 13 female hauling grounds on the southeastern end of Amchitka Island Recent studies involving netting and collecting animals off~hore indicate that male areas remain discrete off shore Usually these areas are off major points of land and pods of males often form in kelp beds directly off shore from the hauling ground

Female areas on the other hand are not discrete at all Any area that is not a male area can be considered a female area Some areas may be more attractive to females with pups or certain areas may be more favorable for hauling out but more females than males vill be found almost everyvthere except in the discrete male areas Therefore the main problem is to locate the male areas in et population He have no evidence of any shift in the location of male areas over a period of time so once an area is identified the information should remain useful for management purposes for many years Lensink (1962) and Kenyon (1969) correctly identified all of the male areas on southeastern Amchitka Extensive harves and netting have not turned up any new areas

Identification of Hale Areas by Pup Counts

In June 1968 a survey of most of Tanaga and Kanaga Islands was made by skiff to gather information to be used in planning harvests from those islands During this survey females with pups were counted separately from single animals No special effort was put into identifying pups fuen a female obviously had a pup it vas recorded Large pups often were separate from the females and some otters were seen only briefly in vaves kelp or at a distance Therefore many pups probably were missed and the counts should be considered minimal 6 and 7 present the counta by area A similar count was attempted by helicopter around Adak but for various reasons the results are not considered useable

From this skiff survey a number of areas were judged to be possible male areas Usually these were points or more exposed aTeas vhere relatively large numbers of single animals but no pups were seen

Shoal Point Kanaga Pass north of Eddy Rock and possibly Naga Point and Cape Tusik were suspected male areas on Kanaga Island Cape Sudak and Cape Amagalik were considered male areas on Tanaga Island and the area around Hazard Point and Lltnnoy Rock in Kanaga Pass was believed to blend with the Eddy Rock area

The best way to confirm the presence of male areas is to collect animals from these areas Harvests were conducted on Kanaga in October 1968 and on

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7A

Sexual Segregation - 3 - March 1 1971

Tanaga in May 1970 The sex ratios of the animals harvested are compared with the skiff survey pup counts in Table 5 The harvest areas are fairly broad while male areas are usually only a few hundred yards wide As a result a harvest area may include both a male area and portions of female areas to either side

The hunting pressure is seldom even over an area Often a good portion of the kill in an area would come from one group of rocks Therefore the information in Table 5 must be tempered with some knmrledge of how the hunting pressure was distributed

Kanaga Island

Shoal Point - While a few more females than males were taken in this area the percentage of males is considerably higher than one would expect in a female area Because weather vas less than ideal much of the hunting pressure ~vas directed to areas other than Shoal Point Most of the females probably came from these areas Therefore Shoal Point is considered to be a male area$

Naga Poin~- There is no strong evidence that this is a male area While no pups were recorded on the survey the total number of otters counted was not high Fairly extesive hunting of the area under good conditions yielded a sex ratio that would be typical of a female area Therefore it is assumed that there is no male area near Naga Point

Cape Tusik - Cape Tusik was suspected as a male area more because of the nature of the area than arry counts The count included areas to both sides of the Cape which could contain females Very little of the hunting was done at the Cape Therefore judgement on this area will have to wait until more information is available Extensive hunting indicates that no male areas exist east of Cape Tusik to Cape Chlanak

Kanaga Pass - From the count it appears that a male area exists in the Pass however the harvest area extended to Hive Rock and included a large portion of female area (Fig 6) The area is too broad as is shown in Table 5 gtvhere the number of pups counted for the whole area is relatively high The results of the harvest confirm the conclusions drawn from the count While the sex ratio for the entire area is roughly equal most of the males came from Kanaga Pass Some females without pups came from that general area but the majority of the females came from the area northeast of the Pass

The Tanaga harvest showed the male areas more distinctly for several reasons The total harvest was higher the entire count area was covered in the harvest and the hunting pressure was recorded more precisely Also as will be shown later sexual segregation is more pronounced in the spring

Cape Sudak - Most of the hunting pressure was concentrated on the tip of the Cape Some pressure was exerted on the area back to~mrd Pendant Point which is a female area as shmvn by the count The high percentage of males harvested from this area clearly confirms that the tip of the Cape is a male area

Tab

le

5

Com

pari

son

of

Pup

s C

ou

nte

d w

ith

S

ex R

atio

o

f H

arv

est

Jun

e

1968

A

rea

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rvey

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ber

19

68 H

arv

est

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ps

10

0

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ps

10

0

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ult

s T

ota

l KA

NAGA

IS

LAN

D

Ind

ep

Ott

er

Ind

ep

Ott

er

M

M

F

Rou

nd

Hea

d-S

ho

al P

oin

t 0

85

3

45

6

55

4

71

5

29

Nag

a P

oin

t-K

anag

a B

ay

66

1

64

middot

18

5

81

5

20

5

79

5

Cap

e C

hla

nak

-Cap

e T

usi

k

51

1

93

2

40

7

60

3

53

6

47

Edd

y R

ock

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e R

ock

61

3

4

35

7

64

3

47

1

52

9

Jun

e

1968

A

rea

Su

rvey

H

ay

1970

Har

ves

t

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ps

10

0

Pu

ps

10

0

Ad

ult

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l TA

NA

GA

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LAN

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Ind

ep

Ott

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F

M

F

Cap

e S

ud

ak-P

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ant

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4

60

6

39

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4

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6

Bar

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run

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59

3

3

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ays

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4

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77

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60

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th B

ay

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3

92

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14

3

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7

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uth

Bay

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ock

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8

31

6

7

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91

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09

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73

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0

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25

4

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em R

ock

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lak

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int

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35

6

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6

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int-

SE

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igh

t 6

9

82

1

32

8

68

1

51

8

49

Sexual Segregation - 4 - March~ 1971

Annoy RockHazard Point - The harvest did not confirm this area as a male area However the weather Vas marginal and this area was too rough to hunt Therefore the otters taken in the harvest ~vere from either side of the Point and almost none were taken from the portion that is suspected to be a male area Actually Kanaga Pass is narrow and shallmv Otters feed in all parts of the Pass when weather and tide rips permit This could be considered a single male a~ea that serves both islands

Cape Amagalik - Two harvest areas overlap this area Lash Bay-Inferno Reef and Harem Rock-Kulak Point The kill from both areas strongly reflects the presence of a male area Nost of the hunting pressure was concentrated on Inferno and this is probably the main male area

As shown above vle 1vere able to identify four major male areas from a relatively superficial pup count No male areas were located during the harvests that had not already been identified from the count ~mile more detailed observations including field identification of males would be desirable the pup counting technique appears to be an acceptable method

Identification of

Because male areas are usually on v1ell exposed points it is possible to pick likely areas from a chart and then confirm these areas by collecting animals This method avoids the expense time and danger involved in surface surveys of remote areas Actual collection of animals also provides the most concrete proof of the nature of an area

The follmving is a description of the sex composition of areas for which no skiff count data was available

Delarof Islands - No attempt 1vas made to predict the location of male areas in the Delarof Islands This is a difficult area to work in Ogliuga Skagul and a portion of Ulak Island v1ere hunted on Nay 9 1970 The hunting effort in the afternoon was concentrated around Tag Island and the rocks south of Skagul Island While the entire kill is lumped together most of the males bullJere taken in the afternoon The percentage of males taken for the day is higher than would be expected if only female areas were hunted Most likely there is a male area near Tag Island

Adak Island - Portions of Adak gtvere hunted in 196 7 and 1968 No attempt was made to identify male areas before or during either harvest The harvest figures indicate that no male areas have been hunted The Bay of Islands is a classical female area The results of extensive hunting in the Bay and around Eddy Island indicate that there is definitely no male area there The percentage of males taken between Mid Point and Blind Point vms higher than might normally be expected in a female area however the scarcity of subadult males indicates that no male area exists within this area A large congregation of otters is often seen near Finger Shoals This could be a male area but Ye have no information from there If this is a male area a few stray males from there could account for the slightly higher number of males in the kill

Sexual Segregation - 5 - March 1971

Amchitka Island - While much work has been done on Amchitka Island almost all of the effort has been concentrated on the southeastern end from Crmvn Reefer Point to a fegtv miles northvest of Rifle Range Point During the 1970 harvest an attempt was made to distribute the harvest over previously unstudied areas Two areas were suspected to be male areas These were the rocks off Chitka Point and either Bird Rock or Aleut Point Heather prevented us from hunting the area around Bird Rock and Aleut Point

The sex ratio of the kill does not indicate a male area near Chitka Point however heavy waves prevented hunting off the point Therefore a male area could be there but because no animals were killed there the kill figures would not reflect it

Composition of Female Areas

Kenyon (1969) found that 93 percent of the adult otters and 63 percent of the juvenile otters in female areas were females The May samples from the 1970 harvest agree with this The mean percent of adult females in female areas was 93 percent The mean percent of subadults not including pups was 73 percent Hmmiddot1ever in the September and October samples only 77 percent of the a-dults in female areas were female About 86 percent of the subadults were female

The seasonal difference in the sex ratio of adults is consistant for all samples These seasonal changes are most likely due to adult males moving into female areas to look for estrous females During late winter and spring breeding activity is at the lmmiddot1est point of the year and fev1er females are entering estrus In fall the number of estrous animals is at its peak and presumably more sexually mature males move into the female areas to breed It is interesting to note that there were about three times as many males in female areas in fall as in winter and spring Information from female reproductive tracts indicates that approximately three times as many females would be in estrus at any given time in fall It appears that the number of adult males in a female area is directly proportional to the number of estrous females From the data collected there appear to be 15 to 20 males for each female -vlith enlarged follicles at any given time hmmiddotrever hunter bias might influence this figure

Ages have been estimated for otters harvested in 1968 on the basis of cementum layering These age data indicate that virtually all of the males in discrete female areas are either pups (or very young subadults) or are 7 years old or older No males between the ages of 2 and 6 Here found This strongly implies some form of territorial behavior among actively breeding males Fighting among males has not been observed ofteci hmvever a mature male at the Tacoma Zoo became intolerant of a subadult male in the presence of females They frequently 11 fought 1 although neither seemed to attempt to injure the other Finally the young male had to be removed Under imiddotJild conditions the young male might have moved out of the area without repeated physical encounters

Kenyon (1969) described the breeding behavior of sea otters His description indicated that males patrolled an area and two to four males may pass a given point during a 3 or 4 hour period Vandevere (1970) observed some males

Sexual Segregation - 6 - March~ 1971

apparently defending a territory in California but from his observations t

it appeared that successful breeding males were more mobile Perhaps established breeding males are more tolerant of each other More likely they maintain a separation from each other which limits the number in an area In this way a number of males might be patrolling the sa~e area rather than each establishing a geographical territory This might be advantageous where female concentrations shift from place to place as weather conditions change

Why does the number of adult males in female areas decline when the number of estrous females declines The limiting factor may be competition for estrous females rather than territory size With fewer receptive females competition lvould be greater causing some animals to leave the area Another possible reason may be that males may have a seasonal fluctuation in fertility Lensink (1962) found that all adult males produced sperm at all seasons Limited studies since then support this however it is possible that most of these males were collected in or near female areas Also while a male may produce sperm at all seasons there may be less production at certain times of year

A contributing factor may be that males are capable of feeding in more exposed areas than pregnant females or females with pups With a reduced attraction to the female areas either middotbecause of fewer receptive females or a reduced fertility in the male the male may find it easier to obtain food away from the crowded female areas where competition for food is greatest

The fact that the number of males does appear proportional to the number of estrous females indicates that competition for these females may play an important part in regulating the number of males in female areas

The numbers of subadults in the samples is relatively low so fluctuations in sex ratio may be due to sampling errors However the greater number of subadult males found in female areas in winter and spring may be a true increase made possible by the reduction in the number of breeding males A subadul t male would have fewer encounters -vli th breeding males

There are areas within what would normally be considered female areas where subadult males tend to congregate particularly during the winter These are usually areas v7ith a ma-r-ginal food supply that are not heavily used by females Because there are fewmature females in these areas there are also very few breeding males The subadult males using these areas seem to be transients that find less pressure from breeding males there These areas are used less in summer probably because calrrer weather allows feeding farther off-shore reducing competition for food in the male areas

Examples of areas within female areas that are used by subadult males are Constantine Harbor on Amchitka and possibly Finger Bay on Adak

within Female

In the course of the harvest on Tanags in tIay of 1970 a large number of pregnant females tvith large fetuses were collected at one time Most of

Sexual Segregation - 7 - March 1971

these came from Tidgituk Island This raised the possibility that females may segregate according to reproductive condition Table 6 presents the numbers of females in each stage of the reproductive cycle by area on Tanaga Island More pregnant animals were collected in the South Bay area and within that area most of the females with fetuses weighing over 100 g were collected near Tidgiktuk Island This concentration was evident in the pup counts made in June 1968 when a very high percentage of pups was observed in the same area (Table 5) Host females ~vith large fetuses in May would pup by June

The data do not reveal any other areas of this type on Tanaga Harakov (1965) mentioned a bay on Hedny Island that tvas preferred by pregnant females and females with pups Obviously all females do not go to a specific area to pup but a female vi th a large fetus or small pup probably has more difficulty in exposed areas and seeks more protected areas This tendency can be seen by the casual observer Single animals are more likely to venture off-shore and remain in rougher waters In some areas such as Crown Reefer Point on Amchitka there is a male area near the tip of a point Males congregate directly off-shore However to either side of these male congregations there are females that are not accompanied by a pup and probab do not have large fetuses In this way we may find segregation in a single kelp bed off a point If we set a net near the outer margin of the kelp we will catch almost all males Ho~Vever if we set a net to the side of the bed and closer tmiddoto shore we will catch mostly single females Nets set in a nearby bay will catch more females with pups

Table 6 shoHs that an unusually high number of resorptions were found between Cape Sudak and THin Bays This is probably a reflection of poor physical condition of the otters in that area due to food Other data indicate a continuous decline in body condition from the east side of Adak to Kanaga Pass There is some evidence that condition improves west of Kanaga Pass The animals in the Kanaga Pass vicinity are probably in poorer condition than any of the other populations sampled Therefore the high incidence of resorptions in this area should not be considered an example of segregation within the population

ition of Male Areas

As mentioned earlier the harvest areas ivere broader than any male area Harvest areas containing male areas also contained portions of female areas Our identification of the location at rhich each animal vas collected is not precise enough to determine the exact composition of these rnale areas Kenyons (1969) data from winter and early spring harvests are more precise His data indicate that at that time of year 98 percent of the adults and 80 percent of the ju~veniles using male areas were males Of 128 males in male areas 100 gtvere adults and 28 were juveniles

Experience from netting in the summer and harvesting in fall indicates that the percentage of males remains very high all year While adult females may be very close to male areas it appears unlikely that a significant number regularly use these areas at any time year nile adult males enter female areas regularly and in predictable numbers for a specific purpose females probably ~nter male areas only occasionally in stray movements

le 6

R

epro

du

ctiv

e S

tag

e o

f S

exu

ally

Mat

ure

S

ea O

tters

Har

ves

ted

on

Tan

aga

Isla

nd

-by

Are

a -

May

1

97

0

Un

imp

lan

ted

Im

pla

nte

d

Pre

gn

ant

Pre

gp

ant

Fet

us

Wei

gh

t C

lass

A

rea

No

np

reg

nan

t N

o

No

1

2 3

4 5

Res

orp

tio

n

To

tal

A

Su

dak

-Pen

dan

t P

oin

t 12

7

26

8 30

0

2 0

3 2

2 27

( B

arn

es

Po

int-

Ho

t S

pri

ng

s B

ay

12

7 24

10

36

3

0 1

4 2

2 29

B

( (

Bar

nes

P

oin

t-T

run

k P

oin

t 1

1

6 25

7

29

0 3

1 3

0 2

24

( T

run

k P

oin

t-T

win

Bay

s 7

9 39

7

30

2 1

1 1

2 2

23

c ( (

Haz

ard

P

oin

t-T

win

Bay

s 4

3 27

4

36

0 0

1 1

2 11

( T

win

B

ays-

Her

d

Roc

k 1

2

8 28

9

31

1 1

1 Lf

2

29

D

( ( T

win

B

ays-

So

uth

Bay

2

9 45

9

45

3 0

1 1

Lf

1 20

( S

ou

th B

ay-L

ash

B

ay

6 9

29

16

51

1 0

0 9

6 1

31

E

( ((

So

uth

Bay

-Har

em R

ock

17

14

31

14

31

2 0

1 5

6 45

(

F ( (

L

ash

B

ay-I

nfe

rno

Ree

f 4

1 9

6 55

2

1 1

0 2

11

( G

(

Har

em R

ock

-Ku

lak

Po

int

12

6 29

3

14

1 0

1 1

0 21

H

Ku

lak

Po

int-

SE

B

igh

t 13

9

28

10

31

2 0

1 2

5 32

Bra

cket

s in

dic

ate

o

ver

lap

pin

g are

as

Sexual Segregation - 8 - March 1971

Juveniles of both sexes probably move more randomly than adults lihile definite sexual segregation exists among younger animals it is not as pronounced as in adults Because subadult males are tolerated less by breeding males they are more strictly confined to small areas than females of the same age This has been demonstrated in fall harvests where extensive hunting over a large area will turn up only an occasional subadult male Then ~v-hen the hunters move to a definite male area large numbers of subadult males are taken in a very short time and in a very small area

While Kenyons data indicate that 22 percent the males in male areas are subadults our experience indicates that this percentage may be quite a bit higher at least in fall Such a seasonal change in the age composition of males sounds reasonable We have demonstrated that adult males move into female areas in greater numbers in the fall perhaps tripling their numbers in these areas At the same time fewer subadult males are found in the female areas Presumably the adult males are coming from male areas and the subadults driven from the female areas move to the same male areas The result vmuld be a higher percentage of young males in male areas in fall than in spring

The total number of males of all ages in female areas doubles in fall About 13 percent of all animals taken in these areas in spring were ~ales while 25 percent ta-ken in fall were males This suggests that the total number of males in male areas declines in fall There are more adults leaving than subadults entering the area

Sex Ratio

With the sexes segregating and with the degree segregation changing seasonally and betmiddotween age groups it is extremely difficult to get a completely random harvest As a result we have not been able to determine the sex ratio of any population as a vhole Hith female areas being larger and in more protected areas where hunting is easier there is a strong tendency to take more females than males particularly in sp-ring ~Je have shown that through a concentrated effort the percentage of males harvested can be increased however even then the lowest percentage of females taken in recent harvests ivas 62 percent on Kanaga Island in October 1968 Similar results occur in capturing operations for transplants

On all islands studied there are fewer male areas thltm female areas The male areas are very small usually only a few hundred yards wide The male feeding areas off male areas are also very narrow although they may extend farther off-shore In short that portion of the total available sea otter habitat included in male areas is very smalL

All evidence indicates that there are more females in the population than males While the percentage of is probably greater than that indicated by most harvest statistics it appears that at least 60 perc2nt of the sea otters in the populations studied are females

There are probably several factors contributing to this uneven sex ratio First our reproductive data indicate that 56 percent of the pups are females at birth Secondly Kenyon (1969) has found a higher mortality rate among

Sexual Segregation - 9 - March 1971

newly weaned males There is also some evidence that males might not live as long as females These factors could easily account for the preponderance of females in the population

It is possible that the higher rate of mortality in juvenile males is in part due to the breeding behavior of adult males and the resulting sexual segregation Juvenile males tend to be restricted to male areas or areas of marginal feeding habitat where there is little competition from breeding males During the ltvinter the feeding activity of young males would tend to be confined to small areas close to shore in male areas Therefore they may be subjected to greater competition for food than females of the same age which have a broader area in ltvhich to feed Kenyon (1969) found that subadult males vere less hardy in captivity than subadult females Therefore we can not adequately evaluate the influence of segregation on mortality

The sex ratio probably varies from one population to the next Very little juvenile mortality occurs in expanding populations eliminating that source of sexual selection However there is some evidence that males are more numerous among the first animals to repopulate a new area While a high percentage of males might be found on a newly repopulated island the loss of these males tvould alter the sex ratio of the parent population even though juvenile mortality might not yet be a significant factor

If the nThuber of breeding males is regulated by the number of estrous females an even sex ratio might produce a surplus of sexually mature males This surplus would not be beneficial to the population and could be detrimental This would permit the situation of an unbalanced sex ratio to evolve

The segregation of sexes and ages and the composition of sea otter populations is complex Unless we can understand the situation we cannot fully evaluate the effects of mortality or habitat changes

REPRODUCTION IN THE FElIALE SEA OTTER

by Karl Schneider Narch 1971

A total of 1358 sea otter reproductive tracts collected between 1967 and 1970 have been examined Ovaries were sliced with a razor blade and examined macroscopically Uteri Here macroscopically examined both internally and Each tract vas classified as to its stage in the reproductive cycle

The follovJing criteria were used in classification

Nulliparou~- Horns of uterus small thin and smooth no corpora lutea or corpora albicantia although there is occasionally evidence of a past ovulation but no evidence of implantation

Nultiparous - Uterus thicker corpora lutea andor corpora albicantia present Includes primiparous animals (these are not readily separated from multiparous animals)

Anestrus - Largest follicle under 3 5 rnm no corpus luteum

Proestrus - Follicles over 35 mm

Es - Follicles approximately 10 rrm

Implan~elt_ Pregnant_ - Visible s~velling in uterine horn usually with fetal membranes embryo or fetus visible Corpus luteum usually over 10 mm

- Uterine horn sti11 noticeab enlarged fresh roughshy~=-=--=~

al scar newly formed corpus albicans with some luteal tissue remaining

The appearance of the uterus ~ras used to confirm the classification The thickness arrd of the horns thickness and consistency of the uterine walls coloration of the of the horns and condition of the rugae change with each stage In most cases it is possible to guess the condition of the tract by a superficial examination of its exterior This appearance was used only to confirm what was indicated by structures in the ovary hmvever

Table 1 smnmarizes the reproductive condition of the sexually mature females in the that are enough for comparison throughout the year In the following discussion reported by Sinha et (1966) and Kenyon (1969) are included 7here possible The January and Harch are from these studies

Tab

le

l Re

p iv

e co

nd

itio

n o

f se

xual

ly m

atur

e se

a o

tters

_

IF

etus

Wei

--------------+

__

_

ln~a

ctive

ov

arie

s 1

Act

ive ovari~

ht C

Jas

s

jmiddot

I P

ost

-I P

roes

tru

s U

nim

pl

lmpl

D

ates

L

ocat

ioQ

1A

nest

rus

Part

um

Res

orpt

ion

l +

E

stru

s P

reo

Pr

eQ

2 3

lJ

S

~

4-8

1970

T

anag

a 1

51

41

0

10

104

17

8 10

33

30

4 (1

68)

(1

35)

(3

3)

(3 3

) (2

89)

(34~

(5

6)

(27

) (3

3)

(11

4)(

10

9)

May

9

19

70

Del

arof

Is

18

14

1

4 13

27

10

0

3 6

8 77

4

) (1

82)

( l

3)

(5

(16

9)

( o

) (1

30

) (3

9)

(78

)(1

04

)

10-1

219

70 A

mch

ltka

I

34

18

3 8

27

48

5 6

2 23

12

13

8

( 0

(24

6)

(13

0)

(22

) (5

8)

(19

6)

(34

8)

(36

) (4

3)

(14

) (1

67)

(8

7)

~lune

23

middotmiddot ~middot~~middot~Amchitka

1

17

2 1

2 4

18

1 2

3 5

7 44

A

ugus

t 13

196

8 (3

86)

(4

5)

(23

) (4

5)

(90

) (4

09)

(2

3)

(45

) (6

8)

(11

3)(

15

8)

July

6-

~dgtAmchitka

1

17

1 0

4 14

10

0

1 1

2 6

46

Aug

ust

819

69

(36

9)

( 2

0 2

) (8

6)

(30

4)

(21

9)

(22

) (2

2)

(43

)(1

32

)

Sep

t 1

0-17

A

dak

72

5 0

16

31

40

6 6

8 9

11

164

1967

(4

39

) ( 3

1)

(98

) ( 1

89)

(

4)

(3 7

) (3

7)

(49

) (5

5)

(6

7)

Sep

t

25

-A

mch

itk

a l

55

6

0 18

19

17

4

0 0

0 3

115

OcL

6

19

67

(4

7~8)

(5

2)

(15

7)

(16

5)

(14

8)

(35

) (8

7)

(26

)

Oct

o 1

2-1

5

Kan

aga

I

58

6 1

13

31

31

4 4

7 11

5

140

1968

(4

13)

(4

3)

(o 7

) (9

3)

(22

2)

(22

2)

(29

) (2

9)

(50

) (7

8)

(36

)

Oct

18

-20

A

dak

l

46

6 0

1 24

13

2

3 0

4 4

100

1968

(L

16 0

) ( 6

0)

(11

0)

(24

0)

(13

0)

(20

) (3

0)

0)

(40

)

Num

bers

in

re

nth

esis

are

pe

rcen

t se

xu

ally

mat

ure

fem

ales

F

etus

wei

ght

clas

s 1

=

0-l

g

2 =

1-l

Og

3

= 1

0-lO

Og

4

= 1

00-lO

OO

g

5 =

Tra

nsp

lan

t m

ort

alit

ies

(all

o

ther

sam

ples

fr

om

har

ves

ts)

Reproduction - 2 - March 1971

There may be some problems associated with comparing samples taken from different islands and in different years but allowing for sampling errors the data for the most part fit a definite pattern indicating that the annual cycle remains fairly constant The one outstanding exception is the number of pregnant animals in the t1m summer samples In 1968 there were many more implanted pregnancies than unimplanted pregnancies and in 1969 the situation was reversed hmvever the total number pregnant was almost identical These are the t s being relatively small collected over a longer of time and consisting of transplant mortalities An average of the two samples fits the pattern established by the other samples Relatively large samples are necessary because all stages of reproduction are found at all times of the year The period from November to early January is not represented hmmiddotJever things are changing so rapidly during that period that any information from that time may be confusing unless a large sample was collected in a very short time

The information in Table 1 is shmvn graphically vJith Kenyons (1969) winter information in Figs 1 and 2 It has often been demonstrated that sea otters breed and pup at all times of the year A number of observers have noticed that roore pups are born in the spring aJd sumrrter than at other times and there have been conflicting reports about breeding times Kenyon (1969) was the first to demonstrate seasonal changes in pregnancy rates but only his January and Yarch sallples gtvere large enough to be at all reliable

Figs 1 and 2 demonstrate that there are cons le secsonal in the numbers of animals in each s reproductive cycle There is a definite period of increased bree activity and a definite period of increased pupping Hmmiddotlever ~ some anii1als are in each s at all times of the year and these periods of increased breeding and pupping do not have distinct boundaries

Each curve is influenced by two factors so it is difficult to isolate the magnitude of any single factoc For example the of implanted pregnancies vill increase ss blastocysts and decrease as births occur The birth rate may be increas but if the number of implantations increases at a greater rate the implanted curve will still rise The animal remains in the anestrus implanted pregnant~ and unimplanted pregnant categories for a relatively long time and there may be considerable carry-over from one sample to the next Because the animal a relatively short time in the proestrus-estrus and post partum categories there is little if any carry-middotover from one sample to the next and the changes betgtmiddotTeen are more likely to be absolute changes The mean fetus weight class prob does not mean much because it does not take the actual number of animals in each class into account ~hen there are many implanted pregnancies ard the mean lmiddotTeigh t class is lovJ there actually may be more Class 5 fetuses in the population than vhen there are fetv irr1planted pregnancies and the mean middotleight class is high Therefore the actual percentages of all sexually mature females middotlith Class 1 ald Class 5 fetuses have been plotted in 3 These curves represent the actual number of fetuses in the population Because tha sample sizes become quite small vhen the fetuses are divi(12d into classes and the surruner sample is believ2d

2

~ _s ~

~

c U)

~-1

r

~J

lt

c])

Reproduction - 3 - Narch 1971

to be biased tvo curves for each class are shmvn The solid curves approximate the data while the broken curves represent subjective adjustments to correct for sampling errors

cussion of the Annual

The number of animals entering estrus rises in the fall At this time the pregnancy rate is at its lowest point and a high percentage of anestrous (nonpregnant) animals are in the population Breeding activity increases and at this time the number of sexually mature males in 1female areas increases

The number of unimplanted pregnancies begins to rise sharply in October and November At the smne time the nlli~ber of anestrous animals drops as these animals enter estrus breed and become pregnant Thj_s is the peak breeding season

The birth rate is at its louest this period as is shovm by the lmv number of post-partum a1imals and the lovr number of large fetuses

By late November and December the pregnant rate is increas even though the birth rate is slightly This is due to ne7

blastocysts implanting as is shown the increase in Class 1 fetuses

By January the period of t breedLng activity has declined and the number of unimplanted animals ceaches a peak as the number of blastocysts implanting equals the number of The lanted pregnancy rate then begins to declLne as fevJer animals breed and more implant The greater rate of antation is reflected in a rise in the Class 1 fetuses as well as an overall increas in the er of lanted pregnancies However the rate of implantation is partially obscured by an increased birth rate removing animals from the total of lanted pregnancies This increase in the birth rate is shmm in the rise in post-partum females and a rise in Class 5 fetuses

This condition of a low rate of breeding high rate of implantation and increasing rate of birth lasts from February to Hay

At this point He come to the least reliable data The sumner are smaller gtJere collected over a period of seven weeks and were from animals that died as the result of capture and handling

Therefore tvhile the data shaH that the post-partum rate decreases sh indicating a reduction in the birth rate~ the number of Class 5 fetuses

a peak indicating a very high birth rate The true birth rate is probably some1middothere in betigtJeen Females 1-ith very young pups (hence are post-partum) are vlary and probably less likely to be caught in nets Females with fetuses appear to be more likely to from handling In several cases~ twisting of the reproductive tract by a fetus vJas the actual cause of death Anestrous females (including post partum) on the other hand are less likely to die

Reproduction - 4 - March 1971

There is no question that the birth rate remains high during June and July Kenyons (1969) sum11er sample is limited but shows a large number of Class 5 fetuses Again the sample is biased in favor of pregnampIt animals but not to large fetuses only His field counts as vJell as observations by most other observers indicate an increase in the nUIlber of dependent young througbout the surnmer The number of Lmted pregnancies declines throughout the suTimer tvhile the unimplanted pregnancy rate declines at a slmver rate and breeding remains fairly constant at a lm-v level This indicates that through June there are more implantations than conceptions hmvever the birth rate is substantially greater than the implantation rate By late July and August there are relatively fe1r implantations taking place as shmm by a leveling of the unimplanted pregnant rate and a lotr number of Class 1 fetuses The number of Class 5 fetuses declines as a result of births during June July and August

Therefore the season of greatest pupping may be quite broad running from April through August possibly with the peak occurring in late May or June Until a better summer sample is available tve cannot pin dmm the peak of this pupping season with certainty (See Fetal GrmmiddotJth Rate for further discussion on breeding and pupp peaks)

If the information were precise we could compare subtracting numbers of pups born and numbers of conceptions to deterrnine the actual percent of ferrales breeding implantLng ltmd pupping in each month Unfortunately the data are not Each curve is influenced by more than one factor and ue do not knmmiddotr for sure the magnitude of any of these factors Hmmiddotrever by comparing some of the major samples e may be able to get a rough idea of how much higher the birth rate is at one time of year than another

The combined Mty and the combined September and October samples provide the best comparison Both are large samples One occurs near the peak of pupp and a lmv point in breeding and the other is near the low point in pupping and the peak of b

The number of Height Class 5 fetuses~ post-partum femaless and proestrousshyestrous females come closest to represent a particular event The time spent in each category is comparatively short

There were approximately three times as many Class 5 fetuses in Hay thanmiddot in Septerqber-October There Here also about three times as wany postshypartum females in May In tember-October there were two and a half to three times as many proestrous-estrous animals as in Hay It ~middotwuld appear that the peaks of breeding and pupping are roughly three times as high as the lowest points

At the lmmiddot12st period of pupping 2 to 3 percent of the females have Class 5 fetuses and should pup middotJithin a month If vm assume the post-partum stage lasts L 5 to 2 months (see Recovery of the Uterus) it also appears that 2 to 3 percent pup in the ltwrest months of pupping

Reproduction - 5 - March 1971

Similarly about 3 percent of the females have enlarged follicles during the lmvest period of breeding He do not knmv how long this period lasts hovever

At the peak of pupping about 10 of the females have Class 5 fetuses and should pup in the next month About 15 are post partum indicating about 8 births per 100 females in the preceding month

These percentages are prob2bly slightly high because of hunter bias avoiding females with pups It appeatmiddots that at least 2 to 3 percent of the mature females breed and 2 or 3 percent have pups in any month of the year In the peak breedingmonths such as September and October perhaps 7 to 10 percent breed and similarly in peak pupping months sud1 as Nay 7 to 10 percent of the mature females pup

Gestation Period

Barabash-Nikiforov (1947) listed the gestation period of 8 to 9 months This apparently was based on the case where a fentele mated in captivity as reported by i-Ialkovich (19 37) According to Barabash-Nikiforov a pup was born 8 months later although it died While it was fully formed~ it may have been premature

A number of births have occurred in captive animiddotmals held by the of Fish and Game in recent years In all cases the pu-ps ~-ere sc-middot1all usually less tha1 1600 g but still than the smallest pups found in the -rild All died after birth

The gestation period of 9 months has been repeated in the literature but apparently these statemsnts are based on Barabash~Nikiforol (19lf7)

Lensink (1962) also estimated a gestation period of 8-9 months based on field observations He felt that the peak of breeding was in August or September and the of pupping ~Alas in l-1arch or April

It has been demonstrated that the gestation of sea otters a relatively long period of tation (Sinha 1966 Kenyon 1969 and present study) Therefore any attempt to estimate the length of gestation by examinatioD of tracts must take both the unimp1anted and Janted periods into accour1t

Kenyon (1969) presented an estimate of the implanted period by assuming that the cube roots of fetus weights fall in a straight line after the embryo is established (Hugget and Hiddas 1951) and by that the European otter and Aluerican river otter would have similar fetal growth rates By plotting tenn fetus on a line established by the European otter he estimated an gestation of about 120 days not including the period for establishment of the ewbryo

Kenyon (1969) then an estimate made by Dr D G Chapman using the method of Hugget and ~-Jiddas (1951) v7here a regression line is fitted

Reproduction - 6 - March 1971

to the cube roots of the raeail of the mean weights of fetuses in each of five vreight clas3es plotted t the mean time of year

From the regression coefficient ob he estimated an implanted period of 154 days By taking the mean of the pregnant animals that are unimplanted pregnant estimated the tmimplanted period as about 75 months 1Ulmv-ing a half month for establishment the embryo he estimates a total gestation of 12-13 months

The data used by Chapman included a January-February sample of 26 a March-April sample of 49 and a Nay-August of only 9 The surtmler sample is more biased than the others and quite small yet the estimate relies heavily on this sa~ple when fetus size is the greatest

Table 2 presents a neTw- estimate of the length of the implanted period using the method used by Chapman but including data from the present study in addition to Chapmans data Table 3 presents a similar estimate of the length of the unimplanted period

The following is a comparison of the two estimates

Unimplanted Period 230 days 66 days Establishment of Embryo 15 15 days Implanted Period 154_~~Yrgt~ _73 d~s

Total Gestation Period 399 d2ys 154 days or about 13 months 5 months

Obviously there is some problem tiith the the technique or both A similar calculation mcde before the Hay sample had been collected produced an estimate of 109 for the implanted period and about 8 months for the total ation period

This method of calculation assur1es that there are definite peaks in breeding and pupping If breeding and pupping occurred evenly throughout the year the average fetus size 1vould remain constant thruughout the year and the technique would not tvork An ideal situation v10uld be when all individuals breed and pup 1vi thin very short periods Sea otters do have but are not well defined Very large s v-ould be required in such a case because there are ahvays fetuses of all sizes and the periods of maximum breeding are broad

Obviously Chapmans sample vas inadequate llthile the number of fetuses available for the latest estimate seems large and are well distributed throughout the year the a_ctual nu~ober in any cne fetus vJeight class at any one time of year is small A look at Table 1 shows that there is little consistency in many of the classes (Fig 3 shmvs that Class 1 and 5 fetuses do fit a pattern)

Table 2 Estimated Length of Period

Percent of Fetuses in each Height Class

Time of Collection After January

1 month 31 15 15 38 0

3 months 18 12 22 35 12

5 months 18 8 8 36 30

7 months 3 8 16 29 45

9 months 20 7 10 40 23

10 months

14 18 12 33 23

---~--

r1ean Time in Nonths After January 50 57 53 58 70

Cube Root of Umveighted Mean 063 17 36 7lf lllf

Specific Grmth Velocity 0169day EstirGated Implanted Period = 73 days

Table 3 Length of Unimplanted Period

Months After Unimplanted Unimplanted Percent January Pregnant Pregnant Unimplanted

1 month 35 26 58

3 months 49 49 so

5 months 128 179 42

7 months 22 37 37

9 months 50 57 47

10 months 55 44 56

Unweighted

48

Reproduction - 7 - March 1971

All of the estimates very heavily on su111mer samples 1vhen the fetuses are largest These sauples are knmm to be biased probably in favor of larger fetuses There are more fetuses at this time of year but it is exaggerated in these SCilllples

The latest estimate is not necess better than Chapmans even though it uses more and larger samples The velocity is twice as high as that calculated for fur seal and a 5 month gestation period is not consistent with other information available

This technique for estimating the lanted gestation period is not capable of providing a reliable estimate with the available data Therefore the 12 to 13 month estimate and any calculations based on it are meaningless

Much of the discrepancy lies in the estimate of the unimplanted period The present estimate that 48 percent of all pregnancies throughout the year are unimplanted based on an even distribution of relatively large samples and is probably more accurate than Chaprnan 1 s estimate of 60 percent

There are several other ways of guessing at the length of gestation Kenyons (1969) estillate of an lanted period of 4 months using the cube roots of term fetus weightscannot be relied upon entirely but it may be closer than the other estimates If gtmiddotJe estimate the uninplanted period from this using the 48 percent ted factor and allow for the establishment of the embryo we get a total gestation of about 85 months

Lensinks (1962) method of detennining the and peak pupping periods and taking the time between the peaks as the total gestation period has some merit TJnile Lens ink tried to determine these peaks by field observations we can determine them more accurately using the condition of reproductive tracts

Lensink felt that the peak of breeding middotlas in August or September The data in Fig 2 indicate that he may seen the beginning of the period of est breeding activity but that the probably in October or possibly November Lensink estimated period in larch or April Again the data indicate that he v1as s the beginning of the period of greatest pupping but the actual peak is probably in June or July At this time the number of Class 5 fetuses is greatest The nucJber of post-partum females probably and the number of implanted pregnant females is dropping from its If the gestation period of

11 11the average animal lasts from October or November to June or July we get a period of 8 or 9 months tThile Lensink 1 s was early 1 the duration agrees vith the present data

The peak of implantation when the greatest number of blastocysts are implanting can also be roughly estimated from 1 and 3 In February the number of unimplanted middot is dropping and the number of implanted pregnancies at this time th2 number of Class 1 fetuses probably reaches a

Reproduction - 8 - March 1971

Therefore it appears that the unimplanted period lasts 4 to 5 months and the implanted period from 4 to 5 months 1vi th a total gestation period of 8 to 9 months It should be recognized that these estimates are based on general trends and not distinct Therefore they are approximate However the relative length of the unimp and implanted p2riods agrees with the percentages noted previous and this estimate agrees vlith the one based on Kenyons (1969) comparison vdth other species of otters

If the gestation period ~Jere 12 months the percentage of pregnant animals would remain constant throughout the year If it vJere only slightly longer or slightly shorter there would be only a slight fluctuation unless there were a distinct breeding period Fig 4 indicates a substantial fluctuation indicating a gestation period substantially shorter or longer than 12 months

The peak of pregnancy occurs in February after the period of t breeding activi The luH point on the pregnancy curve occurs after those animals giving birth dur the period of t pupping have pupped That is the t point on the pregnancy curve occurs after the main breeding and the lmves ~ point is after the main pupping period These periods actually last for 4 or 5 months so these points are well after the peaks of breedtng and pupping

The time betbulleen the peak when the pregnant and the lmr point Hhen the fewest are the average gestation period This is about 85 months (Fig 4) The anestrus curve (Fig 1) shmvs the same thing betng a negative of the pregnancy curve

Hith three different methods we have estimated a gestation period of 8 to 9 months Hith the unimplanted period roughly equal to the implanted period This agreas bullmiddotlith the observed period in captivity (BarabashshyNikiforov 194 7) and with field estimates Any gestation period differirg greatly from this does not fit the data Conceivably one could apply a 20 to 21 month period to the SaTEpound information however if this 1vere the case females would have to breed vJhile accomp by a pup or be on a four year reproductive cycle This prospect stretches the imagination

There is a definite possibility that the gestatton period varies in sea otters as in land ot te~cs This most likely -middotwuld occur through variations in the unimplantec1 period The above discussion refers to the average gestation period Until better information to the contrary is obtained we must continue to assume that the average period is 8 to 9 months

Fetal Rate

the estimates of the gestation curve are corrects Kenyons (1969) Fig 4 should approximate the grouth curve of the fetus Fig 5 duplicates this and shmvs the actual Heights Fig 6 shous the same information more graphically It is possible to esttmgtte the of time in each v7eight classlt Fig 6 does not take the period for establishment of the embryo into account so Height Class 1 may be longer than shmm perhaps 15 days if we accept Chapcnans (Kenyon 1969) es e

--

3 0

~

shy

----- -shy --middotmiddot~middotmiddot

___ ___1__

_

(middot-)

_ ft -) J ~)___

~-

3

Reproduction - 9 - IYarch 1971

The short time span of Classes 2 and 3 explains the relatively small numbers and the lack of a consistent pattern found in those classes (Table 1)

Using this curve it should be possible to predict the time of birth and with less precision the time of Luplantation and conception of any particular fetus By plot the estimated birth dates of a large number of fetuses rve should be able to drav curves Ttlhich approximate the breeding implantation and pupping rates of the population throughout the year

Unfortunately there is overlap bet1veen samples collected at different times of year Some of the samples are too small to provide a comparison We vould have to give eight to each in order to combine them Therefore only the ttay September and October samples are used lfuere the September and October samples overlap t1lt10 separate points are used Fig 7 presents the estimated birth dates grouped by month Fig 8 separates the dates into half month groups February and Narch are not represented and April is an estimate based on post-partum females rather than fetus size (see Recovery of the Uterus after Parturition)

The peak in this curve may be exaggerated It has been shmvn previously that fetus size is less consistent bet-Jeen samples than the other categories (Table 1) because the sample size of each grouping is small Therefore the exact shape of the curve may not be correct If however we assume the approximate of the peak is correct we can construct a model of the based on our estimates of the gestation period besed on a fetal grmJth rate estimated from this estimated gestation Therefore our reasoning would be some~hat circular if -Je used thjs to prove the estimate Hmvever we can see how this model fits the rest of the data A good fit would tend to support the model and the estimates on which it is based

For this model He use the curve in 8 to represent births We assume a gestation period of 8 months -rith the unimplanted and implanted periods each lasting 4 months TheTefore v8 draw curves identical to the birth curve but moved backward 4 months for implantation of the blastocyst and 8 months for conception (Fig 9)

A comparison of the peaks in Fig 9 with the data in Figs 1 2 and 3 and the narrative b on those indicates a fairly close fit The data indicat that the peaks may ~ctually occur slightly later than indicated in 9 but in the timing and distances bet7een the peaks fit

The sharpness of the birth curve indicates that the breeding in1plantatimiddotm and pupping peaks mey be more distinct than indicated by the curves in Figs 1 and 2 It is possible to fit curves with more abrupt changes to the data Fig 10 shows the same data lith the curves drawn to more closely fit the model tfuether these curves are more accurate or not is open to debate The data on Hhich the birth curve is based is not that strong The numbers in each month are relatively snall If such distinct did occur it seems likely that field observers ~vould have seen the effects Nost field observations indicate broader peaks that are not gtvell defined

2

G

X

middot ~ -- (-- ) r~ ~ - - -- ~

yen~ r- ~--~

3

i__middot -~---

~

gt

- cshy

Reproduction - 10 - March 1971

~e of _Sexual Maturity

Tables 4 - 7 list the frequency of nmnbers of corpora albicantia and corpora lutea found in females of different areas It should be recognized that the ages may not al-1ays be exact Animals ATTichi tka to become sexually mature Hhen about 4 years old It appbull~ars that more animals may become mature earlier on Adak The information is too limited to dratmiddotJ any definite conclusions from this hmmiddotJever the food availabili ty is probably better at Adak and an earlier age of sexual 1naturity may be a response to better nutrition

Ovulation

From Tables 4 - 7 it can be seen that the maximum number of corpora albicantia for each age roughly the numb~r of years after sexual maturity that some animals ovulated once every year after maturity It can be assumed that many corpora albicantia are invisible macroscopically after several years particularly if they are not remnants of a corpus luteum of pregnancT Therefore it Ls possible that most females ovulate every year This poss ili ty is also supported by the high incidence of large follicles in lactating females (TabL~ 10)

faximum

Tables 4 ~ 7 indicate that at t some females continue to breed at a very old age The samples are teo small to determine 1rhether a t number become unproductive at any point

Ovulation

Many mustelids are induced ovulatoTs

On September 14 1967 a female vJas tsd Hhile copulating She appeared to have Oilnlated shortly before Because sea otters may copulate several times over a period of two or three (Kenyon 1969) ovulation may have been induced by a previous

ficance of

Delayed lon is a charact8ristic of the reproductive cycle of most mustelids Several theories on the ccdv of a delay have been advanced Most assume that spring is the most favorable time of year for survival of nemiddotJborn young but that either a wir1ter breeding season is not favorable or that the presence of a Ttlale vhich occurs only during the

season is for survival of the young of the previous pregnancy (1963) discusse the evolution of delayed implantation in mustelids and points out that there are exceptions even in arctic areas middotJhere time of birth be consLdered more may be born at any tin1e Scheduling the and made possible by delayed mey be advantageous but is not necessary

----

a - o lt bull y bull bull l w laquo bull _ ~ ~

Table 4 Frequency of occurance of Numbers of Corpora Albicantia Amchitkll - June-August 1968

N U M B E R 0 F C 0 R P 0 R A A L 8 I C A N T A

I

~I -AGE -~ 1 I 1084 6 92 5 7(Years) 1 middot shy

- 0-l - ~~ ~- middot~middot- middotmiddotmiddotmiddotmiddot

1-2

2

3

4 1 I

2 (1)

6

middot5

1 (1)1

2 (l) 17

1 ( 1 )

2 ( 1)

18

l (1) I ( 1 ) 9

10 1 ( 1) L ~j( 2 1 ( 1 )

2 ( l) 1

12

11

1 (1)

I I (I)

1

2 ( 1)

13

1 ( 1 )

2

114

115 I116

17

I 18

I I

I

119 I

Numbers inside parentheses =Number of individuals with corpus luteum

11

Table s Frequency of oc~urance of Numbers of Corpora A1bicantfa _Kanaga - October 1968

N U M B E R 0 F C 0 R P 0 R A Al B CANT IA - I I I

AGE 84 I I

5 I l 6 7z 3(Years) 1

l rbull ~middot--

--- shy

0-l

J l-2

~ 2

3

4 2 ( 1)

5 3

Ibullbull 6 6 (3)

4 (4)7

8 4

3 9 2 ( 1 )

j iI 10

I 1 (1)11

12

13

14

1 ( 1)

16

15

17

18

Numbers irisi

-

J

(Plus 2 with corpus 1 uteum but no corpora a 1 b i cant i a)4(1)

1

1 (1)

5 (1)

10(5)

3

3 1

3 (2)

2 ( 1) 3 (3)

1 ( 1)

2

1 ( 1 ) 5 ( 1)

1 ( 1 ) 3 ( 1)

1

3

2 ( 1 )

1 (l)

1 (1)4 ( 1)

2) 1 (1) 3 (3) 4 (1) 12 ( 1 ( 1 )

1 ( 1)

1

2

1 ( 1)

1 ( 1)

1

I I parentheses = Number of individuals with corpus luteum

11

Table 6 Frequency of occurance of Numbers of Corpora Albicantia Mid Pt-~Blind Pt~ Adak- October 1968

N U ~1 B E R 0 F c 0 R P 0 R A A L B I C A NT I A

-1AGE

~

9 1084 75 6Years) 1 2 3 - shy ----- --middot shy~--

- _ Q~L

1-2

(Plus one with_corpus luteum but no corpus a1bicans)2 1 l I I I I I ) (One with corpus luteum but no corpus albicans 3

4 2 (1)

l5

16 1 (1)

1 1)7

8

2 (1)9

10

111

12

I13

14

15 1 (1) 16

17

18

1 ( 1)

1 (l)

1

1

1

2

1

1

1

1(1 )

1

1 (1)

1 (1)

1

I

1

1

1

1

-I

Numbers Inside rentheses = Number of individuals with corpus luteum

1

Table 7- Frequency of occurance of Numbers of Corpora A1bicantia Three Arm Bay - Bay of Islands Adak - October 1968

AGE (Years)

Q-1

t-2

2

3

4

s 6

7

8

9

10

11

12

13

14

15

16

17

18

Numbers

N U ~ B E R 0 F C 0 R P 0 R A A l B C A N T I A

~L 3 4 slE - 1middotshy

7 1~8 __J~l_11 shy

1-

One -Ji th corpus 1uteum but no corpora _a 1bicantia I I I I I I I

One vJi th corpus luteum but no corpora al bicantia

12 I I (Plus one with corpumiddots luteum but no corpora albicantia)1 (1) 3

1 (1) 2 (1)

1 2 (2) l 1

1 (1) 4 (1)

2 (1)

2 1 )

1 ( 1 ) 1

2 (1)

2 (2)

l22 3

3 ( 1)2 21 ( 1)

2 ( 1)

2 ( 1)

l ( 1)1

1 1

1 I ll

I

I I

inside parentheses Nurnber of individuals with corpus luteurn

Reproduction - 11 - March 1971

The sea otter lives in an area of relatively uniform temperatures However storms tend to be more frequent and violent in fall and winter Survival may be better in the late spring and sunrmer Therefore there may be some advantage to birth in the late spring However there does not appear to be a great deal of advantage to a breeding season at any particular time of year

Wright (1963) pointed out that most musteHds lant after daylight begins to increase usually around February This is the period when sea otters implant at the t rate If the time of implantation is influenced by daylight the period when the greatest number of births occur could be shorter than the equivalent breeding period The data do not show this although the possibility cannot be ruled out

The fact that substantial numbers of sea otters are breeding implanting and pupping at all times of the year indicates that hile there may be some advantage to certain events taking place at a particular time of year that advantage is not great enough for a distinct breeding season to evolve

Sea otters exhibit certain characteristics that are comnon to most mustelid breeding cycles but these chfracteris tics are not as well developed as in most es

It is interesting to note that Hright (1963) correlated the molt of Mus with the implanted period TI1e molt began around the

ion and ended around parturi tioD

Sea otters molt all year and some are implanted pregnant at all times At this time e cannot say uhether any between molt and pregnancy exists

Fetal

Kenyon (1969) found that the number of caudally presented fetuses roughly equalled the nunber of cephali presented fetuses He suggests that this indicates a lack of tatim1 for birth occurring in water and infers that sea otters must give birth on land

Fetuses of all 1veights in the present study also appear to be randomly oriented however 97 Class 5 fetuses (1000 g and over) showed 60 percent cephalic ion while only 40 percent shovJed caudal presentation

The orientation of large fetuses should be a better indicator of orientation at birth Smaller fetuses may position Therefore it appears that more sea otters are born head first than tail first

If you accept Kenyons premise that caudal presentation is necessary or at least beneficial to birth in water this more recent information supports the supposition that birth occurs on land The little information available indicates that birth does occur on lando but there have been unconfirmed reports of birth in water

Reproduction - 12 - tltIarch 1971

At least some sirenians are born head first indicating that caudal presentation is not necessary North of Unimak Island a large population of sea otters lives off-shore There is little evidence that any numbers come ashore It seems likely that many of these animals are born in the water

Of 305 implant pregnancies 138 fetuses (45 percent) had implanted in the left horn and 167 (55 percent) in the right horn Most of the difference was in the 1970 Amchitka sample ~fithout this sarnple 48 percent implanted in the left horn and 52 percent in the right horn Kenyon (1969) found an equal number in each horn If a real difference exists it is probably exaggerated by the 1970 A~1chitka sample

Out of 305 implanted pregnencies the point of implantation was on the same side of the reproductive tract as the corpus luteum in all but three instances In one case the corpus lutaum was in the left ovary and the fetus in the right porn In the second case the corpus luteum in the right ovary but the fetus was in the left horn In the third case there was a corpus luteurn in each ovary and tHo fetuses (of different sexes) in the left horn

It appears that blastocysts rarely cross from one horn to the other

Because there is usually a long period of time parturition and the next est_rus is little if any inhibitory effect on ovulation caused by the corpus of the previous pregnancy The ovary producing the largest follicle appears to be random and there does not appear to be the tendency for a pregnancy to be from the opposite ovary from the previous pregnancy In many cases there are many more corpora albicantia in one ovary than in the other

When a loses a pup shortly after birth and enters estrus before the uterus has conpletely recovered and before the corpus luteum has completely degenerated there may be some inhibitory effect Of the ll+ such cases identified six had large follicles in the same ovary as the new corpus albicans and in the opposite ovary It appears that any inhibitory effect by the corpus luteum only lasts for a short time

A few cases bullv-ere found vJhere one ovary was not developed or othenvise not capable of producing ova Because one ovary may support repeated pregnancies such animals may be as productive as those ~vith both ovaries active

Kenyon (1969) reported that most implantations occur within the central third of the uterine horn This held true for the animals in the present study they may implant In one case with a near term fetus the placenta covered the at the base of the horn blocking the exit of the fetus

Reproductive - 13 - March 1971

Birth iltleigh t

Barabash-Nikiforov (1947) listed the vJeight of a ne-v1born sea otter as 2000 g Kenyon (1969) presents the best quantitative data published to this time He found a maximum fetus weight of 1869 g 1-Thi1e he found pups as small as 1020 g those under 3 lb (14 kg) had died shortly after birth He estimated that successful birth weights range from 3 to 5 1b dr 1 4 to 2 3 and for purposes of calculation assumes an average birth weight of 1850 g to 1900 g

The weights of Weight Class 5 fetuses the smallest pups found living in the wild and three pups born in captivity (all died within 24 hours) are presented in Fig 11 These data tend to support Kenyons (1969) estimate of the ~veight at birth Few living pups middotHeighing less than 1350 g or about 3 lb and fevJ fetuses over 2000 g ( 4 5 lb) are found Kenyons upper estimate appears to be slightly high although there are extreme cases in both directions One pup weighed only 2 lb (900 and one fetus could not be weighed accurately on available equipment but weighed over 2500 g

One female with a 4 lb (1816 g) pup still had the placenta attached to the uterine wall and must have given birth shortly before being collected

From 11 it appears that most pups are born when they weigh in the neighborhood of 1800 to 1900 g in most of the populations sanpled Hmvever the and Delarof s indicate a bith Jei of between 1600 and 1700 g The Delarapound sample is srEall but the Tanaga sample the largest containing over a third of the large fetuses collected and was collected at a time of year when the birth rate is very high

This may be a reflection of the phys condition of the adult females and indirectly a reflection of food availab ty There are other factors indicating that the population is declining because of overpopulation

This lov1er birth yeight raises the question of middotwhether the pups are born earlier or whether are in poorer condition when born In an effort to gain some insight into this the fetal of Class 4 and 5 fetuses from Tanaga were plotted against their total lengths (Fig 12) middot Fetuses from other eastern Adak which appears to be a very healthy population were checked this curve fonaed by the Tanaga fetuses In all cases they fell Jell Jithin the limits of the Tanaga data indicating that there is no difference in the of fetuses of a given length It seems unlikely that a reduction in the rate of weight gain vould be completely proportional to a reduction of linear growth It appears that the rate of growth of fetuses is relatively constant but that females in poor physical condition may not be able to support as large a fetus as those in good condition so parturition occurs earlier

This may have sone effect on information on this

Fig 12 demonstrates some other interesting points There is no difference in the ratio and the maximum fetus size of males and females

~

(

~

-s 2 0 C

U)

0

--

ez ()

~) I) middot _t t- (

j -~middot -~

Reproduction - 14 - March 1971

This infeTs although dolt2s not prove an identical grmgtlth rate afld birth weight for males and females The circles on Fig 12 show the ~eight-length ratio of pups These fall below the curve formed by fetus weight-length ratios indicating that pups of a given length are some-Jhat heavier than fetuses of the sarne length This indicates an increase in weight immediately after parturition

Sex Ratio at Birth

Kenyon (1969) found that of 58 fetuses for which sex could be determined~ 45 percent vJere male and percent were female He assumed an equal sex ratio until a larger sample could be obtained

In the present study 111 fetuses were male and 144 ~vere female When this sample is combined with Kenyonts the sex ratio of 313 fetuses is 44 percent male and 56 percent female

Sex Ratio of the

In the process of harvesting capturing animals for transplants and delineating male and femalen areas it has become apparent that there are more females in the population than males Our general ion is that perhaps 60 to 65 percent of the sea otters are females Kenyon (1969) demonstrated that more juvenile males than females are found dead on the beaches of Amchitka lhe age structure of harvested in 1968 indicates that males may not live as long as fetnales

The lower birth rate higher juvenile mortality rate and slightly shorter life expectancy of males could account for the unbalanced sex ratio in the population

The sex ratio may vary from one island to another Presumably there -vmuld be more females in a population that has reached the carrying capacity of the habitat where juvenile mortality is higher There is some indication that males are the first to expand into new areas of unoccupied habitat Therefore in recently populated anas one might find more males However this situation vould be

Lit

The normal litter size of the sea otter is one This has been recorded by almost every observer since Steller Barabash-Nikiforov (194 7) reported twins in utero and mentioned other reports of twin fetuses from the early days of SnoF (1910) recorded a set of neHborn tdns In more recent studies~ both Sinha et al (1966) and Kenyon (1969) found reproductive tracts with two corpora lutee but in all cases only one fetus las present In thousands of hours of observation by many observers no tbullvin pups have been reported

In the present study 24 instances of multiple ovulations were recorded Five of these resulted in hv-in and one in triplet fetuses The information is summarized in Table 8

Table 8 Multiple Corpora Lutea Fetuses

Total Pregnant Females 565

Implanted Pregnancies 316

Unimplanted with 2 CL (corpora lutea) 9

Implanted Pregnancies with ) L CL and 1 Fetus 8

Implanted Pregnancies Vlith 3 CL and 1 Fetus 1

Implanted Pregnancies gtvith 2 CL and 2 Fetuses 5

Implanted Pregnancies middotwith 3 CL and gt Fetuses 1

Total Nultiple Ovulations

I

Percent Nultiple Ovulations 42

Percent of Pregnancies tvith Nultiple Fetuses 19

Corpora Lutea per Pregnancy 105

Fetuses per 102

Reproduction - 15 - lvlarch 1971

From these data it appears that in over 4 percent of the estrus cycles more than one ovulation takes place Of these about half result in the development of more than one fetus In most cases the were of a relatively large size and probably would have been born normally

All multiple fetuses appeared to be the result separate ovulations All had separate placentas and in some cases were different sexes Four tracts with twins had both fetuses in the same horn one had one in each horn and the tract with triplets had one fetus in one horn and t~vo

in the other Another tract had skeletal remains of triplets in one horn that were being resorbed Another tract appeared to have had five fetuses in one horn but they were almost completely resorbed One might infer that physically there is not enough room in one horn to accomn10date more than two fetuses for any length of time

With the exception of the newborn tvins reported by Snow (1910) there are no docu_mented records of female sea otters with tvin pups There are occasional unconfirmed reports of twin pups but as Kenyon (1969) points out a female may tolerate a pup in addition to her ovm but it is unlikely that a female could care for more than one pup It is unlikely that more than one pup survi~es In any case orily 2 percent of the pregnancies produce more than one pup 1middot1ultiple births cannot be a significant factor in the productivity of sea otter populations

The presence of a corpus luteum without gross evidence of implantation t-7as assumed to be an unimplanted pregnancy No attempt ~vas made to locate unimplanted blastocysts As a result e have little information concerning the survival of the blastocyst Of 206 reproductive tracts classified as nulliparous on the basis of the appearance of the uterus 12 had one corpus albicens and one had two Some of these may have been large attritic follicles Others may have ovulated and even conceived but implantation did not occur All of these cases represent the failure of the antmals first estrus

A total of 16 resorptions or possible abortions were identified (Table 1) The follmJing is a brief descdption of these ~7i th possible causes of some

Des

Resorption of blastocyst or ovum (corpus luteum 4 degenerating no evidence of ion)

Failure at time of (possibly because 1 of growmiddotth inside horn adjacent to implantation site)

Resorbed or aborted fetus (implantation at end of horn) 3

Resorbed fetus (umbilicus tvTisted tightly) 1

Resorbed fetuses (three or more fetuses in one horn) 2

Resorbed fetus (cause unknmm) 5

Reproduction - 16 - March 1971

Six of these resorptions may have been caused by a lack of room for growth of the placenta because of crowding from other placentas a growth or the end of the uterine horn These six and the one vith the twisted umbilicus are probably the result of random events or accidents

The numbers of resorptions are too small to accurately measure the frequency of such occurrences although the relatively large number in the Tanaga sample (Table 1) may be a reflection of the physical condition of the animals Part of this number is probably due to the fact that there are more pregnant animals in the sample In general the samples with the most pregnancies also conta~~ned the most resorptions Half of the Tanaga resorptions can be attributed to random events but five vere still due to unknown causes Eight of the 10 resorptions fo1md on Tanaga came from a 20-mile stretch of shore about 25 percent of the harvest area

While concrete conclusions cannot be drawn from these data they do raise the possibiliy that under certain conditions 5 percent of the pregnancies may fail

near Parturition

Collecting methods were such that the presence of a pup with a female was often overlooked Therefore the lack of a pup with a post-partum or lactating female did not mean that the pup had died However 14 females shmving of recent pregnancy were estrus presumably having lost their pup shortly before or shortly parturition

Again the magnitude of this mortality cannot be determined but it appears to be of the same order as the in utero mortality

after

Reproductive tracts were subjectively classified as post-partum or anestrous on the basis of degeneration of the corpus luteum (or appearance of the corpus albicans) the size of the horn of pregnancy and appearance of the placent scar In an effort to get an idea of how long it takes for the tract to return to normal the size of pups accompanying females in each category is listed in Table 9

It appears that females pass the subjective boundary from post-middotpartum to anestrus Hhen the pup about 10 lb and is around 75 to 80 em long

We do not know a great deal about the early growth of a sea otter pup but judging from the cur1e e9tablished based on cementum deposition of older animals and from tooth eruption a 10 lb pup is probably in its second month of life

Table 9 Pups Accompanying Pos artum Females

Sex llleight

M 2 lb F 3 F 4 M 5 F 55 F 625 F 775 M 10 F 11

Total Length

47 em 52 56 60 65 65 70 81 (borderline post partum-anestrous) 82 (late post-partum)

Pups Accompanying Anestrous Females

F 10 lb 78 em M 11 79 M 12 8l F 13 83 M 13 92 H 16 83 M 17 91 1 19 87 M 19 90 F 21 107 F 22 96 1 22 101 M 2Llmiddot 97 1 26 100 F 28 103

Reproduction - 17 - Narch 1971

Frequency of Breeding

Steller (1751) reported female sea otters with ne~vborn pups also accompanied by another pup that seemed to be no more than a year old Nurie (1940) recorded an instance of copulation by a female accompanied by a pup and Jones (1952) caught a young pup simultaneously gtvith a copulating pair Lensink (1962) felt that few females breed until the pup is a year old but mentions females with small pups also accompanied by large pups

I believe that some of thes2 observations may be misinterpreations of the situation Females will occasionally leave small pups for a time often near other animals The pups Lensinkmentions would be 2 years old and probably veigh 30-35 lb Sea otters are gregarious and subadults may remain near other animals appearing to be accompanying their mother I suggest that most of Stellers and Lens inks large pups ere such cases

Females gtvi th pups probably do breed occasionally but the most recent evidence indicates that this rarely occurs Kenyon (1969) records no instances of females accompanied by pups copulating and states that he found no females- knm7n to be accompanied by a pup that were This holds true for the animals collected in the present study However the nature of the harvesting operation is such that many females accompanied by pups were not recorded as such Therefore we are forced to look at lactating and assume that they were accompanied by a pup at the time of collection or shortly before le 10 sm1marizes the reproductive condition of the lactat females In the 1967 and 1968 samples some animals vith enlarged marmary middotmre listed as lactating As a result some females with near tenrr fetuses Here included Because this condition is considered to be associated with the unborn pup rather than a previous pup these animals were with the anestrous animals in le 10 In 1969 and 1970 the presence of milk was used as the sole criterion and no term animals we-rmiddote in the sample

Of 246 lactating females only one had an ted fetus (excluding the near term animals from 1967-68) Several others had small corpora lutea indicating that they recently become or possibly had large

vhich had luteinized but tvere not actually pregnant The remainder had follicles

This information that accompanied by a pup rnay enter proestrus and even ovulate but that they only rarely mate That they enter estrus is subs iated by the f2ct that the number of corpora albicantia in many tracts equals the age of the amplimal minus the three years prior to nBturity (see les 4-7) This indicates that large follicles tend to develop year after sexual nl8turi ty is reached whether the female is accompanied by a pup or notlt

Some of those lactating females with s1all corpora lutea may have recently weaned or othenvise lost a pup and are already pregnant again Malkovich (1937) took a pup least 3 months old) away in cap She mated 12 days later here she had the male It appears that a female enters estrus shortly after Heaning

Table 10 Reproductive Condition of Lactating Females

Location

Bay of Is Adak I Sept 1967 9 1 100

Allchitka I Sept-Oct 1967 17 2 105

Mid Pt - Blind Pt Oct 1968 19 3 136 Adak I

Bay of Is Adak I Oct 1968 22 4 154 3

Kanaga I Oct 1968 32 s- 135

Al1chitka I July-Aug 1969 4 2 333

iTanaga I May 1970 56 10 152

Delarof Is May 1970 18 3 143

Amchitka I May 1970 36 3 77

---~--middot--middot----~-~----~----middot~-middot--~----~----- -------~-~ middotmiddot--middot-shy

TOTAL 213 33 134

Anestrous or pregnant with term fetus

]_ Large follicles or corpus luteum present

3 Includes one implanted pregnancy with small fetus

Reproduction - 18 - March 1971

The question of hml long it takes for a female to enter estrus again after losing a small pup arises Eleven tracts of the 1970 sample and ttvo of the 1968 Kanaga Sampnple shmJed signs of being pregnant recently) but were in proestrus These animals had an enlarged horn usually middotlith a slightly pigmented area but no actual placental scar a11d a recently fonned corpus albicans However they also had enlarged follicles and the tvalls of the uterus were typical of proes trous animals Another animal was similar but appeared to have already ovulated and a corpus luteum was present In these cases the female has lost a pup either through abortion or vithin the first veeks after parturition and has started into another estrus cycle Three or four of these had follicles vhich may have started to become attritic This may be because the uterus had not recovered sufficiently from the last pregnancy

The implication of this information is that with a normal pregnancy and successful rearing of a pup the female becomes pregnac1t only after the pup has been weaned This n~y be a year after parturition However if the pregnancy fails or if a pup dies even at birth the female may become pregnant again in a few weeks rather than _waiting the full year

Kenyon (1969) suggests that the period middotbetween and the next estrus may long because he found animals that tvere not lactating but had a faint placental scar and an indistinct but recent corpus albicans These animals ltJere cLtssi as anestrus in the present study They are undoubtedly bet1veen veaning of their last pup and the next estrus The number of suh animals is t durirg Sspte12~ cnd October At this time the rate of estrus increases a1d the number of anestrous animals decreases sharply the next feJ months Therefore many of these anestrous anirrals become pregnant 1lithin a month or two Kenyon (1969) suggests that if the female keeps the pup for about a year that the period between births may be smreltmiddotJhat greater than two years He is assuming a 12-13 month gestation period However the present study shows that the estimate of a 12-13 month tation period was based on an inadequate sample Estimates in the present study put this period at closer to 8 or 9 months Presumably the anestrous nonlactating females are in the 3 to 4 month period

This does not change Kenyons (1969) point that the and the next estrus may long It indicates that in a normal cycle it may be months Hmvever as it vvas shmvn above~ it is possible for a female to enter estrus very shortly after losing a pup

We have demonstrated that sea otters breed at all times of year and it is possible for a female to become pregnampJt again after losing a pup sooner than she vould have if the pup had survived HmmiddotJever we have also demonstrated that annual of b and pupping occur and that these are similar in different years and in different populations The assumption can be made that the average length of the entire reproductive cycle is some multiple of a year

Kenyon (1969) assumes that tbe pup remains with its mother for about a year ~-Ieights and measurements cementum deposition skull changes~

Reproduction - 19 - March 1971

observations in the wild and in captivity all support this assumption although it cannot be said with certainty that it averages 12 months It could be s tly ITDre or less

Assuming about 12 months for rearing the pup 8 to 9 months gestation plus a fev months rest bet~veen and the next estrus the average reproductive cycle takes two years

This is supported by Fig 4 which shows that slightly over half the mature females are pregnant in a year Kenyon (1969) pointed out that his January and Harch samples gtvere biased against females with pups a1d therefore in favor of pregnant females because of selective hunting This bias applies to the fall sa~ples as well The summer samples were biased similarly because pregnant females appeared to be less resistant to handling during capture Therefore the percent of pregnant females is shown to be higher than lt actually Has -rhile no correction factor can be applied to all samples the evidence indicates that the percent of pregnaJt females is usually around 10-middot15 lmrer than in the sample For example the 1970 Alnchitka sltLrnple t-ras less biased and was about 9 percent lower than the Tanaga sample vrhich is l)nm-rn to be more biased Also on a June skiff survey 7 percent of the animals counted had pups that were readily identified If ~re assune that this is the nunber of animals avoided it can be calculated that 15 of the sexually mature females all of vhich are t middotTen~ avoided Actually this will vary with the hunter weather time of year etc Hmvever it indicates the order of magnitude of the bias

Using this as a rough correctton for the data in 4 we find that about half or slightly fe-Jer of the sexually mature females are pregnant each year or as indicated ly the average female has a pup every two years

Reproduction - 20 - March 7 1971

Conclusions

1 While sea otters may breed or pup at any time of year there are seasonal fluctuations in the illliTlbers of females in each reproductive stage The pattern of these fluctuations remains the same from year to year and population to population

2 Breeding activity is highest in September October and November

3 More implantations occur during January February and March than at other times of year

4 The birth rate is highest during April Y~y and June

5 Tlvo and a half to three times as many females breed during the peak breeding months as during the months of lowest breeding activity

6 Similarly two and a half to three times as many females pup during peak pupping months

7 During the period of lovest b activity seually mature females breed at a rate of 2 to 3 percent per month During the peak of breeding 7 to 10 percent per month breed

8 During the period of lov1est pupping activity 2 or 3 percent per month pup and during the peak of pupping 7 to 10 percent per month pup

9 The gestation period about 8 to 9 months on the average About half of this time is ~~ ted period

10 Female sea otters become sexually mature ihen 3 to 4 years old

11 Females may ovulate on an average of once a year after sexual maturity is reached

12 Females continue to breed at Cn old age

13 Sea otters retain certain characteristics common to the reproductive cycles of many other mustelids however exceptions are common Scheduling of the reproductive cycle during the year may have some advantages or it may be a trait that has not been entirely lost through evolution

14 Sixty percent of the near term fetuses are cephalicly oriented and 40 percent are candllly oriented

15 Sea otters probably birth both on land and in the water

16 Blastocysts rarely cross from one uterine horn to the other

17 Consecutive pregnanctes may occur in the same horn The side of pregnancy appears random

Reproduction - 21 - March 1971

18 Implantation usually occurs in the central third of the horn but it may occur any~rhere in the horn

19 Birth weights may range from 900 g to over 2500 g however most pups weigh 1800 g to 1900 g at birth

20 Females from areas gtvhere the population is declining because of food shortages may pup earlier and as a result the pups 1 birth weight may average 200 g lmver than nonnal

21 Male and female fetuses gr01middot1 at the same rate ampJd are the same size at birth

22 Body condition of the female has little effect on the growth rate of the fetus

23 There is a rapid ~Jeight gain shortly after parturition

24 The sex ratio at birth is 44 percent male and 56 percent female

25 The normal litter size is one hmmiddotever in 2 percent of the implanted pregnancies two or fetuses survive to near term

26 Multiple ovulations occur in 4 percent of the estrus cycles

27 es do not appeErc- to be able to more than two fetuses in a single horn

28 Up to 5 percent of the p may fail dne to in after implantation

29 An unknown but probably significant number of first pregnancies fail before implantation

30 The uterus is usually considered recovered from a pregnancy when the pup vleighs about 10 lb and is probably in its second month of life

31 Sexually matu1middote females normally have one pup every two years

32 Females vrith a pup may ovulate but rarely mate

33 Slightly less than half of the sexually mature females are in any given year

34 A female may enter estrus within a few weeks of losing a pup Consequently she may become agaln gtvi thout waiting the same length of time as if the pup hacl survived

35 Hhen a pup is weaned normally there is a longer period perhaps 3 or 4 months between weaning and the next estrus

Reproduction - 22 - March 1971

CITED

Barabash-Nikiforov I I 1947 The sea otter (Kalan) Soviet Ministrov RSFSR Translated from Russian by Dr A Birron and Z S Cole Israel Program for Scientific Translation 1962 227 pp

Hugget A St G and W F Widdas 1951 The relationship betlveen marmnalian foetal -reight and conception age Jour Physiology 144306-317

Kenyon K W 1969 The sea otter in the eastern Pacific Ocean USFWS North American Fauna No 68

Malkovich T A 1937 The sea otter in captivity Priroda No 381-87 Translated by J T Naximovitch 1966 Fisheries Research Board of Canada Nanaimo BC Translation Series 657

Sinha A A C H Conavay and K t-J Kenyon 1966 Reproduction in the female sea otter J Hildl Ngrrit 39(1)121-130

Snow H J 1910 In forbidden seas Edward Arnold London 303 pp

Wright P L 1963 Variations in reproductive cycles in North Arrierican mustelids In Enders A C ed Delayed tation Univ of Chicago Press 318 pp

  • SEA OTTER STUDIES - 1970
    • Surveys
    • Harvests and Transplants
      • Sexual Segregation in Sea Otter Populations
      • Reproduction in the Female
        • AERIAL COUNT OF SEA OTTERS- SOUTHSIDE OF THE ALASKA PENINSULA
          • Description of Conditions During March 1970 Aerial Count of Sea Otters
          • Discussion of Results
          • Sanak Island-Sandman Reefs
          • Shumagin Islands
          • Sutwick Area
          • Augustine Island-Cape Douglas
            • SUMMARY OF SEA OTTER SURVEYS
            • Amagat 1 to False Pass Ikatan Peninisula Cape Lazaref
            • Sanak Is
            • Deer 1
            • Pavlof Is
            • Northern Shumagin Is
            • Cold Bay 1
            • Kupreanof Peninsula
            • Mitrofania 1 to Kupreanof Pennisula
Page 4: Sea otter studies - 1970 - Alaska

Description of Conditions During March 1970 Aerial Count of Sea Otters

March 20 Aniakchak Bay to Chignik Lagoon (1022 AM- 1240 PM) Conditions were generally fair with a heavy chop The south side of Sutwick Island and the area between the island and mainland were poor with heavy surf Visibility was very good inside Kujulik Bay however many animals were scattered throughout the area On two passes ~cross the bay 12 and 27 were seen therefore many were probably missed Similar scattering occured in Chignik Bay The overall count for the area is probably low

Pavlof Bay to Cold Bay (130PM-200PM) Conditions poor with heavy chop and squalls The count was superficial however a second superfi cia 1 count of this a rea was made on March 21 and only one otter was seen in Cold Bay

Cold Bay to Caee Lazaref (310PM -420PM) Conditions very poor fair in a few spots Heavy chop dense kelp

March 21 Northern Shumagin Islands (943AM- 1137 AM) Poo~ with chop on windward sides Fai_r on leeward sides (westerly wind) however air turbulence forced us away from high cliffs Heavy surf created very poor conditions on the south side of Unga

Pavlof Islands (1145 AM- 125ff PM) Conditions similar to those in the Shumagins

Chignik Lagoon to Pavlof Bay (305 PM- 535 PM) Conditions varying from fair to poor with chop and surface glare Count concentrated on points and better looking areas Most deep bays not completely surveyed

March 22 Sanak Islands (849AM- 1020 AM) Conditions fair to poor with chop on south side and surface glare on north side Animals scattered and very difficult to see Many otters on rocks Count is probably very low

Sandman Reefs ( 1025 AM~ 1115 AM) Conditions similar to Sanak With the exception of one pod of 450+ the animals were widely scattered and very difficult to pick out Only those very close to the aircraft could be seen Bad squalls moved in and the wind increased preventing a complete count of Deer Island and the eastern half of the reefs

March 23 Wide Bay to Aniakchak Bay (1123 AM- 140PM) Conditions generally good at first becoming fair after Cape Providence with a light chop and some surface glare Otters in Amber and Aniakchak Bays were widely scattered and many may have been missed

Port Heiden to Naknek_ Superficial counts were made from Ugashik Bay to Naknek on March 19 and from Port Heiden to Egegik on March 23 These were made under poor conditions by flying outside the surf line The water was choppy and full of silt No sea otters were seen

11arch 27 Shaw Island to Puale Bay (1030 AM - 1205 PM) Conditions very good to excellent High overcast and very little wind down to Katmai Bay After this the conditions deteriorated rapidly to fair and the count had to be stopped because of a combination of bad weather and lack of fuel As shown on the charts the count was concentrated on the points and islands however the good visibility allowed us to see well into the bays We feel certain no concentrations were missed

The numbers of sea otters counted are presented in the Table Specific locations and numbers are shown on the charts Where coverage of an area was not complete the approximate path of the aircraft is shown

Discussion of Results

Surveys along the Alaska Peninsula have been fragmentary and most of those made were done under less than ideal conditionsmiddot There are a number of reasons for this Weather tends to be poor along the south shore where squall5 form along the mountains Areas where an aircraft can refuel are north of the mountains and there are relatively few passes therefore range of the aircraft is a problem Also there is much shallow water off shore in the area from the Shumagin Islands to Sanak Island Coverage of this area is difficult and many animals are undoubtably missed

The present survey covered most of the area however conditions were such that most of the counts are probably quite low Despite these problems and a lack of continuity we have a fairly good picture of the recovery of sea otter populations in this area

Reports from various individuals are available from the 1930s and 1940s 11ajor surveys by the Fish and Wildlife Service in 1951 (Jones) 1957 (Lensink) 1962 and 1965 (Kenyon and Spencer) covered at least portions of the area In 1969 the southern Shumagin Islands were counted by the Alaska Department of Fish and Game and the present survey covered most of the rest of the area

Basically there are four distinct population nuclei in the area These centered around the Sanak Island-Sandman Reef area the Shumagin Island area the Sutwick Island area and the Augustine Island-Cape Douglas area The following discussion is based on information from reports by Lensink and Kenyon and from Alaska Department of Fish and Game surveys middot

~anak Island-Sandman Reefs

Small numbers of sea otters have been reported at Sanak Island since 1922 No reports came from the Sandman Reefs until 1942 In 1948 27 were sighted at

----Cherni Island The 1951 aerial survey showed 65 around Sanak and 97 in the Sandman Reefs In 1957 251 were seen around Sanak and 508 around the Sandman Reefs In addition two were seen in the Pavlof Islands however few were on the mainland In 1962 548 were counted in the Sanak area and 638 in the Sandman Reefs None were reported from the mainland or Pavlof lsland~however

much of the increase was in the northern area around Deer Island The 1962 survey was probably the best being made under excel lent conditions

AERIAL COUNT OF SEA OTTERS MARCH 1970

Partial or Location Count Date Camp 1ete Count Visibility

Cape Lazaref - Cold lsecty_

Cape Lazaref 3 320 Complete Poor

lkatan Peninsula 71 320 Complete Poor

False Pass - Amagat I 66 320 Camp 1ete Poor

Cold Bay 321 Partial Poor

TOTAL 141

Sanak Islands 239 322 Camp 1ete Fair to Poor

Sandman Reefs

Clubbing Rocks 12 322 Complete Fair to Poor

Cherni I amp Rocks 495+ 322 Complete fair to Poor

Goose I 7 322 Complete Fair to Poor

Hay I 18 322 Camp lete Fair to Poor

Hunt I 2 322 Complete Fair to Poor

Deer I 322 Partial Fair to Poor~

TOTAL 568+

Pavlof Islands

Inner 11iasik 2 321 Camp 1ete Fair to Poor

Outer lliasik 16 321 Camp 1ete Fair to Poor

Goloi 2 321 Complete Fair to Poor

Dolgoi 67 321 Complete Fair to Poor

Poperechnoi 29 321 Complete Fair to Poor

Ukolnoi 2 321 Complete Fair to Poor

yenWosnesenski 4 321 Complete Fair to Poor

TOTAL 122

Partial or Location Count Date Comp 1ete Count vis ib j 1i ty

Northern Shumagin Islands

Unga 184 321 Complete Fair to Poor

Popof 52 321 Complete Fair to Poor

Korovin 46 321 Complete Fair to Poor

Karpa __2t 321 Complete Fair to Poor

TOTAL 286

Cold ~ to Beaver ~ 0 320-21 Partial Poor

Beaver Bay to Kupreanof Pen

Beaver Bay 2 321 Partial Fair to Poor

Cape A 1 iaks in 4 321 Partial Fair to Poor

Guillemot I 16 321 Partial Fair to Poor

__1Kupreanof Peninsula 321 Partial Fair to Poor

TOTAL 23

Kupreanof Pen to Castle Cape 0 321 Partial Fair to Poor

AntJrrCastle Cape to-1-Ji e ~

Castle Cape-Castle Bay 16 321 Partial Fair to Poor

Chignik Bay 118 320 Complete Fair

Nakchamik I 5 320 Complete Fair

Hook Bay 13 320 Complete Fair

Cape Kum 1i un 88 320 Complete Fair

Kujul ik Bay 1199 320 Complete Very Good

Univikshak I 62 320 Complete Fair

Cape Kuml ik 54 320 Complete Fair to Poor

Sutwick I 14 320 Complete Fair to Poor

Cape Agutka 323 Complete Fair~

TOTAL 1766

Partial or Location Count Date Comp 1ete Count vis i b i 1i ty

Cape Kunmik 13 323 Complete Fair

Naka 1i kok Bay amp offshore islands 16 323 Complete Fair

Chiginagak Bay 5 323 Complete Fair

Agripina amp lmuya Bay 105 323 Complete Good

Wide Bay to Puale Bay N 0 T S U R V E Y E D

Puale Bay to c Kubugakl i __]_ 327 Partial Fair

TOTAL 146

Kashvik Bay to L Chiniak 0 327 Partial Very Good to Excellent

Cape Chiniak to Shaw _I

Shakun Is amp Rocks 71 327 Partial Very Good to Exce 11 ent

Kiukpal ik J to Shaw 1 0 327 Partial Very Good to Excellent

TOTAL 71

In the present survey which was made under relatively poor conditions 239 were seen in the Sanak area and 568+ in the Sandman Reefs The latter count Was incomp 1ete and conditions were such that the number of otters was more a function of time spent counting rather than area covered Obviously many were missed As a result not much can be said about total numbers The main change evident is that 141 were seen along the mainland and eastern portion of Unimak Island and 122 were seen in the Pavlof Islands This indicates that a fairly extensive movement northward and along the Peninsula is occuring

A remnant population probably remained along the south side of Sanak Island in the early 1900bulls By the late l940 1 s they began spreading into the Sandman Reefs This northward expansion has continued to the present time There is still unoccupied or sparsely populated habitat along the mainland shore and in the Pavlof Islands There should be continued expansion of this population for a number of years although the numbers around Sanak Island and the western Sandman Reefs may have already reached a peak

With the arrival of substantial numbers in the Pavlof Islands this population is probably on the verge of mixing with the Shumagin Island populshyation No doubt some individuals have moved back and forth in the past but the two populations are almost continuous at the present time

As in past surveys few otters were seen around the north side of Sanak Island and Caton Island This is probably poor habitat for otters The main population is around the south and west sides The higher count on the west end is probably more a result of intensive searching rather than a higher population

Shumagin Is 1 ands

This has been considered the largest of the four populations in the Alaska Peninsula area The most recent counts have not been adequate enough to show any major increase in numbers in the last decade however major changes in distribution have occured which are very similar to the pattern mentioned in the Sanak-Sandman population

There were occasional reports of sea otters in the Shumagins in the 1930s By 1947 Victor Scheffer estimated that 500 1 ived around Simeonof Island In 1953 Hooper counted 633 around Simeonof and Little Koniuj i Island The 1957 survey showed 1829 Most of these were- in the southern islands Five individuals were scattered in the northern islands and one was on the mainland shore near Elephant Point The 1962 survey totaled only 1352 The animals were scattered well off shore and the count was low However the count on Nagai increased from 149 to 338 indicating a continued northward expansion

In 1969 1510 were counted in the southern islands under relatively poor conditions An additional 286 were counted in the northern islands in the present survey and 23 were seen along the mainland north of the Shumagins Again survey conditions were not good

There is a very clear expansion of the population from a center near Simeonof Substantial numbers now occur on all the islands and repopulation of the mainland is occuring The population in the northern islands should continue to increase and most of the habitat on the mainland is not occupied

There is no evidence of an increase in total numbers since 1957middot However the last two surveys have been less than ideal and the large shallow off shore areas have not been covered adequately Considering the survey conditions and the obvious extentions of range it is almost certain that the populations from San-ak Island to the Shumagins have continued to increase In general it appears that the southern islands and reefs have become fully populated and the northern areas are just developing significant populations The entire a rea may be comp 1ete 1 y repopu 1 a ted in the next 10 years This wi 11 depend largely on the status and potential of the off-shore areas

Sutwi ck Area

Reports of sea otters near Sutwick Island are available since 1936 This population was far removed from other populations and is probably a remnant left after hunting ceased In 1951 Jones counted 388 between Cape Kuml iun and Cape Kunmik Most were near Sutwick Island In 1957 889 were counted Nineteen of these were between Cape Kunmik and Cape Providence indicating some northeastward expansion In 1962 949 were seen with individuals straying as far as Cape lgvak and one between there and Cape Kuliak In 1965 a stray otter was seen at Kinak Bay A major shift in the population from Sutwick into Kujulik Bay occured This may be the result of time of year and weather

In the present survey 1766 were counted between Castle Cape and Cape ~~~gtf~aip the majority were in Kujulik Bay Large numbers have moved

~~ranging as far as Cape Kubugakl i The area from Wide Bay to Puale Bay was not surveyed because of weather

The total count for the population was 1912 even though conditions were less than ideal throughout the bcunt and we feel many were missed It is possible that the increase in numbers is entirely due to reproduction assuming a 10 percent annual increase However it is difficult to compare surveys

There is still room for expansion in both directions from this population The greater movement to the northeast is probably the result of better habitat The population around Kujulik Bay is very dense and a large movement of animals may occur if competition for food becomes serious Otherwise we should expect to see continued steady expansion into adjacent areas for a number of years

to come

Augustine Island-Cape Douqlas

For some time a population has existed around the Augustine Island area at the mouth of Cook In 1 et In 1948 approxImate I y 50 sea otters vvere reported from Augustine Island Reports of sightings have been made from Shaw Island to Tuxedni Bay In 1957 Spencer counted 40 at Augustine and one at Shaw Island Lensink counted 52 on Augustine in 1959 but he considered it a poor count In 1965 Kenyon counted 18 on Augustine and 101 in the Shaw Island-Cape Douglas area In March of 1969 Loren Flag saw 62 around Augustine and in May 1969 Jim Faro counted 130 another observer saw about 30 some of which were probably not tallied by Faro

On the present survey Augustine Island could not be surveyed because of weather No otters were seen around Cape Douglas but 71 were seen in Shakun Islands and rocks near the abandoned village of Kaguyak These were probably from the Cape Douglas group

It appears that the animals move about in the area To date no complete survey of the area has been made at one time Until this is done 1ittle can be said about the population other than that several hundred probably occur there

The following table is a summary of sightings and counts made by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game These data were collected by different individuals using different types of aircraft under varying conditions of weather A strict comparison of numbers should not be made from this table

SUM

MAR

Y OF

SE

A OT

TER

SURV

EYS

(Com

pile

d by

p

op

ula

tio

n f

rom

L

ensi

nk

(196

2 T

hes

is

K

enyo

n 0

96

2 amp

196

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(

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92

bull

middot

34

middot

middotbull~middot_

--

J middot

middotmiddotmiddot

61

712

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-

71

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2812~bull

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c

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9 _

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55

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61

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24

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(I~0 0--

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Hook Bay Chignik Bay

13 118

-1 shy

shy

shy

bull

I +

Castle Bay - Castle Cape 16 Nakcharriik I 5 -- Katmai Reef 0

Total 152

middot

~ middot-

bull -middot

bull

-middot

Cape Agutka Cape Kum 1 i k Sutwick I~ Kujul ik Bay Cape Kum 1 i un Unavikshak 1

Total

X

-------8

197 54 14

1199 88 62

1614

X

E c 1 ~

(l) a 10

u

0 ---_ +-

Q) u

gt-sect OJ al middotshy

gt 10 0 c 1

middot- a a middot- (l)1 a Olfil lt(U

bull c

)

r r -middot

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middot i 1~ bull middot- i ~ shy

i

- t ~-middot IImiddot

- _ ~~---Q middot- ---- ~ - ~ ~ --~ G lt-~ bulllt00 middot __ ~-middot middot-middot +~ yen_-_____ _ ---

_-y _ -

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bull ~

v

4gt

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rd ltgt

~~ lltgtJltJshy

rJgt

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9

J J

0

0

lt

bull

1gt ~

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1

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I

I I

I amp~middot

smiddotmiddot I

l bullt

middot

99

10

i

Sg

9

75

~-

MARINE MAMMALS SURVEY - BRISrOL BAY TO AKUTAN ISLAND June 2 1970

OBSERVER J Vania J Faro

PILOT George Tibbets Jr

AIRCRAFr Piper Azetex

middotWEATdER Partly cloudy throughout flight wind 10 miles per hour or less visibility generally excellent One bad area and that was at Akutan Island Air was turbulant there and we were unable to go completely around the island because of fog in Akutan Pass Water had slight ripple throughout flight and ocassionally there was glare but overall conditions for sightinmiddot=r animals was good

Departed King Salmon 1115 AM Returned to King Salmon 745 PM

FLIGHT PA~1 middotAfter leaving King Salmon we went over to the north side of the Alaska Peninsula and proceeded down the coast flying at 300 to 600 feet of altitude Generally we flew aboumiddott one mile off the beach At Port Heiden Moller and Cinder River we checked the outer sand bars for seals and some of the inner bars where I knew seal hauled out

We did not cover Izembeck Laroon very well Instead we flew offshore and checked Amak Island We then fueled up at Cold Bay Leaving Cold Bay we continued dmvn the north side of the peninsula (sea otter sighted) down to Cape Sharichef and crossed over to Akutan Island flying along the south side of it We were able to get around Cape Morgan and fly doNn the beach a few miles but then had to turn back becausmiddote of fo9

The flight path was then eastward to the north side of Tigalda Island At Ugamak Island we circled it completely flying at about 300 to 500 feet We completed the survey at this point and returnt~d to King Salmon generally following the route we had taken on the flight down

SEA OTTER SIGHTINGS

Seven pods of sea otter were seen south of Amak Island These were photographed and later counted from middotthe photographs Beshycause of the density in the larger pods and the fact that some animals in the smaller pods were diving when the pictures were taken some individuals may have been missed

The number counted in each pod is as follows

1071 520 357 68 36 75 30

TOTAL 2157

Several hours later the pods had drifted four or five-miles northeastward (see map) bull In addition 1 one sea otter was seen at Amak Island and sixteen near Tigalda Island

171

-amp

7

7

7 58

57

53

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8 A

l

ltc-~)

53

51 ~9

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)lt

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45

56

4

6

52

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(lbASC

4

8 M

s 1

7 4

2

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4

5

39

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middot-

oo

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60

46

-

-

(

56

88

89

8

8

10

80

85

45

s

SS

II

87

89

78

4

43

95

75 S

it

92

65

98

42

93

82

84

87

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85

82

77

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32

34

81

74

66

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31

41

36

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50

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43

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31

40

37

40

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6

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7 Q

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47

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52

32

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56

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43

41

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35

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152

45

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54

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55

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ps

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t 32

03

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52

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56

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57

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47

56

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e r

ips

57

47

55

56

54

50

5

0

j

48

5

49

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-

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_

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i51

54

_

bullmiddot

-middotmiddot

middotmiddotmiddot-

53

51

52

bull

-

5-

48

bull9

-middotmiddotmiddot

middotmiddot-

54

58

54

53

51

56

53

32

rip

s 43

36 ~

44

(~

39

3

6

313

8

4

44

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middotmiddot

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middotmiddotshy

53

S

Sit 51

53

54 ~

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I middot

bull

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46

bull

49

)

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(

47

middot~~_

middot

38

5

6

535

3

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48

45

39

61

61

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de r

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58

56

54

54

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6

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shy

SEA OTTER COUNT BARREN ISLANDS and SHUYAK - AFOGNAK ISLANDS

June 9 1970

On June 9 1970 an aerial survey of sea otter was flown in the Barren Islands and in the AfognakShuyak Islands area A Gruman Goose was used with Schneider serving as the only observer Offshore coverage was poor Conditions of visibility were as follows

Barren Islands- 1140 AM to 1225 PM- Water calm moderate surface glare Conditions verygood Count complete

Ban Island to Pernosa Bay- 1255 to 135PM- Conditions similar to Barren Islands but surface glare worse in spots Conditions generally good

Marmot Island to Latax Rocks - 255 to 345PM- Increased ripples on surface Glare Bad Conditions fair Count around Sea Otter Island complete although some otter may have been scattered offshore Remainder of count very superficial

Area

BARREN ISLANDS East Amatuli Island West Amatuli Island Sugarloaf Island Ushagat Island Rocks south of Ushacat Sud Island Nord Island

AFOGNAK-SHUYAK ISLANDS Ban Island-Shuyak Strait Pernosa Bay Marmot Island Sea Otter Island area East side of Shuyak Jest side of Shuyak Latax Rocks-Dark Island

TOTAL

TOTAL

Number of Sea Otters

0 2 0

76 200

29 0

307

18 6 3

121 0

33 26

207

Complete or Visibility Partial Count

Very good Complete ll If II

It II

II

II IJ

Good Complete II

Fair Partial If Complete II Partial II II

II

i o j

~ )Ilt

lt6gt iltj- cgtcgt Jgt

-bull

lt- j )Ilt I

9

I

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I

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J

64

7

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675

6

57

55

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4

50

5

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55

5

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61

62

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5

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4

66

48

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52

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41

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63

5

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13

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19

29

71

20

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51

SEA OTTER COUNTS - KENAI June J970 to January 1971

Carl Divinyi flew aerial surveys of seals along the outside of the

Kenai Peninsula on a monthly basis He recorded all sea otters sighted

on these surveys The surveys did not cover the entire coast line and

the type of aircraft weather conditions and observers varied from one

survey to another Therefore the seperate counts cannot be compared

However when viewed in total they indicate the areas used by sea otters

and the relative abundance of otters in each area

The attached table presents the counts by broad areas

--

SEA

OTT

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CO

UN

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F K

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70

AERIAL SURVEY Prince William Sound

April 1970

Between April 15 and 18 1970 Carl Divinyi and Julius Reynolds fletr an aerial survey of most of Prince William Sound using a Cessna 185 The survey tras primarily to locate seals however sea otter were counted The following is a summary of the survey

I departed Anchorage on April 14 via Alaska Airlines and arrived in Cordova at 600 AM tJeather conditions prevented flying so Julius and I drove the local road network looking for anything in the way of wildlife

April 15 vleather was a little better than yesterday so we decided to try surveying along the east side of the Sound up to Valdez Arm We started the survey at Bomb Point and flew the coast and offshore islands up to Galena Bay

Observations Seal - Scattered small groups with the majority of the animals being in Port Fidalgo There were no noticeable concentrations (50 on up) of seals

Sea Otter - A total of 105 otter were counted with the majority (60) being located in St Matthews Bay The remainder of the animals were concentrated between Red Head and Knmvles Head outside of Port Gravina

Sea Lion- Small scattered groups in Port Fidalgo (60-80 total)

April 16 Weather inclement - no flying

April 17 We flew Hawkins Hinchinbrook Montague and Green Islands Very few seal- many otter and two large concentrations of sea lions (See maps for numbers and locations of seals sea otter and sea lion) We also counted 5 otter around Kayak Island

April 18 Flew those islands from Montague Strait to Icy Bay and from Port Bainbridge to Knight Island Passage (See maps for numbers and locations of seals sea otter and sea lions)

Julius Reynolds continued the survey along the mainland from Knight Island Passage to Esther Island and around Knight Ingot and Eleanor Islands

The area from Esther Passage to Valdez Arm and Culross Perry Lone Naked Peak and Storey Islands were not surveyed

SEA OTTER COUNTED Prince William Sound

April 1970

Number of Area Sea Otter Date

PORT BAINBRIDGE - ICY BAY Bainbridge Island 30 418 Evans Island 14 418 Elrington Island 4 418 Latouche Island 46 418 Whale Bay - Icy Bay 39 418

CHENEGA ISLk~D - MAIN BAY Chenega Island and Dangerous Passage 33 52 Crafton Island 2 52

PORT NELLIE JUAN - ESTHER ISLAND Blackstone Bay 1 52 Culross Island NS

ESTHER PASSAGE - VALDEZ ARM NS

GALENA BAY FISH BAY 103 415

FISH BAY - GRAVINA POINT 1 415

GRAVINA POINT - CORDOVA 1 415

HAWKINS ISLfu~D 1 4l7

HINCHINBROOK ISLfu~ 99 417

EGG ISLAND 2 417

MONTAGUE ISLAND 259 417

GREEN AND LITTLE GREEN ISLANDS 102 4J7

KNIGHT ISLAND 136 52

INGOT ISLAND 6 52

ELEANOR ISLAND 3 52

SMITH ISLAND 4 52

SEAL ISLAND 0 52

APPLEGATE ROCK 1 52

PERRY ISLAND NS

(continued) SEA OTTER COUNTED Prince William Sound

April 1970

Number of Area Sea Otter Date-

LONE ISLAND NS

NAKED ISLAND NS

PEAK ISLAND NS

STOREY ISLAND NS

KAYAK - WINGHAM ISLAND 5 417

TOTAL 892

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PRE-TRANSPLANT SURVEY OF SEA OTTERS Green 1 andMontague 1 -July 171970

On July 17 1970 an aerial survey of sea otters was made in the area

around Green Island and the adjacent coast of Montague Island The purpose

of the survey was to locate concentrations of animals prior to a capturing

operation which began the next day A Dornier twin-engine arrcraft was used

with Schneider Reynolds and Somerville acting as observers The sky was

overcast almost no wind and the sea calm However heavy rain showers intershy

fered with forward visibility and the counting conditions were only fair on

the average Conditions vmiddotere especially poor in Port Chalmers and Zaikof Bay

The count began at 210pm and ended at 345 pm

The numbers and locations of otters are shown on the map In addition

16 sea otters were seen on a superficial look at Constantine Harbor on Hinchshy

in brook Island and a pod of 35 were seen three miles east of Johnstone Pt

Most of the area included in the survey was traveled repeatedly in a

skiff over the next four days Durlng this time the weather became calm and

clear for the first time in several days and the bulk of the sea otters shifted

from the coast of Montague out into the shallow areas between there and Green 1

and to Green and Channel Islands It was obvious that many otters had been missed

Fn the aerial sur~ey and that the population in the area is quite dense A

total of 48 sea otters were captured in three days of netting

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SEA OTTER SURVEY Salisbury Sound to Cape Spencer

August 23-25~ 1970

On August 23 1970 an aerial search for sea otters was made from Salisbury Sound to Torch Bay in the area north of Sitka A Helie Courier aircraft was used with Karl Schneider Alan Courtright and

middotWalt Cunningham serving as observers

The water on the outside coast was too rough for good visibility however it was calmer in the lee of rocks and islands and visibility was fair to good

On the initial survey only two otters were seen These were both in Surge Bay on Yakobi Island While returning after completing the count we located approximately 25 in the same area we saw the first two About 15 of these including at least one pup were in a single pod This illustrates how relatively large numbers of otters can be missed from the air The fact that none were seen in other areas does not mean that established populations do not exist elsewhere

On August 24 and 25 a search of the release area north of Khaz Bay was made by the skiff While flying into Kla) Bay for this seach two otters were seen near the old mine of Chichagof Two miners who have been working there reported that two otters have been there off and on since May

Rough weather and outboard problems restricted tl the search area~

and the effectiveness of the search in that area The following sightings were made

82470 I Adult West of Granite Islands 1 Adult South of Granite Islands

11 0A~dgtzl~4JVflup) middot1 In rocks SE of ranite Islands 82570 I Adult -n Southwest of Gran~te Islands

These sightings represent from four to seven animals

Reports over the last year indicate that sea otters regularly move in and out of Kla~ Bay and occasionally as far in as Sisters Lake bull

The fact that the count in the Khaz Bay area Has less than that made in 1969 does not mean much The animals appeared to be scattered during the present count the search did not include all of the nearby sea otter habitat and survey conditions were less than ideal

The population in Surge Bay appears to be separate and is probably well established Two otters were reported in the same location in January 1966 after only 23 otters had been released near Khaz Bay In 1969 two were seen in the same spot from the ait

While no estimate of the population of sea otters in the area can be made from the available information is evident that sea etters can be found along the entire west coast of Chichagof Island and that concentrations occur near Black Island and the Granite Islands north of Khaz Bay and in Surge Bay on Yakobi Island

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55

HARVEST AND TRANSPLANTS - 1970

Harvest

In May of 1970 a sea otter harvest was conducted on Tanaga Island the middotDelarof Islands and Amchitka Island The numbers to come from each island

and the distribution of the kill around each island were planned in advance Conservative estimates of the population tvere made and the total harvest numbers were based on these estimates The harvest level for Tanaga and the Delarof Islands was set at 15 percent assuming that the next harvest in these areas would be three years later so the rate of harvest would be 5 percent per year Amchitkas harvest level was set at 10 percent assuming an annual harvest and a harvest rate of 10 percent per year

The harvest plan was based on information collected on a June 1968 skiff survey for Tanaga on the 1965 USFWS aerial survey and the 1969 ADFampG aerial

survey for the Delarofs and on 1968 helicopter survey by the USFWS for Amchitka There is good evidence that all the population estimates tvere low especially for the Delarofs

Hunting was done from avo skiffs with two men each Both men shot when possible A 117-foot vessel was used for skinning and living quarters Table 1 shows the schedule of the hunt with the daily harvest numbers

Table 2 presents the projected and actual harvest numbers by area Table 3 presents the numbers killed pelts saved specimen and pelt numbers used and the sex ratio of the kill for each area Figures 1 2 and 3 show the actual numbers and locations in more detail tveather was the main factor causing deviations from the original harvest plan Possible male areas were identified on the 1968 skiff survey on Tanaga An attempt was made to concentrate more pressure on these areas to achieve a more balanced sex ratio This effort is reflected in the sex ratio of the kill A high degree of sexual segregation occurs at this time of year and without specific pressure on male areas we could expect 85 percent females to be taken as occurred on Amchitka With the effort the female percentage was reduced to 65 percent on Tanaga In the Delarofs a male area which had not been identified previously was hit and the effect was much the same

Transplants

Two transplant operations took place in 1970 The first was done in the same manner as the 1968 and 1969 transplants A total of 86 otters tvere captured at Amchitka One aircraft load was shipped out Twenty-nine were released in Oregon and 30 in vashington The animals were held in floating pens at the release sites for several days to recover from the trip Survival appeared to be excellent

The second operation took place in Prince William Sound The Canadian research vessel G B Reed came to the Green Island-Port Chalmers area with tanks built on deck Forty-six sea otters were captured Approximately 44 were alive when the vessel left Prince William Sound however only 14 survived to be released off Vancouver Island

Table 1

April 20

middotMay 1

May 2

May 3

May 4

May 5

May 6

May 7

May 8

May 9

May 10

May 11

May 12

May 13

Schedule of the 1970 harvest

1030 pm depart Homer

Midnight arrive Adak MV Active

Refuel and prepare boat

Arrive off Tanaga at noon

Complete skinning pack hides etc in pm

Delarofs

Spent am getting water at Amchitka

Clean up boat take down shelter pack gear

900 am arrive Amchitka fly to Anchorage

Killed 53

138

88

131

143

53

144

43

79

83

otter

(am)

(pm)

(by mid afternoon)

Table 2 Projected and actual harvest numbers May 1970

Projected Actual Tana~a Island Harvest Harvest

Bumpy Point - Hot Springs Bay 200 191

Hot Springs Bay - Twin Bays 50 50

Twin Bays - Cape Sasmik 50 ) ) 199

Cape Sasmik - Tower 120 )

Tower Tanaga Bay 180 166

TOTAL 600 606

Delarof Islands

Gareloi 20-25 0

Kavalga Ogliuga and Skagul 75-100 125

Ulak and Amatignak 35-55 19

TOTAL 150 144

Amchitka Island saved for

USFWS Counting Units 1 amp 2 100 transplant

3 30 82

4 50 36

5 120 87

TOTAL 300 205

Table 3 Details of sea otters harvested in May 1970

Tanaga Island

Total kill 606

Number of pelts saved 598

Number of small pup pelts discarded 8

Specimen numbers SO 70-4 to SO 70-609 Pelt numbers 3198-3769 and 3771-3796

(Seal 3770 void - damaged)

Approximate sex ratio - 220 males 386 females or approximately 64 percent females

Delarof Islands

Total kill 144

Number of pelts saved 138

Number of pups discarded 6

Specimen numbers SO 70-610 to SO 70-753 Pelt numbers 3797 to 3934

Sex ratio- 44 males 100 females or approximately 69 percent females

Amchitka Island

Total kill 205

Number of pelts saved 194

Number of pups discarded 11

Specimen numbers SO 70-754 to SO 70-958 Pelt numbers 3935 to 4128

Sex ratio- 27 males 178 females or approximately 867 percent females

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A combination of bad weather a lack of time to let the otters recover from their capture and problems with the holding facilities probably caused the high rate of mortality

Summary of Harvests and Transplants

Table 4 summarizes the sea otters removed from Alaskan populations in the middot last 20 years An attempt has been made to remove 300 otters from Amchitka each year since 196 7 Originally this number was set as 10 percent of the population which was conservatively estimated to toal 3000 Recent USFtfS helicopter counts of up to 3927 show that this estimate definitely tvas low In the past the removal of animals was from the southeas tern half of the island The 1970 harvest was purposely concentrated toward the northwestern end in order to spread out the pressure and to learn something about the population in that area

The numbers of animals released in all transplant attempts by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game are presented in Table 5

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30

SEXUAL SEGREGATION IN SEA OTTER POPULATIONS

by Karl Schneider March 1971

Workers have recognized for some time that sea otters tend to segregate by sex Lensink (1962) described this segregation around the southeastern end of Amchitka Island and identified three male areasn and three female areas He speculated that females used areas of more favorable habitat and that younger males were excluded by territorial males scattered throughout the female areas The implication is that male areas contain younger or at least nonterritorial males

Marakov (1965) mentions sexual segregation around Medny Island in the USSRs Commander Islands

Kenyon (1969) gives a more complete description of the sexual segregation around the southeastern end of Amchitka and supports it with quantitative data from harvested animals

Both Lens ink (1962) and Kenyon (1969) w_ere primarily describing hauling grounds used by different sexes While Kenyons harvested animals included otters near shore neither study provided much information on the use of off-shore areas

A knmmiddotlledge of the degree of sexual segregation and the location of male and female areas is important to the management of sea otter populations Harvests must be regulated to avoid putting too much pressure on one segment of the population Then capturing animals for transplants it may be necessary to set nets in specific areas to obtain the desired sex ratio case of a localized kill of otters such as when oil spills occur it is important to know the distribution of sexes to evaluate the importance of kill to the population

In

the

By the same token much can be learned about the distribution of sexes through harvest and capture operations The area from which each sea otter was taken during the harvests of 1967 1968 and 1970 was recorded Because a single hunting party might kill 30 otters over up to 10 miles of coast in a few hours it wasnt possible to record the precise locations Therefore the coast was broken up into areas and the numbers killed in each area ltJere totaled

Figs 1 - 5 show the locations of the areas letters correspond to those in Tables 1 - 4 which present the sex and age composition of animals harvested in each area The ages of the 1968 animals 1vere based on cementum deposition and reproductive condition The other samples were broken down using body size In general all males under 25 lb females under 20 lb and both sexes shorter than 100 em were considered dependent pups Females under 35 lb and 120 em and males under lb and 130 em were considered subadults These criteria probably underestimate the age of some animals Cementum deposition information for the 1967 and 1970 samples is not available yet

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Sexual Segregation - 2 - March 7 1971

Information from the 1967 Amchitka harvest and the 1968 1969 and 1970 Amchitka transplant capturing operations is not comparable and is not presented here However knmv-ledge gained from these operations has contributed to our understanding of sea otter distribution and this knowledge will be used in the following discussion In general this data confirms Kenyons (1969) observations for the Amchitka area although these summer and fall collections indicate a higher percentage of males in female areas than Kenyon reported from his winter and spring samples

Kenyon (1969) states that female areas are more numerous and less discrete than male areas He identified three male hauling grounds and 13 female hauling grounds on the southeastern end of Amchitka Island Recent studies involving netting and collecting animals off~hore indicate that male areas remain discrete off shore Usually these areas are off major points of land and pods of males often form in kelp beds directly off shore from the hauling ground

Female areas on the other hand are not discrete at all Any area that is not a male area can be considered a female area Some areas may be more attractive to females with pups or certain areas may be more favorable for hauling out but more females than males vill be found almost everyvthere except in the discrete male areas Therefore the main problem is to locate the male areas in et population He have no evidence of any shift in the location of male areas over a period of time so once an area is identified the information should remain useful for management purposes for many years Lensink (1962) and Kenyon (1969) correctly identified all of the male areas on southeastern Amchitka Extensive harves and netting have not turned up any new areas

Identification of Hale Areas by Pup Counts

In June 1968 a survey of most of Tanaga and Kanaga Islands was made by skiff to gather information to be used in planning harvests from those islands During this survey females with pups were counted separately from single animals No special effort was put into identifying pups fuen a female obviously had a pup it vas recorded Large pups often were separate from the females and some otters were seen only briefly in vaves kelp or at a distance Therefore many pups probably were missed and the counts should be considered minimal 6 and 7 present the counta by area A similar count was attempted by helicopter around Adak but for various reasons the results are not considered useable

From this skiff survey a number of areas were judged to be possible male areas Usually these were points or more exposed aTeas vhere relatively large numbers of single animals but no pups were seen

Shoal Point Kanaga Pass north of Eddy Rock and possibly Naga Point and Cape Tusik were suspected male areas on Kanaga Island Cape Sudak and Cape Amagalik were considered male areas on Tanaga Island and the area around Hazard Point and Lltnnoy Rock in Kanaga Pass was believed to blend with the Eddy Rock area

The best way to confirm the presence of male areas is to collect animals from these areas Harvests were conducted on Kanaga in October 1968 and on

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Sexual Segregation - 3 - March 1 1971

Tanaga in May 1970 The sex ratios of the animals harvested are compared with the skiff survey pup counts in Table 5 The harvest areas are fairly broad while male areas are usually only a few hundred yards wide As a result a harvest area may include both a male area and portions of female areas to either side

The hunting pressure is seldom even over an area Often a good portion of the kill in an area would come from one group of rocks Therefore the information in Table 5 must be tempered with some knmrledge of how the hunting pressure was distributed

Kanaga Island

Shoal Point - While a few more females than males were taken in this area the percentage of males is considerably higher than one would expect in a female area Because weather vas less than ideal much of the hunting pressure ~vas directed to areas other than Shoal Point Most of the females probably came from these areas Therefore Shoal Point is considered to be a male area$

Naga Poin~- There is no strong evidence that this is a male area While no pups were recorded on the survey the total number of otters counted was not high Fairly extesive hunting of the area under good conditions yielded a sex ratio that would be typical of a female area Therefore it is assumed that there is no male area near Naga Point

Cape Tusik - Cape Tusik was suspected as a male area more because of the nature of the area than arry counts The count included areas to both sides of the Cape which could contain females Very little of the hunting was done at the Cape Therefore judgement on this area will have to wait until more information is available Extensive hunting indicates that no male areas exist east of Cape Tusik to Cape Chlanak

Kanaga Pass - From the count it appears that a male area exists in the Pass however the harvest area extended to Hive Rock and included a large portion of female area (Fig 6) The area is too broad as is shown in Table 5 gtvhere the number of pups counted for the whole area is relatively high The results of the harvest confirm the conclusions drawn from the count While the sex ratio for the entire area is roughly equal most of the males came from Kanaga Pass Some females without pups came from that general area but the majority of the females came from the area northeast of the Pass

The Tanaga harvest showed the male areas more distinctly for several reasons The total harvest was higher the entire count area was covered in the harvest and the hunting pressure was recorded more precisely Also as will be shown later sexual segregation is more pronounced in the spring

Cape Sudak - Most of the hunting pressure was concentrated on the tip of the Cape Some pressure was exerted on the area back to~mrd Pendant Point which is a female area as shmvn by the count The high percentage of males harvested from this area clearly confirms that the tip of the Cape is a male area

Tab

le

5

Com

pari

son

of

Pup

s C

ou

nte

d w

ith

S

ex R

atio

o

f H

arv

est

Jun

e

1968

A

rea

Su

rvey

O

cto

ber

19

68 H

arv

est

Pu

ps

10

0

Pu

ps

10

0

Ad

ult

s T

ota

l KA

NAGA

IS

LAN

D

Ind

ep

Ott

er

Ind

ep

Ott

er

M

M

F

Rou

nd

Hea

d-S

ho

al P

oin

t 0

85

3

45

6

55

4

71

5

29

Nag

a P

oin

t-K

anag

a B

ay

66

1

64

middot

18

5

81

5

20

5

79

5

Cap

e C

hla

nak

-Cap

e T

usi

k

51

1

93

2

40

7

60

3

53

6

47

Edd

y R

ock

-Hiv

e R

ock

61

3

4

35

7

64

3

47

1

52

9

Jun

e

1968

A

rea

Su

rvey

H

ay

1970

Har

ves

t

Pu

ps

10

0

Pu

ps

10

0

Ad

ult

s T

ota

l TA

NA

GA

IS

LAN

D

Ind

ep

Ott

er ~ter

M

F

M

F

Cap

e S

ud

ak-P

end

ant

Po

int

0 4

4

60

6

39

4

66

4

33

6

Bar

nes

P

oin

t-T

run

k P

oin

t 7

9

59

3

3

96

7

83

9

17

Tru

nk

Po

int-

Tw

in B

ays

26

4

2

77

9

23

1

40

8

60

D

rin

Bay

s-S

ou

th B

ay

96

1

11

7

3

92

7

14

3

85

7

So

uth

Bay

-Har

em R

ock

14

8

31

6

7

middot93

3

91

9

09

Las

h

Bay

-In

fern

o

Ree

f 1

5

73

0

27

0

74

6

25

4

Har

em R

ock

-Ku

lak

Po

int

0 2

7

64

4

35

6

68

4

31

6

Ku

lak

Po

int-

SE

B

igh

t 6

9

82

1

32

8

68

1

51

8

49

Sexual Segregation - 4 - March~ 1971

Annoy RockHazard Point - The harvest did not confirm this area as a male area However the weather Vas marginal and this area was too rough to hunt Therefore the otters taken in the harvest ~vere from either side of the Point and almost none were taken from the portion that is suspected to be a male area Actually Kanaga Pass is narrow and shallmv Otters feed in all parts of the Pass when weather and tide rips permit This could be considered a single male a~ea that serves both islands

Cape Amagalik - Two harvest areas overlap this area Lash Bay-Inferno Reef and Harem Rock-Kulak Point The kill from both areas strongly reflects the presence of a male area Nost of the hunting pressure was concentrated on Inferno and this is probably the main male area

As shown above vle 1vere able to identify four major male areas from a relatively superficial pup count No male areas were located during the harvests that had not already been identified from the count ~mile more detailed observations including field identification of males would be desirable the pup counting technique appears to be an acceptable method

Identification of

Because male areas are usually on v1ell exposed points it is possible to pick likely areas from a chart and then confirm these areas by collecting animals This method avoids the expense time and danger involved in surface surveys of remote areas Actual collection of animals also provides the most concrete proof of the nature of an area

The follmving is a description of the sex composition of areas for which no skiff count data was available

Delarof Islands - No attempt 1vas made to predict the location of male areas in the Delarof Islands This is a difficult area to work in Ogliuga Skagul and a portion of Ulak Island v1ere hunted on Nay 9 1970 The hunting effort in the afternoon was concentrated around Tag Island and the rocks south of Skagul Island While the entire kill is lumped together most of the males bullJere taken in the afternoon The percentage of males taken for the day is higher than would be expected if only female areas were hunted Most likely there is a male area near Tag Island

Adak Island - Portions of Adak gtvere hunted in 196 7 and 1968 No attempt was made to identify male areas before or during either harvest The harvest figures indicate that no male areas have been hunted The Bay of Islands is a classical female area The results of extensive hunting in the Bay and around Eddy Island indicate that there is definitely no male area there The percentage of males taken between Mid Point and Blind Point vms higher than might normally be expected in a female area however the scarcity of subadult males indicates that no male area exists within this area A large congregation of otters is often seen near Finger Shoals This could be a male area but Ye have no information from there If this is a male area a few stray males from there could account for the slightly higher number of males in the kill

Sexual Segregation - 5 - March 1971

Amchitka Island - While much work has been done on Amchitka Island almost all of the effort has been concentrated on the southeastern end from Crmvn Reefer Point to a fegtv miles northvest of Rifle Range Point During the 1970 harvest an attempt was made to distribute the harvest over previously unstudied areas Two areas were suspected to be male areas These were the rocks off Chitka Point and either Bird Rock or Aleut Point Heather prevented us from hunting the area around Bird Rock and Aleut Point

The sex ratio of the kill does not indicate a male area near Chitka Point however heavy waves prevented hunting off the point Therefore a male area could be there but because no animals were killed there the kill figures would not reflect it

Composition of Female Areas

Kenyon (1969) found that 93 percent of the adult otters and 63 percent of the juvenile otters in female areas were females The May samples from the 1970 harvest agree with this The mean percent of adult females in female areas was 93 percent The mean percent of subadults not including pups was 73 percent Hmmiddot1ever in the September and October samples only 77 percent of the a-dults in female areas were female About 86 percent of the subadults were female

The seasonal difference in the sex ratio of adults is consistant for all samples These seasonal changes are most likely due to adult males moving into female areas to look for estrous females During late winter and spring breeding activity is at the lmmiddot1est point of the year and fev1er females are entering estrus In fall the number of estrous animals is at its peak and presumably more sexually mature males move into the female areas to breed It is interesting to note that there were about three times as many males in female areas in fall as in winter and spring Information from female reproductive tracts indicates that approximately three times as many females would be in estrus at any given time in fall It appears that the number of adult males in a female area is directly proportional to the number of estrous females From the data collected there appear to be 15 to 20 males for each female -vlith enlarged follicles at any given time hmmiddotrever hunter bias might influence this figure

Ages have been estimated for otters harvested in 1968 on the basis of cementum layering These age data indicate that virtually all of the males in discrete female areas are either pups (or very young subadults) or are 7 years old or older No males between the ages of 2 and 6 Here found This strongly implies some form of territorial behavior among actively breeding males Fighting among males has not been observed ofteci hmvever a mature male at the Tacoma Zoo became intolerant of a subadult male in the presence of females They frequently 11 fought 1 although neither seemed to attempt to injure the other Finally the young male had to be removed Under imiddotJild conditions the young male might have moved out of the area without repeated physical encounters

Kenyon (1969) described the breeding behavior of sea otters His description indicated that males patrolled an area and two to four males may pass a given point during a 3 or 4 hour period Vandevere (1970) observed some males

Sexual Segregation - 6 - March~ 1971

apparently defending a territory in California but from his observations t

it appeared that successful breeding males were more mobile Perhaps established breeding males are more tolerant of each other More likely they maintain a separation from each other which limits the number in an area In this way a number of males might be patrolling the sa~e area rather than each establishing a geographical territory This might be advantageous where female concentrations shift from place to place as weather conditions change

Why does the number of adult males in female areas decline when the number of estrous females declines The limiting factor may be competition for estrous females rather than territory size With fewer receptive females competition lvould be greater causing some animals to leave the area Another possible reason may be that males may have a seasonal fluctuation in fertility Lensink (1962) found that all adult males produced sperm at all seasons Limited studies since then support this however it is possible that most of these males were collected in or near female areas Also while a male may produce sperm at all seasons there may be less production at certain times of year

A contributing factor may be that males are capable of feeding in more exposed areas than pregnant females or females with pups With a reduced attraction to the female areas either middotbecause of fewer receptive females or a reduced fertility in the male the male may find it easier to obtain food away from the crowded female areas where competition for food is greatest

The fact that the number of males does appear proportional to the number of estrous females indicates that competition for these females may play an important part in regulating the number of males in female areas

The numbers of subadults in the samples is relatively low so fluctuations in sex ratio may be due to sampling errors However the greater number of subadult males found in female areas in winter and spring may be a true increase made possible by the reduction in the number of breeding males A subadul t male would have fewer encounters -vli th breeding males

There are areas within what would normally be considered female areas where subadult males tend to congregate particularly during the winter These are usually areas v7ith a ma-r-ginal food supply that are not heavily used by females Because there are fewmature females in these areas there are also very few breeding males The subadult males using these areas seem to be transients that find less pressure from breeding males there These areas are used less in summer probably because calrrer weather allows feeding farther off-shore reducing competition for food in the male areas

Examples of areas within female areas that are used by subadult males are Constantine Harbor on Amchitka and possibly Finger Bay on Adak

within Female

In the course of the harvest on Tanags in tIay of 1970 a large number of pregnant females tvith large fetuses were collected at one time Most of

Sexual Segregation - 7 - March 1971

these came from Tidgituk Island This raised the possibility that females may segregate according to reproductive condition Table 6 presents the numbers of females in each stage of the reproductive cycle by area on Tanaga Island More pregnant animals were collected in the South Bay area and within that area most of the females with fetuses weighing over 100 g were collected near Tidgiktuk Island This concentration was evident in the pup counts made in June 1968 when a very high percentage of pups was observed in the same area (Table 5) Host females ~vith large fetuses in May would pup by June

The data do not reveal any other areas of this type on Tanaga Harakov (1965) mentioned a bay on Hedny Island that tvas preferred by pregnant females and females with pups Obviously all females do not go to a specific area to pup but a female vi th a large fetus or small pup probably has more difficulty in exposed areas and seeks more protected areas This tendency can be seen by the casual observer Single animals are more likely to venture off-shore and remain in rougher waters In some areas such as Crown Reefer Point on Amchitka there is a male area near the tip of a point Males congregate directly off-shore However to either side of these male congregations there are females that are not accompanied by a pup and probab do not have large fetuses In this way we may find segregation in a single kelp bed off a point If we set a net near the outer margin of the kelp we will catch almost all males Ho~Vever if we set a net to the side of the bed and closer tmiddoto shore we will catch mostly single females Nets set in a nearby bay will catch more females with pups

Table 6 shoHs that an unusually high number of resorptions were found between Cape Sudak and THin Bays This is probably a reflection of poor physical condition of the otters in that area due to food Other data indicate a continuous decline in body condition from the east side of Adak to Kanaga Pass There is some evidence that condition improves west of Kanaga Pass The animals in the Kanaga Pass vicinity are probably in poorer condition than any of the other populations sampled Therefore the high incidence of resorptions in this area should not be considered an example of segregation within the population

ition of Male Areas

As mentioned earlier the harvest areas ivere broader than any male area Harvest areas containing male areas also contained portions of female areas Our identification of the location at rhich each animal vas collected is not precise enough to determine the exact composition of these rnale areas Kenyons (1969) data from winter and early spring harvests are more precise His data indicate that at that time of year 98 percent of the adults and 80 percent of the ju~veniles using male areas were males Of 128 males in male areas 100 gtvere adults and 28 were juveniles

Experience from netting in the summer and harvesting in fall indicates that the percentage of males remains very high all year While adult females may be very close to male areas it appears unlikely that a significant number regularly use these areas at any time year nile adult males enter female areas regularly and in predictable numbers for a specific purpose females probably ~nter male areas only occasionally in stray movements

le 6

R

epro

du

ctiv

e S

tag

e o

f S

exu

ally

Mat

ure

S

ea O

tters

Har

ves

ted

on

Tan

aga

Isla

nd

-by

Are

a -

May

1

97

0

Un

imp

lan

ted

Im

pla

nte

d

Pre

gn

ant

Pre

gp

ant

Fet

us

Wei

gh

t C

lass

A

rea

No

np

reg

nan

t N

o

No

1

2 3

4 5

Res

orp

tio

n

To

tal

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dak

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dan

t P

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t 12

7

26

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0

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int-

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t S

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7 24

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s 7

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cket

s in

dic

ate

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ver

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as

Sexual Segregation - 8 - March 1971

Juveniles of both sexes probably move more randomly than adults lihile definite sexual segregation exists among younger animals it is not as pronounced as in adults Because subadult males are tolerated less by breeding males they are more strictly confined to small areas than females of the same age This has been demonstrated in fall harvests where extensive hunting over a large area will turn up only an occasional subadult male Then ~v-hen the hunters move to a definite male area large numbers of subadult males are taken in a very short time and in a very small area

While Kenyons data indicate that 22 percent the males in male areas are subadults our experience indicates that this percentage may be quite a bit higher at least in fall Such a seasonal change in the age composition of males sounds reasonable We have demonstrated that adult males move into female areas in greater numbers in the fall perhaps tripling their numbers in these areas At the same time fewer subadult males are found in the female areas Presumably the adult males are coming from male areas and the subadults driven from the female areas move to the same male areas The result vmuld be a higher percentage of young males in male areas in fall than in spring

The total number of males of all ages in female areas doubles in fall About 13 percent of all animals taken in these areas in spring were ~ales while 25 percent ta-ken in fall were males This suggests that the total number of males in male areas declines in fall There are more adults leaving than subadults entering the area

Sex Ratio

With the sexes segregating and with the degree segregation changing seasonally and betmiddotween age groups it is extremely difficult to get a completely random harvest As a result we have not been able to determine the sex ratio of any population as a vhole Hith female areas being larger and in more protected areas where hunting is easier there is a strong tendency to take more females than males particularly in sp-ring ~Je have shown that through a concentrated effort the percentage of males harvested can be increased however even then the lowest percentage of females taken in recent harvests ivas 62 percent on Kanaga Island in October 1968 Similar results occur in capturing operations for transplants

On all islands studied there are fewer male areas thltm female areas The male areas are very small usually only a few hundred yards wide The male feeding areas off male areas are also very narrow although they may extend farther off-shore In short that portion of the total available sea otter habitat included in male areas is very smalL

All evidence indicates that there are more females in the population than males While the percentage of is probably greater than that indicated by most harvest statistics it appears that at least 60 perc2nt of the sea otters in the populations studied are females

There are probably several factors contributing to this uneven sex ratio First our reproductive data indicate that 56 percent of the pups are females at birth Secondly Kenyon (1969) has found a higher mortality rate among

Sexual Segregation - 9 - March 1971

newly weaned males There is also some evidence that males might not live as long as females These factors could easily account for the preponderance of females in the population

It is possible that the higher rate of mortality in juvenile males is in part due to the breeding behavior of adult males and the resulting sexual segregation Juvenile males tend to be restricted to male areas or areas of marginal feeding habitat where there is little competition from breeding males During the ltvinter the feeding activity of young males would tend to be confined to small areas close to shore in male areas Therefore they may be subjected to greater competition for food than females of the same age which have a broader area in ltvhich to feed Kenyon (1969) found that subadult males vere less hardy in captivity than subadult females Therefore we can not adequately evaluate the influence of segregation on mortality

The sex ratio probably varies from one population to the next Very little juvenile mortality occurs in expanding populations eliminating that source of sexual selection However there is some evidence that males are more numerous among the first animals to repopulate a new area While a high percentage of males might be found on a newly repopulated island the loss of these males tvould alter the sex ratio of the parent population even though juvenile mortality might not yet be a significant factor

If the nThuber of breeding males is regulated by the number of estrous females an even sex ratio might produce a surplus of sexually mature males This surplus would not be beneficial to the population and could be detrimental This would permit the situation of an unbalanced sex ratio to evolve

The segregation of sexes and ages and the composition of sea otter populations is complex Unless we can understand the situation we cannot fully evaluate the effects of mortality or habitat changes

REPRODUCTION IN THE FElIALE SEA OTTER

by Karl Schneider Narch 1971

A total of 1358 sea otter reproductive tracts collected between 1967 and 1970 have been examined Ovaries were sliced with a razor blade and examined macroscopically Uteri Here macroscopically examined both internally and Each tract vas classified as to its stage in the reproductive cycle

The follovJing criteria were used in classification

Nulliparou~- Horns of uterus small thin and smooth no corpora lutea or corpora albicantia although there is occasionally evidence of a past ovulation but no evidence of implantation

Nultiparous - Uterus thicker corpora lutea andor corpora albicantia present Includes primiparous animals (these are not readily separated from multiparous animals)

Anestrus - Largest follicle under 3 5 rnm no corpus luteum

Proestrus - Follicles over 35 mm

Es - Follicles approximately 10 rrm

Implan~elt_ Pregnant_ - Visible s~velling in uterine horn usually with fetal membranes embryo or fetus visible Corpus luteum usually over 10 mm

- Uterine horn sti11 noticeab enlarged fresh roughshy~=-=--=~

al scar newly formed corpus albicans with some luteal tissue remaining

The appearance of the uterus ~ras used to confirm the classification The thickness arrd of the horns thickness and consistency of the uterine walls coloration of the of the horns and condition of the rugae change with each stage In most cases it is possible to guess the condition of the tract by a superficial examination of its exterior This appearance was used only to confirm what was indicated by structures in the ovary hmvever

Table 1 smnmarizes the reproductive condition of the sexually mature females in the that are enough for comparison throughout the year In the following discussion reported by Sinha et (1966) and Kenyon (1969) are included 7here possible The January and Harch are from these studies

Tab

le

l Re

p iv

e co

nd

itio

n o

f se

xual

ly m

atur

e se

a o

tters

_

IF

etus

Wei

--------------+

__

_

ln~a

ctive

ov

arie

s 1

Act

ive ovari~

ht C

Jas

s

jmiddot

I P

ost

-I P

roes

tru

s U

nim

pl

lmpl

D

ates

L

ocat

ioQ

1A

nest

rus

Part

um

Res

orpt

ion

l +

E

stru

s P

reo

Pr

eQ

2 3

lJ

S

~

4-8

1970

T

anag

a 1

51

41

0

10

104

17

8 10

33

30

4 (1

68)

(1

35)

(3

3)

(3 3

) (2

89)

(34~

(5

6)

(27

) (3

3)

(11

4)(

10

9)

May

9

19

70

Del

arof

Is

18

14

1

4 13

27

10

0

3 6

8 77

4

) (1

82)

( l

3)

(5

(16

9)

( o

) (1

30

) (3

9)

(78

)(1

04

)

10-1

219

70 A

mch

ltka

I

34

18

3 8

27

48

5 6

2 23

12

13

8

( 0

(24

6)

(13

0)

(22

) (5

8)

(19

6)

(34

8)

(36

) (4

3)

(14

) (1

67)

(8

7)

~lune

23

middotmiddot ~middot~~middot~Amchitka

1

17

2 1

2 4

18

1 2

3 5

7 44

A

ugus

t 13

196

8 (3

86)

(4

5)

(23

) (4

5)

(90

) (4

09)

(2

3)

(45

) (6

8)

(11

3)(

15

8)

July

6-

~dgtAmchitka

1

17

1 0

4 14

10

0

1 1

2 6

46

Aug

ust

819

69

(36

9)

( 2

0 2

) (8

6)

(30

4)

(21

9)

(22

) (2

2)

(43

)(1

32

)

Sep

t 1

0-17

A

dak

72

5 0

16

31

40

6 6

8 9

11

164

1967

(4

39

) ( 3

1)

(98

) ( 1

89)

(

4)

(3 7

) (3

7)

(49

) (5

5)

(6

7)

Sep

t

25

-A

mch

itk

a l

55

6

0 18

19

17

4

0 0

0 3

115

OcL

6

19

67

(4

7~8)

(5

2)

(15

7)

(16

5)

(14

8)

(35

) (8

7)

(26

)

Oct

o 1

2-1

5

Kan

aga

I

58

6 1

13

31

31

4 4

7 11

5

140

1968

(4

13)

(4

3)

(o 7

) (9

3)

(22

2)

(22

2)

(29

) (2

9)

(50

) (7

8)

(36

)

Oct

18

-20

A

dak

l

46

6 0

1 24

13

2

3 0

4 4

100

1968

(L

16 0

) ( 6

0)

(11

0)

(24

0)

(13

0)

(20

) (3

0)

0)

(40

)

Num

bers

in

re

nth

esis

are

pe

rcen

t se

xu

ally

mat

ure

fem

ales

F

etus

wei

ght

clas

s 1

=

0-l

g

2 =

1-l

Og

3

= 1

0-lO

Og

4

= 1

00-lO

OO

g

5 =

Tra

nsp

lan

t m

ort

alit

ies

(all

o

ther

sam

ples

fr

om

har

ves

ts)

Reproduction - 2 - March 1971

There may be some problems associated with comparing samples taken from different islands and in different years but allowing for sampling errors the data for the most part fit a definite pattern indicating that the annual cycle remains fairly constant The one outstanding exception is the number of pregnant animals in the t1m summer samples In 1968 there were many more implanted pregnancies than unimplanted pregnancies and in 1969 the situation was reversed hmvever the total number pregnant was almost identical These are the t s being relatively small collected over a longer of time and consisting of transplant mortalities An average of the two samples fits the pattern established by the other samples Relatively large samples are necessary because all stages of reproduction are found at all times of the year The period from November to early January is not represented hmmiddotJever things are changing so rapidly during that period that any information from that time may be confusing unless a large sample was collected in a very short time

The information in Table 1 is shmvn graphically vJith Kenyons (1969) winter information in Figs 1 and 2 It has often been demonstrated that sea otters breed and pup at all times of the year A number of observers have noticed that roore pups are born in the spring aJd sumrrter than at other times and there have been conflicting reports about breeding times Kenyon (1969) was the first to demonstrate seasonal changes in pregnancy rates but only his January and Yarch sallples gtvere large enough to be at all reliable

Figs 1 and 2 demonstrate that there are cons le secsonal in the numbers of animals in each s reproductive cycle There is a definite period of increased bree activity and a definite period of increased pupping Hmmiddotlever ~ some anii1als are in each s at all times of the year and these periods of increased breeding and pupping do not have distinct boundaries

Each curve is influenced by two factors so it is difficult to isolate the magnitude of any single factoc For example the of implanted pregnancies vill increase ss blastocysts and decrease as births occur The birth rate may be increas but if the number of implantations increases at a greater rate the implanted curve will still rise The animal remains in the anestrus implanted pregnant~ and unimplanted pregnant categories for a relatively long time and there may be considerable carry-over from one sample to the next Because the animal a relatively short time in the proestrus-estrus and post partum categories there is little if any carry-middotover from one sample to the next and the changes betgtmiddotTeen are more likely to be absolute changes The mean fetus weight class prob does not mean much because it does not take the actual number of animals in each class into account ~hen there are many implanted pregnancies ard the mean lmiddotTeigh t class is lovJ there actually may be more Class 5 fetuses in the population than vhen there are fetv irr1planted pregnancies and the mean middotleight class is high Therefore the actual percentages of all sexually mature females middotlith Class 1 ald Class 5 fetuses have been plotted in 3 These curves represent the actual number of fetuses in the population Because tha sample sizes become quite small vhen the fetuses are divi(12d into classes and the surruner sample is believ2d

2

~ _s ~

~

c U)

~-1

r

~J

lt

c])

Reproduction - 3 - Narch 1971

to be biased tvo curves for each class are shmvn The solid curves approximate the data while the broken curves represent subjective adjustments to correct for sampling errors

cussion of the Annual

The number of animals entering estrus rises in the fall At this time the pregnancy rate is at its lowest point and a high percentage of anestrous (nonpregnant) animals are in the population Breeding activity increases and at this time the number of sexually mature males in 1female areas increases

The number of unimplanted pregnancies begins to rise sharply in October and November At the smne time the nlli~ber of anestrous animals drops as these animals enter estrus breed and become pregnant Thj_s is the peak breeding season

The birth rate is at its louest this period as is shovm by the lmv number of post-partum a1imals and the lovr number of large fetuses

By late November and December the pregnant rate is increas even though the birth rate is slightly This is due to ne7

blastocysts implanting as is shown the increase in Class 1 fetuses

By January the period of t breedLng activity has declined and the number of unimplanted animals ceaches a peak as the number of blastocysts implanting equals the number of The lanted pregnancy rate then begins to declLne as fevJer animals breed and more implant The greater rate of antation is reflected in a rise in the Class 1 fetuses as well as an overall increas in the er of lanted pregnancies However the rate of implantation is partially obscured by an increased birth rate removing animals from the total of lanted pregnancies This increase in the birth rate is shmm in the rise in post-partum females and a rise in Class 5 fetuses

This condition of a low rate of breeding high rate of implantation and increasing rate of birth lasts from February to Hay

At this point He come to the least reliable data The sumner are smaller gtJere collected over a period of seven weeks and were from animals that died as the result of capture and handling

Therefore tvhile the data shaH that the post-partum rate decreases sh indicating a reduction in the birth rate~ the number of Class 5 fetuses

a peak indicating a very high birth rate The true birth rate is probably some1middothere in betigtJeen Females 1-ith very young pups (hence are post-partum) are vlary and probably less likely to be caught in nets Females with fetuses appear to be more likely to from handling In several cases~ twisting of the reproductive tract by a fetus vJas the actual cause of death Anestrous females (including post partum) on the other hand are less likely to die

Reproduction - 4 - March 1971

There is no question that the birth rate remains high during June and July Kenyons (1969) sum11er sample is limited but shows a large number of Class 5 fetuses Again the sample is biased in favor of pregnampIt animals but not to large fetuses only His field counts as vJell as observations by most other observers indicate an increase in the nUIlber of dependent young througbout the surnmer The number of Lmted pregnancies declines throughout the suTimer tvhile the unimplanted pregnancy rate declines at a slmver rate and breeding remains fairly constant at a lm-v level This indicates that through June there are more implantations than conceptions hmvever the birth rate is substantially greater than the implantation rate By late July and August there are relatively fe1r implantations taking place as shmm by a leveling of the unimplanted pregnant rate and a lotr number of Class 1 fetuses The number of Class 5 fetuses declines as a result of births during June July and August

Therefore the season of greatest pupping may be quite broad running from April through August possibly with the peak occurring in late May or June Until a better summer sample is available tve cannot pin dmm the peak of this pupping season with certainty (See Fetal GrmmiddotJth Rate for further discussion on breeding and pupp peaks)

If the information were precise we could compare subtracting numbers of pups born and numbers of conceptions to deterrnine the actual percent of ferrales breeding implantLng ltmd pupping in each month Unfortunately the data are not Each curve is influenced by more than one factor and ue do not knmmiddotr for sure the magnitude of any of these factors Hmmiddotrever by comparing some of the major samples e may be able to get a rough idea of how much higher the birth rate is at one time of year than another

The combined Mty and the combined September and October samples provide the best comparison Both are large samples One occurs near the peak of pupp and a lmv point in breeding and the other is near the low point in pupping and the peak of b

The number of Height Class 5 fetuses~ post-partum femaless and proestrousshyestrous females come closest to represent a particular event The time spent in each category is comparatively short

There were approximately three times as many Class 5 fetuses in Hay thanmiddot in Septerqber-October There Here also about three times as wany postshypartum females in May In tember-October there were two and a half to three times as many proestrous-estrous animals as in Hay It ~middotwuld appear that the peaks of breeding and pupping are roughly three times as high as the lowest points

At the lmmiddot12st period of pupping 2 to 3 percent of the females have Class 5 fetuses and should pup middotJithin a month If vm assume the post-partum stage lasts L 5 to 2 months (see Recovery of the Uterus) it also appears that 2 to 3 percent pup in the ltwrest months of pupping

Reproduction - 5 - March 1971

Similarly about 3 percent of the females have enlarged follicles during the lmvest period of breeding He do not knmv how long this period lasts hovever

At the peak of pupping about 10 of the females have Class 5 fetuses and should pup in the next month About 15 are post partum indicating about 8 births per 100 females in the preceding month

These percentages are prob2bly slightly high because of hunter bias avoiding females with pups It appeatmiddots that at least 2 to 3 percent of the mature females breed and 2 or 3 percent have pups in any month of the year In the peak breedingmonths such as September and October perhaps 7 to 10 percent breed and similarly in peak pupping months sud1 as Nay 7 to 10 percent of the mature females pup

Gestation Period

Barabash-Nikiforov (1947) listed the gestation period of 8 to 9 months This apparently was based on the case where a fentele mated in captivity as reported by i-Ialkovich (19 37) According to Barabash-Nikiforov a pup was born 8 months later although it died While it was fully formed~ it may have been premature

A number of births have occurred in captive animiddotmals held by the of Fish and Game in recent years In all cases the pu-ps ~-ere sc-middot1all usually less tha1 1600 g but still than the smallest pups found in the -rild All died after birth

The gestation period of 9 months has been repeated in the literature but apparently these statemsnts are based on Barabash~Nikiforol (19lf7)

Lensink (1962) also estimated a gestation period of 8-9 months based on field observations He felt that the peak of breeding was in August or September and the of pupping ~Alas in l-1arch or April

It has been demonstrated that the gestation of sea otters a relatively long period of tation (Sinha 1966 Kenyon 1969 and present study) Therefore any attempt to estimate the length of gestation by examinatioD of tracts must take both the unimp1anted and Janted periods into accour1t

Kenyon (1969) presented an estimate of the implanted period by assuming that the cube roots of fetus weights fall in a straight line after the embryo is established (Hugget and Hiddas 1951) and by that the European otter and Aluerican river otter would have similar fetal growth rates By plotting tenn fetus on a line established by the European otter he estimated an gestation of about 120 days not including the period for establishment of the ewbryo

Kenyon (1969) then an estimate made by Dr D G Chapman using the method of Hugget and ~-Jiddas (1951) v7here a regression line is fitted

Reproduction - 6 - March 1971

to the cube roots of the raeail of the mean weights of fetuses in each of five vreight clas3es plotted t the mean time of year

From the regression coefficient ob he estimated an implanted period of 154 days By taking the mean of the pregnant animals that are unimplanted pregnant estimated the tmimplanted period as about 75 months 1Ulmv-ing a half month for establishment the embryo he estimates a total gestation of 12-13 months

The data used by Chapman included a January-February sample of 26 a March-April sample of 49 and a Nay-August of only 9 The surtmler sample is more biased than the others and quite small yet the estimate relies heavily on this sa~ple when fetus size is the greatest

Table 2 presents a neTw- estimate of the length of the implanted period using the method used by Chapman but including data from the present study in addition to Chapmans data Table 3 presents a similar estimate of the length of the unimplanted period

The following is a comparison of the two estimates

Unimplanted Period 230 days 66 days Establishment of Embryo 15 15 days Implanted Period 154_~~Yrgt~ _73 d~s

Total Gestation Period 399 d2ys 154 days or about 13 months 5 months

Obviously there is some problem tiith the the technique or both A similar calculation mcde before the Hay sample had been collected produced an estimate of 109 for the implanted period and about 8 months for the total ation period

This method of calculation assur1es that there are definite peaks in breeding and pupping If breeding and pupping occurred evenly throughout the year the average fetus size 1vould remain constant thruughout the year and the technique would not tvork An ideal situation v10uld be when all individuals breed and pup 1vi thin very short periods Sea otters do have but are not well defined Very large s v-ould be required in such a case because there are ahvays fetuses of all sizes and the periods of maximum breeding are broad

Obviously Chapmans sample vas inadequate llthile the number of fetuses available for the latest estimate seems large and are well distributed throughout the year the a_ctual nu~ober in any cne fetus vJeight class at any one time of year is small A look at Table 1 shows that there is little consistency in many of the classes (Fig 3 shmvs that Class 1 and 5 fetuses do fit a pattern)

Table 2 Estimated Length of Period

Percent of Fetuses in each Height Class

Time of Collection After January

1 month 31 15 15 38 0

3 months 18 12 22 35 12

5 months 18 8 8 36 30

7 months 3 8 16 29 45

9 months 20 7 10 40 23

10 months

14 18 12 33 23

---~--

r1ean Time in Nonths After January 50 57 53 58 70

Cube Root of Umveighted Mean 063 17 36 7lf lllf

Specific Grmth Velocity 0169day EstirGated Implanted Period = 73 days

Table 3 Length of Unimplanted Period

Months After Unimplanted Unimplanted Percent January Pregnant Pregnant Unimplanted

1 month 35 26 58

3 months 49 49 so

5 months 128 179 42

7 months 22 37 37

9 months 50 57 47

10 months 55 44 56

Unweighted

48

Reproduction - 7 - March 1971

All of the estimates very heavily on su111mer samples 1vhen the fetuses are largest These sauples are knmm to be biased probably in favor of larger fetuses There are more fetuses at this time of year but it is exaggerated in these SCilllples

The latest estimate is not necess better than Chapmans even though it uses more and larger samples The velocity is twice as high as that calculated for fur seal and a 5 month gestation period is not consistent with other information available

This technique for estimating the lanted gestation period is not capable of providing a reliable estimate with the available data Therefore the 12 to 13 month estimate and any calculations based on it are meaningless

Much of the discrepancy lies in the estimate of the unimplanted period The present estimate that 48 percent of all pregnancies throughout the year are unimplanted based on an even distribution of relatively large samples and is probably more accurate than Chaprnan 1 s estimate of 60 percent

There are several other ways of guessing at the length of gestation Kenyons (1969) estillate of an lanted period of 4 months using the cube roots of term fetus weightscannot be relied upon entirely but it may be closer than the other estimates If gtmiddotJe estimate the uninplanted period from this using the 48 percent ted factor and allow for the establishment of the embryo we get a total gestation of about 85 months

Lensinks (1962) method of detennining the and peak pupping periods and taking the time between the peaks as the total gestation period has some merit TJnile Lens ink tried to determine these peaks by field observations we can determine them more accurately using the condition of reproductive tracts

Lensink felt that the peak of breeding middotlas in August or September The data in Fig 2 indicate that he may seen the beginning of the period of est breeding activity but that the probably in October or possibly November Lensink estimated period in larch or April Again the data indicate that he v1as s the beginning of the period of greatest pupping but the actual peak is probably in June or July At this time the number of Class 5 fetuses is greatest The nucJber of post-partum females probably and the number of implanted pregnant females is dropping from its If the gestation period of

11 11the average animal lasts from October or November to June or July we get a period of 8 or 9 months tThile Lensink 1 s was early 1 the duration agrees vith the present data

The peak of implantation when the greatest number of blastocysts are implanting can also be roughly estimated from 1 and 3 In February the number of unimplanted middot is dropping and the number of implanted pregnancies at this time th2 number of Class 1 fetuses probably reaches a

Reproduction - 8 - March 1971

Therefore it appears that the unimplanted period lasts 4 to 5 months and the implanted period from 4 to 5 months 1vi th a total gestation period of 8 to 9 months It should be recognized that these estimates are based on general trends and not distinct Therefore they are approximate However the relative length of the unimp and implanted p2riods agrees with the percentages noted previous and this estimate agrees vlith the one based on Kenyons (1969) comparison vdth other species of otters

If the gestation period ~Jere 12 months the percentage of pregnant animals would remain constant throughout the year If it vJere only slightly longer or slightly shorter there would be only a slight fluctuation unless there were a distinct breeding period Fig 4 indicates a substantial fluctuation indicating a gestation period substantially shorter or longer than 12 months

The peak of pregnancy occurs in February after the period of t breeding activi The luH point on the pregnancy curve occurs after those animals giving birth dur the period of t pupping have pupped That is the t point on the pregnancy curve occurs after the main breeding and the lmves ~ point is after the main pupping period These periods actually last for 4 or 5 months so these points are well after the peaks of breedtng and pupping

The time betbulleen the peak when the pregnant and the lmr point Hhen the fewest are the average gestation period This is about 85 months (Fig 4) The anestrus curve (Fig 1) shmvs the same thing betng a negative of the pregnancy curve

Hith three different methods we have estimated a gestation period of 8 to 9 months Hith the unimplanted period roughly equal to the implanted period This agreas bullmiddotlith the observed period in captivity (BarabashshyNikiforov 194 7) and with field estimates Any gestation period differirg greatly from this does not fit the data Conceivably one could apply a 20 to 21 month period to the SaTEpound information however if this 1vere the case females would have to breed vJhile accomp by a pup or be on a four year reproductive cycle This prospect stretches the imagination

There is a definite possibility that the gestatton period varies in sea otters as in land ot te~cs This most likely -middotwuld occur through variations in the unimplantec1 period The above discussion refers to the average gestation period Until better information to the contrary is obtained we must continue to assume that the average period is 8 to 9 months

Fetal Rate

the estimates of the gestation curve are corrects Kenyons (1969) Fig 4 should approximate the grouth curve of the fetus Fig 5 duplicates this and shmvs the actual Heights Fig 6 shous the same information more graphically It is possible to esttmgtte the of time in each v7eight classlt Fig 6 does not take the period for establishment of the embryo into account so Height Class 1 may be longer than shmm perhaps 15 days if we accept Chapcnans (Kenyon 1969) es e

--

3 0

~

shy

----- -shy --middotmiddot~middotmiddot

___ ___1__

_

(middot-)

_ ft -) J ~)___

~-

3

Reproduction - 9 - IYarch 1971

The short time span of Classes 2 and 3 explains the relatively small numbers and the lack of a consistent pattern found in those classes (Table 1)

Using this curve it should be possible to predict the time of birth and with less precision the time of Luplantation and conception of any particular fetus By plot the estimated birth dates of a large number of fetuses rve should be able to drav curves Ttlhich approximate the breeding implantation and pupping rates of the population throughout the year

Unfortunately there is overlap bet1veen samples collected at different times of year Some of the samples are too small to provide a comparison We vould have to give eight to each in order to combine them Therefore only the ttay September and October samples are used lfuere the September and October samples overlap t1lt10 separate points are used Fig 7 presents the estimated birth dates grouped by month Fig 8 separates the dates into half month groups February and Narch are not represented and April is an estimate based on post-partum females rather than fetus size (see Recovery of the Uterus after Parturition)

The peak in this curve may be exaggerated It has been shmvn previously that fetus size is less consistent bet-Jeen samples than the other categories (Table 1) because the sample size of each grouping is small Therefore the exact shape of the curve may not be correct If however we assume the approximate of the peak is correct we can construct a model of the based on our estimates of the gestation period besed on a fetal grmJth rate estimated from this estimated gestation Therefore our reasoning would be some~hat circular if -Je used thjs to prove the estimate Hmvever we can see how this model fits the rest of the data A good fit would tend to support the model and the estimates on which it is based

For this model He use the curve in 8 to represent births We assume a gestation period of 8 months -rith the unimplanted and implanted periods each lasting 4 months TheTefore v8 draw curves identical to the birth curve but moved backward 4 months for implantation of the blastocyst and 8 months for conception (Fig 9)

A comparison of the peaks in Fig 9 with the data in Figs 1 2 and 3 and the narrative b on those indicates a fairly close fit The data indicat that the peaks may ~ctually occur slightly later than indicated in 9 but in the timing and distances bet7een the peaks fit

The sharpness of the birth curve indicates that the breeding in1plantatimiddotm and pupping peaks mey be more distinct than indicated by the curves in Figs 1 and 2 It is possible to fit curves with more abrupt changes to the data Fig 10 shows the same data lith the curves drawn to more closely fit the model tfuether these curves are more accurate or not is open to debate The data on Hhich the birth curve is based is not that strong The numbers in each month are relatively snall If such distinct did occur it seems likely that field observers ~vould have seen the effects Nost field observations indicate broader peaks that are not gtvell defined

2

G

X

middot ~ -- (-- ) r~ ~ - - -- ~

yen~ r- ~--~

3

i__middot -~---

~

gt

- cshy

Reproduction - 10 - March 1971

~e of _Sexual Maturity

Tables 4 - 7 list the frequency of nmnbers of corpora albicantia and corpora lutea found in females of different areas It should be recognized that the ages may not al-1ays be exact Animals ATTichi tka to become sexually mature Hhen about 4 years old It appbull~ars that more animals may become mature earlier on Adak The information is too limited to dratmiddotJ any definite conclusions from this hmmiddotJever the food availabili ty is probably better at Adak and an earlier age of sexual 1naturity may be a response to better nutrition

Ovulation

From Tables 4 - 7 it can be seen that the maximum number of corpora albicantia for each age roughly the numb~r of years after sexual maturity that some animals ovulated once every year after maturity It can be assumed that many corpora albicantia are invisible macroscopically after several years particularly if they are not remnants of a corpus luteum of pregnancT Therefore it Ls possible that most females ovulate every year This poss ili ty is also supported by the high incidence of large follicles in lactating females (TabL~ 10)

faximum

Tables 4 ~ 7 indicate that at t some females continue to breed at a very old age The samples are teo small to determine 1rhether a t number become unproductive at any point

Ovulation

Many mustelids are induced ovulatoTs

On September 14 1967 a female vJas tsd Hhile copulating She appeared to have Oilnlated shortly before Because sea otters may copulate several times over a period of two or three (Kenyon 1969) ovulation may have been induced by a previous

ficance of

Delayed lon is a charact8ristic of the reproductive cycle of most mustelids Several theories on the ccdv of a delay have been advanced Most assume that spring is the most favorable time of year for survival of nemiddotJborn young but that either a wir1ter breeding season is not favorable or that the presence of a Ttlale vhich occurs only during the

season is for survival of the young of the previous pregnancy (1963) discusse the evolution of delayed implantation in mustelids and points out that there are exceptions even in arctic areas middotJhere time of birth be consLdered more may be born at any tin1e Scheduling the and made possible by delayed mey be advantageous but is not necessary

----

a - o lt bull y bull bull l w laquo bull _ ~ ~

Table 4 Frequency of occurance of Numbers of Corpora Albicantia Amchitkll - June-August 1968

N U M B E R 0 F C 0 R P 0 R A A L 8 I C A N T A

I

~I -AGE -~ 1 I 1084 6 92 5 7(Years) 1 middot shy

- 0-l - ~~ ~- middot~middot- middotmiddotmiddotmiddotmiddot

1-2

2

3

4 1 I

2 (1)

6

middot5

1 (1)1

2 (l) 17

1 ( 1 )

2 ( 1)

18

l (1) I ( 1 ) 9

10 1 ( 1) L ~j( 2 1 ( 1 )

2 ( l) 1

12

11

1 (1)

I I (I)

1

2 ( 1)

13

1 ( 1 )

2

114

115 I116

17

I 18

I I

I

119 I

Numbers inside parentheses =Number of individuals with corpus luteum

11

Table s Frequency of oc~urance of Numbers of Corpora A1bicantfa _Kanaga - October 1968

N U M B E R 0 F C 0 R P 0 R A Al B CANT IA - I I I

AGE 84 I I

5 I l 6 7z 3(Years) 1

l rbull ~middot--

--- shy

0-l

J l-2

~ 2

3

4 2 ( 1)

5 3

Ibullbull 6 6 (3)

4 (4)7

8 4

3 9 2 ( 1 )

j iI 10

I 1 (1)11

12

13

14

1 ( 1)

16

15

17

18

Numbers irisi

-

J

(Plus 2 with corpus 1 uteum but no corpora a 1 b i cant i a)4(1)

1

1 (1)

5 (1)

10(5)

3

3 1

3 (2)

2 ( 1) 3 (3)

1 ( 1)

2

1 ( 1 ) 5 ( 1)

1 ( 1 ) 3 ( 1)

1

3

2 ( 1 )

1 (l)

1 (1)4 ( 1)

2) 1 (1) 3 (3) 4 (1) 12 ( 1 ( 1 )

1 ( 1)

1

2

1 ( 1)

1 ( 1)

1

I I parentheses = Number of individuals with corpus luteum

11

Table 6 Frequency of occurance of Numbers of Corpora Albicantia Mid Pt-~Blind Pt~ Adak- October 1968

N U ~1 B E R 0 F c 0 R P 0 R A A L B I C A NT I A

-1AGE

~

9 1084 75 6Years) 1 2 3 - shy ----- --middot shy~--

- _ Q~L

1-2

(Plus one with_corpus luteum but no corpus a1bicans)2 1 l I I I I I ) (One with corpus luteum but no corpus albicans 3

4 2 (1)

l5

16 1 (1)

1 1)7

8

2 (1)9

10

111

12

I13

14

15 1 (1) 16

17

18

1 ( 1)

1 (l)

1

1

1

2

1

1

1

1(1 )

1

1 (1)

1 (1)

1

I

1

1

1

1

-I

Numbers Inside rentheses = Number of individuals with corpus luteum

1

Table 7- Frequency of occurance of Numbers of Corpora A1bicantia Three Arm Bay - Bay of Islands Adak - October 1968

AGE (Years)

Q-1

t-2

2

3

4

s 6

7

8

9

10

11

12

13

14

15

16

17

18

Numbers

N U ~ B E R 0 F C 0 R P 0 R A A l B C A N T I A

~L 3 4 slE - 1middotshy

7 1~8 __J~l_11 shy

1-

One -Ji th corpus 1uteum but no corpora _a 1bicantia I I I I I I I

One vJi th corpus luteum but no corpora al bicantia

12 I I (Plus one with corpumiddots luteum but no corpora albicantia)1 (1) 3

1 (1) 2 (1)

1 2 (2) l 1

1 (1) 4 (1)

2 (1)

2 1 )

1 ( 1 ) 1

2 (1)

2 (2)

l22 3

3 ( 1)2 21 ( 1)

2 ( 1)

2 ( 1)

l ( 1)1

1 1

1 I ll

I

I I

inside parentheses Nurnber of individuals with corpus luteurn

Reproduction - 11 - March 1971

The sea otter lives in an area of relatively uniform temperatures However storms tend to be more frequent and violent in fall and winter Survival may be better in the late spring and sunrmer Therefore there may be some advantage to birth in the late spring However there does not appear to be a great deal of advantage to a breeding season at any particular time of year

Wright (1963) pointed out that most musteHds lant after daylight begins to increase usually around February This is the period when sea otters implant at the t rate If the time of implantation is influenced by daylight the period when the greatest number of births occur could be shorter than the equivalent breeding period The data do not show this although the possibility cannot be ruled out

The fact that substantial numbers of sea otters are breeding implanting and pupping at all times of the year indicates that hile there may be some advantage to certain events taking place at a particular time of year that advantage is not great enough for a distinct breeding season to evolve

Sea otters exhibit certain characteristics that are comnon to most mustelid breeding cycles but these chfracteris tics are not as well developed as in most es

It is interesting to note that Hright (1963) correlated the molt of Mus with the implanted period TI1e molt began around the

ion and ended around parturi tioD

Sea otters molt all year and some are implanted pregnant at all times At this time e cannot say uhether any between molt and pregnancy exists

Fetal

Kenyon (1969) found that the number of caudally presented fetuses roughly equalled the nunber of cephali presented fetuses He suggests that this indicates a lack of tatim1 for birth occurring in water and infers that sea otters must give birth on land

Fetuses of all 1veights in the present study also appear to be randomly oriented however 97 Class 5 fetuses (1000 g and over) showed 60 percent cephalic ion while only 40 percent shovJed caudal presentation

The orientation of large fetuses should be a better indicator of orientation at birth Smaller fetuses may position Therefore it appears that more sea otters are born head first than tail first

If you accept Kenyons premise that caudal presentation is necessary or at least beneficial to birth in water this more recent information supports the supposition that birth occurs on land The little information available indicates that birth does occur on lando but there have been unconfirmed reports of birth in water

Reproduction - 12 - tltIarch 1971

At least some sirenians are born head first indicating that caudal presentation is not necessary North of Unimak Island a large population of sea otters lives off-shore There is little evidence that any numbers come ashore It seems likely that many of these animals are born in the water

Of 305 implant pregnancies 138 fetuses (45 percent) had implanted in the left horn and 167 (55 percent) in the right horn Most of the difference was in the 1970 Amchitka sample ~fithout this sarnple 48 percent implanted in the left horn and 52 percent in the right horn Kenyon (1969) found an equal number in each horn If a real difference exists it is probably exaggerated by the 1970 A~1chitka sample

Out of 305 implanted pregnencies the point of implantation was on the same side of the reproductive tract as the corpus luteum in all but three instances In one case the corpus lutaum was in the left ovary and the fetus in the right porn In the second case the corpus luteum in the right ovary but the fetus was in the left horn In the third case there was a corpus luteurn in each ovary and tHo fetuses (of different sexes) in the left horn

It appears that blastocysts rarely cross from one horn to the other

Because there is usually a long period of time parturition and the next est_rus is little if any inhibitory effect on ovulation caused by the corpus of the previous pregnancy The ovary producing the largest follicle appears to be random and there does not appear to be the tendency for a pregnancy to be from the opposite ovary from the previous pregnancy In many cases there are many more corpora albicantia in one ovary than in the other

When a loses a pup shortly after birth and enters estrus before the uterus has conpletely recovered and before the corpus luteum has completely degenerated there may be some inhibitory effect Of the ll+ such cases identified six had large follicles in the same ovary as the new corpus albicans and in the opposite ovary It appears that any inhibitory effect by the corpus luteum only lasts for a short time

A few cases bullv-ere found vJhere one ovary was not developed or othenvise not capable of producing ova Because one ovary may support repeated pregnancies such animals may be as productive as those ~vith both ovaries active

Kenyon (1969) reported that most implantations occur within the central third of the uterine horn This held true for the animals in the present study they may implant In one case with a near term fetus the placenta covered the at the base of the horn blocking the exit of the fetus

Reproductive - 13 - March 1971

Birth iltleigh t

Barabash-Nikiforov (1947) listed the vJeight of a ne-v1born sea otter as 2000 g Kenyon (1969) presents the best quantitative data published to this time He found a maximum fetus weight of 1869 g 1-Thi1e he found pups as small as 1020 g those under 3 lb (14 kg) had died shortly after birth He estimated that successful birth weights range from 3 to 5 1b dr 1 4 to 2 3 and for purposes of calculation assumes an average birth weight of 1850 g to 1900 g

The weights of Weight Class 5 fetuses the smallest pups found living in the wild and three pups born in captivity (all died within 24 hours) are presented in Fig 11 These data tend to support Kenyons (1969) estimate of the ~veight at birth Few living pups middotHeighing less than 1350 g or about 3 lb and fevJ fetuses over 2000 g ( 4 5 lb) are found Kenyons upper estimate appears to be slightly high although there are extreme cases in both directions One pup weighed only 2 lb (900 and one fetus could not be weighed accurately on available equipment but weighed over 2500 g

One female with a 4 lb (1816 g) pup still had the placenta attached to the uterine wall and must have given birth shortly before being collected

From 11 it appears that most pups are born when they weigh in the neighborhood of 1800 to 1900 g in most of the populations sanpled Hmvever the and Delarof s indicate a bith Jei of between 1600 and 1700 g The Delarapound sample is srEall but the Tanaga sample the largest containing over a third of the large fetuses collected and was collected at a time of year when the birth rate is very high

This may be a reflection of the phys condition of the adult females and indirectly a reflection of food availab ty There are other factors indicating that the population is declining because of overpopulation

This lov1er birth yeight raises the question of middotwhether the pups are born earlier or whether are in poorer condition when born In an effort to gain some insight into this the fetal of Class 4 and 5 fetuses from Tanaga were plotted against their total lengths (Fig 12) middot Fetuses from other eastern Adak which appears to be a very healthy population were checked this curve fonaed by the Tanaga fetuses In all cases they fell Jell Jithin the limits of the Tanaga data indicating that there is no difference in the of fetuses of a given length It seems unlikely that a reduction in the rate of weight gain vould be completely proportional to a reduction of linear growth It appears that the rate of growth of fetuses is relatively constant but that females in poor physical condition may not be able to support as large a fetus as those in good condition so parturition occurs earlier

This may have sone effect on information on this

Fig 12 demonstrates some other interesting points There is no difference in the ratio and the maximum fetus size of males and females

~

(

~

-s 2 0 C

U)

0

--

ez ()

~) I) middot _t t- (

j -~middot -~

Reproduction - 14 - March 1971

This infeTs although dolt2s not prove an identical grmgtlth rate afld birth weight for males and females The circles on Fig 12 show the ~eight-length ratio of pups These fall below the curve formed by fetus weight-length ratios indicating that pups of a given length are some-Jhat heavier than fetuses of the sarne length This indicates an increase in weight immediately after parturition

Sex Ratio at Birth

Kenyon (1969) found that of 58 fetuses for which sex could be determined~ 45 percent vJere male and percent were female He assumed an equal sex ratio until a larger sample could be obtained

In the present study 111 fetuses were male and 144 ~vere female When this sample is combined with Kenyonts the sex ratio of 313 fetuses is 44 percent male and 56 percent female

Sex Ratio of the

In the process of harvesting capturing animals for transplants and delineating male and femalen areas it has become apparent that there are more females in the population than males Our general ion is that perhaps 60 to 65 percent of the sea otters are females Kenyon (1969) demonstrated that more juvenile males than females are found dead on the beaches of Amchitka lhe age structure of harvested in 1968 indicates that males may not live as long as fetnales

The lower birth rate higher juvenile mortality rate and slightly shorter life expectancy of males could account for the unbalanced sex ratio in the population

The sex ratio may vary from one island to another Presumably there -vmuld be more females in a population that has reached the carrying capacity of the habitat where juvenile mortality is higher There is some indication that males are the first to expand into new areas of unoccupied habitat Therefore in recently populated anas one might find more males However this situation vould be

Lit

The normal litter size of the sea otter is one This has been recorded by almost every observer since Steller Barabash-Nikiforov (194 7) reported twins in utero and mentioned other reports of twin fetuses from the early days of SnoF (1910) recorded a set of neHborn tdns In more recent studies~ both Sinha et al (1966) and Kenyon (1969) found reproductive tracts with two corpora lutee but in all cases only one fetus las present In thousands of hours of observation by many observers no tbullvin pups have been reported

In the present study 24 instances of multiple ovulations were recorded Five of these resulted in hv-in and one in triplet fetuses The information is summarized in Table 8

Table 8 Multiple Corpora Lutea Fetuses

Total Pregnant Females 565

Implanted Pregnancies 316

Unimplanted with 2 CL (corpora lutea) 9

Implanted Pregnancies with ) L CL and 1 Fetus 8

Implanted Pregnancies Vlith 3 CL and 1 Fetus 1

Implanted Pregnancies gtvith 2 CL and 2 Fetuses 5

Implanted Pregnancies middotwith 3 CL and gt Fetuses 1

Total Nultiple Ovulations

I

Percent Nultiple Ovulations 42

Percent of Pregnancies tvith Nultiple Fetuses 19

Corpora Lutea per Pregnancy 105

Fetuses per 102

Reproduction - 15 - lvlarch 1971

From these data it appears that in over 4 percent of the estrus cycles more than one ovulation takes place Of these about half result in the development of more than one fetus In most cases the were of a relatively large size and probably would have been born normally

All multiple fetuses appeared to be the result separate ovulations All had separate placentas and in some cases were different sexes Four tracts with twins had both fetuses in the same horn one had one in each horn and the tract with triplets had one fetus in one horn and t~vo

in the other Another tract had skeletal remains of triplets in one horn that were being resorbed Another tract appeared to have had five fetuses in one horn but they were almost completely resorbed One might infer that physically there is not enough room in one horn to accomn10date more than two fetuses for any length of time

With the exception of the newborn tvins reported by Snow (1910) there are no docu_mented records of female sea otters with tvin pups There are occasional unconfirmed reports of twin pups but as Kenyon (1969) points out a female may tolerate a pup in addition to her ovm but it is unlikely that a female could care for more than one pup It is unlikely that more than one pup survi~es In any case orily 2 percent of the pregnancies produce more than one pup 1middot1ultiple births cannot be a significant factor in the productivity of sea otter populations

The presence of a corpus luteum without gross evidence of implantation t-7as assumed to be an unimplanted pregnancy No attempt ~vas made to locate unimplanted blastocysts As a result e have little information concerning the survival of the blastocyst Of 206 reproductive tracts classified as nulliparous on the basis of the appearance of the uterus 12 had one corpus albicens and one had two Some of these may have been large attritic follicles Others may have ovulated and even conceived but implantation did not occur All of these cases represent the failure of the antmals first estrus

A total of 16 resorptions or possible abortions were identified (Table 1) The follmJing is a brief descdption of these ~7i th possible causes of some

Des

Resorption of blastocyst or ovum (corpus luteum 4 degenerating no evidence of ion)

Failure at time of (possibly because 1 of growmiddotth inside horn adjacent to implantation site)

Resorbed or aborted fetus (implantation at end of horn) 3

Resorbed fetus (umbilicus tvTisted tightly) 1

Resorbed fetuses (three or more fetuses in one horn) 2

Resorbed fetus (cause unknmm) 5

Reproduction - 16 - March 1971

Six of these resorptions may have been caused by a lack of room for growth of the placenta because of crowding from other placentas a growth or the end of the uterine horn These six and the one vith the twisted umbilicus are probably the result of random events or accidents

The numbers of resorptions are too small to accurately measure the frequency of such occurrences although the relatively large number in the Tanaga sample (Table 1) may be a reflection of the physical condition of the animals Part of this number is probably due to the fact that there are more pregnant animals in the sample In general the samples with the most pregnancies also conta~~ned the most resorptions Half of the Tanaga resorptions can be attributed to random events but five vere still due to unknown causes Eight of the 10 resorptions fo1md on Tanaga came from a 20-mile stretch of shore about 25 percent of the harvest area

While concrete conclusions cannot be drawn from these data they do raise the possibiliy that under certain conditions 5 percent of the pregnancies may fail

near Parturition

Collecting methods were such that the presence of a pup with a female was often overlooked Therefore the lack of a pup with a post-partum or lactating female did not mean that the pup had died However 14 females shmving of recent pregnancy were estrus presumably having lost their pup shortly before or shortly parturition

Again the magnitude of this mortality cannot be determined but it appears to be of the same order as the in utero mortality

after

Reproductive tracts were subjectively classified as post-partum or anestrous on the basis of degeneration of the corpus luteum (or appearance of the corpus albicans) the size of the horn of pregnancy and appearance of the placent scar In an effort to get an idea of how long it takes for the tract to return to normal the size of pups accompanying females in each category is listed in Table 9

It appears that females pass the subjective boundary from post-middotpartum to anestrus Hhen the pup about 10 lb and is around 75 to 80 em long

We do not know a great deal about the early growth of a sea otter pup but judging from the cur1e e9tablished based on cementum deposition of older animals and from tooth eruption a 10 lb pup is probably in its second month of life

Table 9 Pups Accompanying Pos artum Females

Sex llleight

M 2 lb F 3 F 4 M 5 F 55 F 625 F 775 M 10 F 11

Total Length

47 em 52 56 60 65 65 70 81 (borderline post partum-anestrous) 82 (late post-partum)

Pups Accompanying Anestrous Females

F 10 lb 78 em M 11 79 M 12 8l F 13 83 M 13 92 H 16 83 M 17 91 1 19 87 M 19 90 F 21 107 F 22 96 1 22 101 M 2Llmiddot 97 1 26 100 F 28 103

Reproduction - 17 - Narch 1971

Frequency of Breeding

Steller (1751) reported female sea otters with ne~vborn pups also accompanied by another pup that seemed to be no more than a year old Nurie (1940) recorded an instance of copulation by a female accompanied by a pup and Jones (1952) caught a young pup simultaneously gtvith a copulating pair Lensink (1962) felt that few females breed until the pup is a year old but mentions females with small pups also accompanied by large pups

I believe that some of thes2 observations may be misinterpreations of the situation Females will occasionally leave small pups for a time often near other animals The pups Lensinkmentions would be 2 years old and probably veigh 30-35 lb Sea otters are gregarious and subadults may remain near other animals appearing to be accompanying their mother I suggest that most of Stellers and Lens inks large pups ere such cases

Females gtvi th pups probably do breed occasionally but the most recent evidence indicates that this rarely occurs Kenyon (1969) records no instances of females accompanied by pups copulating and states that he found no females- knm7n to be accompanied by a pup that were This holds true for the animals collected in the present study However the nature of the harvesting operation is such that many females accompanied by pups were not recorded as such Therefore we are forced to look at lactating and assume that they were accompanied by a pup at the time of collection or shortly before le 10 sm1marizes the reproductive condition of the lactat females In the 1967 and 1968 samples some animals vith enlarged marmary middotmre listed as lactating As a result some females with near tenrr fetuses Here included Because this condition is considered to be associated with the unborn pup rather than a previous pup these animals were with the anestrous animals in le 10 In 1969 and 1970 the presence of milk was used as the sole criterion and no term animals we-rmiddote in the sample

Of 246 lactating females only one had an ted fetus (excluding the near term animals from 1967-68) Several others had small corpora lutea indicating that they recently become or possibly had large

vhich had luteinized but tvere not actually pregnant The remainder had follicles

This information that accompanied by a pup rnay enter proestrus and even ovulate but that they only rarely mate That they enter estrus is subs iated by the f2ct that the number of corpora albicantia in many tracts equals the age of the amplimal minus the three years prior to nBturity (see les 4-7) This indicates that large follicles tend to develop year after sexual nl8turi ty is reached whether the female is accompanied by a pup or notlt

Some of those lactating females with s1all corpora lutea may have recently weaned or othenvise lost a pup and are already pregnant again Malkovich (1937) took a pup least 3 months old) away in cap She mated 12 days later here she had the male It appears that a female enters estrus shortly after Heaning

Table 10 Reproductive Condition of Lactating Females

Location

Bay of Is Adak I Sept 1967 9 1 100

Allchitka I Sept-Oct 1967 17 2 105

Mid Pt - Blind Pt Oct 1968 19 3 136 Adak I

Bay of Is Adak I Oct 1968 22 4 154 3

Kanaga I Oct 1968 32 s- 135

Al1chitka I July-Aug 1969 4 2 333

iTanaga I May 1970 56 10 152

Delarof Is May 1970 18 3 143

Amchitka I May 1970 36 3 77

---~--middot--middot----~-~----~----middot~-middot--~----~----- -------~-~ middotmiddot--middot-shy

TOTAL 213 33 134

Anestrous or pregnant with term fetus

]_ Large follicles or corpus luteum present

3 Includes one implanted pregnancy with small fetus

Reproduction - 18 - March 1971

The question of hml long it takes for a female to enter estrus again after losing a small pup arises Eleven tracts of the 1970 sample and ttvo of the 1968 Kanaga Sampnple shmJed signs of being pregnant recently) but were in proestrus These animals had an enlarged horn usually middotlith a slightly pigmented area but no actual placental scar a11d a recently fonned corpus albicans However they also had enlarged follicles and the tvalls of the uterus were typical of proes trous animals Another animal was similar but appeared to have already ovulated and a corpus luteum was present In these cases the female has lost a pup either through abortion or vithin the first veeks after parturition and has started into another estrus cycle Three or four of these had follicles vhich may have started to become attritic This may be because the uterus had not recovered sufficiently from the last pregnancy

The implication of this information is that with a normal pregnancy and successful rearing of a pup the female becomes pregnac1t only after the pup has been weaned This n~y be a year after parturition However if the pregnancy fails or if a pup dies even at birth the female may become pregnant again in a few weeks rather than _waiting the full year

Kenyon (1969) suggests that the period middotbetween and the next estrus may long because he found animals that tvere not lactating but had a faint placental scar and an indistinct but recent corpus albicans These animals ltJere cLtssi as anestrus in the present study They are undoubtedly bet1veen veaning of their last pup and the next estrus The number of suh animals is t durirg Sspte12~ cnd October At this time the rate of estrus increases a1d the number of anestrous animals decreases sharply the next feJ months Therefore many of these anestrous anirrals become pregnant 1lithin a month or two Kenyon (1969) suggests that if the female keeps the pup for about a year that the period between births may be smreltmiddotJhat greater than two years He is assuming a 12-13 month gestation period However the present study shows that the estimate of a 12-13 month tation period was based on an inadequate sample Estimates in the present study put this period at closer to 8 or 9 months Presumably the anestrous nonlactating females are in the 3 to 4 month period

This does not change Kenyons (1969) point that the and the next estrus may long It indicates that in a normal cycle it may be months Hmvever as it vvas shmvn above~ it is possible for a female to enter estrus very shortly after losing a pup

We have demonstrated that sea otters breed at all times of year and it is possible for a female to become pregnampJt again after losing a pup sooner than she vould have if the pup had survived HmmiddotJever we have also demonstrated that annual of b and pupping occur and that these are similar in different years and in different populations The assumption can be made that the average length of the entire reproductive cycle is some multiple of a year

Kenyon (1969) assumes that tbe pup remains with its mother for about a year ~-Ieights and measurements cementum deposition skull changes~

Reproduction - 19 - March 1971

observations in the wild and in captivity all support this assumption although it cannot be said with certainty that it averages 12 months It could be s tly ITDre or less

Assuming about 12 months for rearing the pup 8 to 9 months gestation plus a fev months rest bet~veen and the next estrus the average reproductive cycle takes two years

This is supported by Fig 4 which shows that slightly over half the mature females are pregnant in a year Kenyon (1969) pointed out that his January and Harch samples gtvere biased against females with pups a1d therefore in favor of pregnant females because of selective hunting This bias applies to the fall sa~ples as well The summer samples were biased similarly because pregnant females appeared to be less resistant to handling during capture Therefore the percent of pregnant females is shown to be higher than lt actually Has -rhile no correction factor can be applied to all samples the evidence indicates that the percent of pregnaJt females is usually around 10-middot15 lmrer than in the sample For example the 1970 Alnchitka sltLrnple t-ras less biased and was about 9 percent lower than the Tanaga sample vrhich is l)nm-rn to be more biased Also on a June skiff survey 7 percent of the animals counted had pups that were readily identified If ~re assune that this is the nunber of animals avoided it can be calculated that 15 of the sexually mature females all of vhich are t middotTen~ avoided Actually this will vary with the hunter weather time of year etc Hmvever it indicates the order of magnitude of the bias

Using this as a rough correctton for the data in 4 we find that about half or slightly fe-Jer of the sexually mature females are pregnant each year or as indicated ly the average female has a pup every two years

Reproduction - 20 - March 7 1971

Conclusions

1 While sea otters may breed or pup at any time of year there are seasonal fluctuations in the illliTlbers of females in each reproductive stage The pattern of these fluctuations remains the same from year to year and population to population

2 Breeding activity is highest in September October and November

3 More implantations occur during January February and March than at other times of year

4 The birth rate is highest during April Y~y and June

5 Tlvo and a half to three times as many females breed during the peak breeding months as during the months of lowest breeding activity

6 Similarly two and a half to three times as many females pup during peak pupping months

7 During the period of lovest b activity seually mature females breed at a rate of 2 to 3 percent per month During the peak of breeding 7 to 10 percent per month breed

8 During the period of lov1est pupping activity 2 or 3 percent per month pup and during the peak of pupping 7 to 10 percent per month pup

9 The gestation period about 8 to 9 months on the average About half of this time is ~~ ted period

10 Female sea otters become sexually mature ihen 3 to 4 years old

11 Females may ovulate on an average of once a year after sexual maturity is reached

12 Females continue to breed at Cn old age

13 Sea otters retain certain characteristics common to the reproductive cycles of many other mustelids however exceptions are common Scheduling of the reproductive cycle during the year may have some advantages or it may be a trait that has not been entirely lost through evolution

14 Sixty percent of the near term fetuses are cephalicly oriented and 40 percent are candllly oriented

15 Sea otters probably birth both on land and in the water

16 Blastocysts rarely cross from one uterine horn to the other

17 Consecutive pregnanctes may occur in the same horn The side of pregnancy appears random

Reproduction - 21 - March 1971

18 Implantation usually occurs in the central third of the horn but it may occur any~rhere in the horn

19 Birth weights may range from 900 g to over 2500 g however most pups weigh 1800 g to 1900 g at birth

20 Females from areas gtvhere the population is declining because of food shortages may pup earlier and as a result the pups 1 birth weight may average 200 g lmver than nonnal

21 Male and female fetuses gr01middot1 at the same rate ampJd are the same size at birth

22 Body condition of the female has little effect on the growth rate of the fetus

23 There is a rapid ~Jeight gain shortly after parturition

24 The sex ratio at birth is 44 percent male and 56 percent female

25 The normal litter size is one hmmiddotever in 2 percent of the implanted pregnancies two or fetuses survive to near term

26 Multiple ovulations occur in 4 percent of the estrus cycles

27 es do not appeErc- to be able to more than two fetuses in a single horn

28 Up to 5 percent of the p may fail dne to in after implantation

29 An unknown but probably significant number of first pregnancies fail before implantation

30 The uterus is usually considered recovered from a pregnancy when the pup vleighs about 10 lb and is probably in its second month of life

31 Sexually matu1middote females normally have one pup every two years

32 Females vrith a pup may ovulate but rarely mate

33 Slightly less than half of the sexually mature females are in any given year

34 A female may enter estrus within a few weeks of losing a pup Consequently she may become agaln gtvi thout waiting the same length of time as if the pup hacl survived

35 Hhen a pup is weaned normally there is a longer period perhaps 3 or 4 months between weaning and the next estrus

Reproduction - 22 - March 1971

CITED

Barabash-Nikiforov I I 1947 The sea otter (Kalan) Soviet Ministrov RSFSR Translated from Russian by Dr A Birron and Z S Cole Israel Program for Scientific Translation 1962 227 pp

Hugget A St G and W F Widdas 1951 The relationship betlveen marmnalian foetal -reight and conception age Jour Physiology 144306-317

Kenyon K W 1969 The sea otter in the eastern Pacific Ocean USFWS North American Fauna No 68

Malkovich T A 1937 The sea otter in captivity Priroda No 381-87 Translated by J T Naximovitch 1966 Fisheries Research Board of Canada Nanaimo BC Translation Series 657

Sinha A A C H Conavay and K t-J Kenyon 1966 Reproduction in the female sea otter J Hildl Ngrrit 39(1)121-130

Snow H J 1910 In forbidden seas Edward Arnold London 303 pp

Wright P L 1963 Variations in reproductive cycles in North Arrierican mustelids In Enders A C ed Delayed tation Univ of Chicago Press 318 pp

  • SEA OTTER STUDIES - 1970
    • Surveys
    • Harvests and Transplants
      • Sexual Segregation in Sea Otter Populations
      • Reproduction in the Female
        • AERIAL COUNT OF SEA OTTERS- SOUTHSIDE OF THE ALASKA PENINSULA
          • Description of Conditions During March 1970 Aerial Count of Sea Otters
          • Discussion of Results
          • Sanak Island-Sandman Reefs
          • Shumagin Islands
          • Sutwick Area
          • Augustine Island-Cape Douglas
            • SUMMARY OF SEA OTTER SURVEYS
            • Amagat 1 to False Pass Ikatan Peninisula Cape Lazaref
            • Sanak Is
            • Deer 1
            • Pavlof Is
            • Northern Shumagin Is
            • Cold Bay 1
            • Kupreanof Peninsula
            • Mitrofania 1 to Kupreanof Pennisula
Page 5: Sea otter studies - 1970 - Alaska

11arch 27 Shaw Island to Puale Bay (1030 AM - 1205 PM) Conditions very good to excellent High overcast and very little wind down to Katmai Bay After this the conditions deteriorated rapidly to fair and the count had to be stopped because of a combination of bad weather and lack of fuel As shown on the charts the count was concentrated on the points and islands however the good visibility allowed us to see well into the bays We feel certain no concentrations were missed

The numbers of sea otters counted are presented in the Table Specific locations and numbers are shown on the charts Where coverage of an area was not complete the approximate path of the aircraft is shown

Discussion of Results

Surveys along the Alaska Peninsula have been fragmentary and most of those made were done under less than ideal conditionsmiddot There are a number of reasons for this Weather tends to be poor along the south shore where squall5 form along the mountains Areas where an aircraft can refuel are north of the mountains and there are relatively few passes therefore range of the aircraft is a problem Also there is much shallow water off shore in the area from the Shumagin Islands to Sanak Island Coverage of this area is difficult and many animals are undoubtably missed

The present survey covered most of the area however conditions were such that most of the counts are probably quite low Despite these problems and a lack of continuity we have a fairly good picture of the recovery of sea otter populations in this area

Reports from various individuals are available from the 1930s and 1940s 11ajor surveys by the Fish and Wildlife Service in 1951 (Jones) 1957 (Lensink) 1962 and 1965 (Kenyon and Spencer) covered at least portions of the area In 1969 the southern Shumagin Islands were counted by the Alaska Department of Fish and Game and the present survey covered most of the rest of the area

Basically there are four distinct population nuclei in the area These centered around the Sanak Island-Sandman Reef area the Shumagin Island area the Sutwick Island area and the Augustine Island-Cape Douglas area The following discussion is based on information from reports by Lensink and Kenyon and from Alaska Department of Fish and Game surveys middot

~anak Island-Sandman Reefs

Small numbers of sea otters have been reported at Sanak Island since 1922 No reports came from the Sandman Reefs until 1942 In 1948 27 were sighted at

----Cherni Island The 1951 aerial survey showed 65 around Sanak and 97 in the Sandman Reefs In 1957 251 were seen around Sanak and 508 around the Sandman Reefs In addition two were seen in the Pavlof Islands however few were on the mainland In 1962 548 were counted in the Sanak area and 638 in the Sandman Reefs None were reported from the mainland or Pavlof lsland~however

much of the increase was in the northern area around Deer Island The 1962 survey was probably the best being made under excel lent conditions

AERIAL COUNT OF SEA OTTERS MARCH 1970

Partial or Location Count Date Camp 1ete Count Visibility

Cape Lazaref - Cold lsecty_

Cape Lazaref 3 320 Complete Poor

lkatan Peninsula 71 320 Complete Poor

False Pass - Amagat I 66 320 Camp 1ete Poor

Cold Bay 321 Partial Poor

TOTAL 141

Sanak Islands 239 322 Camp 1ete Fair to Poor

Sandman Reefs

Clubbing Rocks 12 322 Complete Fair to Poor

Cherni I amp Rocks 495+ 322 Complete fair to Poor

Goose I 7 322 Complete Fair to Poor

Hay I 18 322 Camp lete Fair to Poor

Hunt I 2 322 Complete Fair to Poor

Deer I 322 Partial Fair to Poor~

TOTAL 568+

Pavlof Islands

Inner 11iasik 2 321 Camp 1ete Fair to Poor

Outer lliasik 16 321 Camp 1ete Fair to Poor

Goloi 2 321 Complete Fair to Poor

Dolgoi 67 321 Complete Fair to Poor

Poperechnoi 29 321 Complete Fair to Poor

Ukolnoi 2 321 Complete Fair to Poor

yenWosnesenski 4 321 Complete Fair to Poor

TOTAL 122

Partial or Location Count Date Comp 1ete Count vis ib j 1i ty

Northern Shumagin Islands

Unga 184 321 Complete Fair to Poor

Popof 52 321 Complete Fair to Poor

Korovin 46 321 Complete Fair to Poor

Karpa __2t 321 Complete Fair to Poor

TOTAL 286

Cold ~ to Beaver ~ 0 320-21 Partial Poor

Beaver Bay to Kupreanof Pen

Beaver Bay 2 321 Partial Fair to Poor

Cape A 1 iaks in 4 321 Partial Fair to Poor

Guillemot I 16 321 Partial Fair to Poor

__1Kupreanof Peninsula 321 Partial Fair to Poor

TOTAL 23

Kupreanof Pen to Castle Cape 0 321 Partial Fair to Poor

AntJrrCastle Cape to-1-Ji e ~

Castle Cape-Castle Bay 16 321 Partial Fair to Poor

Chignik Bay 118 320 Complete Fair

Nakchamik I 5 320 Complete Fair

Hook Bay 13 320 Complete Fair

Cape Kum 1i un 88 320 Complete Fair

Kujul ik Bay 1199 320 Complete Very Good

Univikshak I 62 320 Complete Fair

Cape Kuml ik 54 320 Complete Fair to Poor

Sutwick I 14 320 Complete Fair to Poor

Cape Agutka 323 Complete Fair~

TOTAL 1766

Partial or Location Count Date Comp 1ete Count vis i b i 1i ty

Cape Kunmik 13 323 Complete Fair

Naka 1i kok Bay amp offshore islands 16 323 Complete Fair

Chiginagak Bay 5 323 Complete Fair

Agripina amp lmuya Bay 105 323 Complete Good

Wide Bay to Puale Bay N 0 T S U R V E Y E D

Puale Bay to c Kubugakl i __]_ 327 Partial Fair

TOTAL 146

Kashvik Bay to L Chiniak 0 327 Partial Very Good to Excellent

Cape Chiniak to Shaw _I

Shakun Is amp Rocks 71 327 Partial Very Good to Exce 11 ent

Kiukpal ik J to Shaw 1 0 327 Partial Very Good to Excellent

TOTAL 71

In the present survey which was made under relatively poor conditions 239 were seen in the Sanak area and 568+ in the Sandman Reefs The latter count Was incomp 1ete and conditions were such that the number of otters was more a function of time spent counting rather than area covered Obviously many were missed As a result not much can be said about total numbers The main change evident is that 141 were seen along the mainland and eastern portion of Unimak Island and 122 were seen in the Pavlof Islands This indicates that a fairly extensive movement northward and along the Peninsula is occuring

A remnant population probably remained along the south side of Sanak Island in the early 1900bulls By the late l940 1 s they began spreading into the Sandman Reefs This northward expansion has continued to the present time There is still unoccupied or sparsely populated habitat along the mainland shore and in the Pavlof Islands There should be continued expansion of this population for a number of years although the numbers around Sanak Island and the western Sandman Reefs may have already reached a peak

With the arrival of substantial numbers in the Pavlof Islands this population is probably on the verge of mixing with the Shumagin Island populshyation No doubt some individuals have moved back and forth in the past but the two populations are almost continuous at the present time

As in past surveys few otters were seen around the north side of Sanak Island and Caton Island This is probably poor habitat for otters The main population is around the south and west sides The higher count on the west end is probably more a result of intensive searching rather than a higher population

Shumagin Is 1 ands

This has been considered the largest of the four populations in the Alaska Peninsula area The most recent counts have not been adequate enough to show any major increase in numbers in the last decade however major changes in distribution have occured which are very similar to the pattern mentioned in the Sanak-Sandman population

There were occasional reports of sea otters in the Shumagins in the 1930s By 1947 Victor Scheffer estimated that 500 1 ived around Simeonof Island In 1953 Hooper counted 633 around Simeonof and Little Koniuj i Island The 1957 survey showed 1829 Most of these were- in the southern islands Five individuals were scattered in the northern islands and one was on the mainland shore near Elephant Point The 1962 survey totaled only 1352 The animals were scattered well off shore and the count was low However the count on Nagai increased from 149 to 338 indicating a continued northward expansion

In 1969 1510 were counted in the southern islands under relatively poor conditions An additional 286 were counted in the northern islands in the present survey and 23 were seen along the mainland north of the Shumagins Again survey conditions were not good

There is a very clear expansion of the population from a center near Simeonof Substantial numbers now occur on all the islands and repopulation of the mainland is occuring The population in the northern islands should continue to increase and most of the habitat on the mainland is not occupied

There is no evidence of an increase in total numbers since 1957middot However the last two surveys have been less than ideal and the large shallow off shore areas have not been covered adequately Considering the survey conditions and the obvious extentions of range it is almost certain that the populations from San-ak Island to the Shumagins have continued to increase In general it appears that the southern islands and reefs have become fully populated and the northern areas are just developing significant populations The entire a rea may be comp 1ete 1 y repopu 1 a ted in the next 10 years This wi 11 depend largely on the status and potential of the off-shore areas

Sutwi ck Area

Reports of sea otters near Sutwick Island are available since 1936 This population was far removed from other populations and is probably a remnant left after hunting ceased In 1951 Jones counted 388 between Cape Kuml iun and Cape Kunmik Most were near Sutwick Island In 1957 889 were counted Nineteen of these were between Cape Kunmik and Cape Providence indicating some northeastward expansion In 1962 949 were seen with individuals straying as far as Cape lgvak and one between there and Cape Kuliak In 1965 a stray otter was seen at Kinak Bay A major shift in the population from Sutwick into Kujulik Bay occured This may be the result of time of year and weather

In the present survey 1766 were counted between Castle Cape and Cape ~~~gtf~aip the majority were in Kujulik Bay Large numbers have moved

~~ranging as far as Cape Kubugakl i The area from Wide Bay to Puale Bay was not surveyed because of weather

The total count for the population was 1912 even though conditions were less than ideal throughout the bcunt and we feel many were missed It is possible that the increase in numbers is entirely due to reproduction assuming a 10 percent annual increase However it is difficult to compare surveys

There is still room for expansion in both directions from this population The greater movement to the northeast is probably the result of better habitat The population around Kujulik Bay is very dense and a large movement of animals may occur if competition for food becomes serious Otherwise we should expect to see continued steady expansion into adjacent areas for a number of years

to come

Augustine Island-Cape Douqlas

For some time a population has existed around the Augustine Island area at the mouth of Cook In 1 et In 1948 approxImate I y 50 sea otters vvere reported from Augustine Island Reports of sightings have been made from Shaw Island to Tuxedni Bay In 1957 Spencer counted 40 at Augustine and one at Shaw Island Lensink counted 52 on Augustine in 1959 but he considered it a poor count In 1965 Kenyon counted 18 on Augustine and 101 in the Shaw Island-Cape Douglas area In March of 1969 Loren Flag saw 62 around Augustine and in May 1969 Jim Faro counted 130 another observer saw about 30 some of which were probably not tallied by Faro

On the present survey Augustine Island could not be surveyed because of weather No otters were seen around Cape Douglas but 71 were seen in Shakun Islands and rocks near the abandoned village of Kaguyak These were probably from the Cape Douglas group

It appears that the animals move about in the area To date no complete survey of the area has been made at one time Until this is done 1ittle can be said about the population other than that several hundred probably occur there

The following table is a summary of sightings and counts made by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game These data were collected by different individuals using different types of aircraft under varying conditions of weather A strict comparison of numbers should not be made from this table

SUM

MAR

Y OF

SE

A OT

TER

SURV

EYS

(Com

pile

d by

p

op

ula

tio

n f

rom

L

ensi

nk

(196

2 T

hes

is

K

enyo

n 0

96

2 amp

196

5 R

epor

ts

amp M

anu

scri

pt)

an

d AD

FampG

Dat

a)

Lo

cati

on

P

re

1951

19

51

1957

19

59

1962

19

65

1969

19

70

AU

GU

STIN

E -

C

DOUG

LAS

Aug

usti

ne

Isla

nd

A

ppro

xim

atel

y 50

(1

948)

Shaw

1

amp

Dou

glas

Are

a S

igh

tin

gs

from

Sh

aw

Is

to

Tux

edn

i B

ay

TOTA

L

40

___

_

41

52

-shy 52

18

_lQ

J

119

130+

-shy

130+

J

71

ALAS

KA

PEN

INSU

LA

c

Kul

iak

-

c

lgva

k

c

lgva

k -

c

Kun

mik

Ani

akch

ak

Bay

amp

Am

ber

Bay

Sutw

i ck

Are

a M

isce

llan

eous

R

epor

ts

Kuj

ul i

k B

ay

amp c

K

uml

ik

Sin

ce

1936

C

Kum

1 i u

n

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viks

hak

Isla

nd

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akch

amik

Is

lan

d

8

355 12

13

0 19 6

581

103

180

1

22

47

109

684 86

Not

S

urve

yed

7(+

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139

197 14

1 2

53

101 62

5

I

Lo

cati

on

P

re

1951

19

51

1957

19

59

1962

19

65

1969

19

70

ALAS

KA

PEN

INSU

LA

(Co

nt

)

Chi

gnik

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ay

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tle

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amp

Cap

e

TOTA

L

SHUM

AGIN

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d S

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re

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of

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to

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f B

ay

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cle

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del

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19

40

0

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388

889 1 2 2 1

149

26

8

~-

7

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338 I105

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118

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1 9

12

23

184 52

46 4

75

232 8 27 6

Lo

cati

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P

re

1951

19

51

1957

19

59

1962

19

65

1969

19

70

SHUM

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(C

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500

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947)

l63

3 (

195 3

)

3 0

220

430

455

160

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183

0

14 3

222

255

294 38

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150 15

296

290

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to

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2

141 4 2 29

67

Lo

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1957

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1962

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1969

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SAN

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27

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middot

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Cape Agutka Cape Kum 1 i k Sutwick I~ Kujul ik Bay Cape Kum 1 i un Unavikshak 1

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1199 88 62

1614

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75

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MARINE MAMMALS SURVEY - BRISrOL BAY TO AKUTAN ISLAND June 2 1970

OBSERVER J Vania J Faro

PILOT George Tibbets Jr

AIRCRAFr Piper Azetex

middotWEATdER Partly cloudy throughout flight wind 10 miles per hour or less visibility generally excellent One bad area and that was at Akutan Island Air was turbulant there and we were unable to go completely around the island because of fog in Akutan Pass Water had slight ripple throughout flight and ocassionally there was glare but overall conditions for sightinmiddot=r animals was good

Departed King Salmon 1115 AM Returned to King Salmon 745 PM

FLIGHT PA~1 middotAfter leaving King Salmon we went over to the north side of the Alaska Peninsula and proceeded down the coast flying at 300 to 600 feet of altitude Generally we flew aboumiddott one mile off the beach At Port Heiden Moller and Cinder River we checked the outer sand bars for seals and some of the inner bars where I knew seal hauled out

We did not cover Izembeck Laroon very well Instead we flew offshore and checked Amak Island We then fueled up at Cold Bay Leaving Cold Bay we continued dmvn the north side of the peninsula (sea otter sighted) down to Cape Sharichef and crossed over to Akutan Island flying along the south side of it We were able to get around Cape Morgan and fly doNn the beach a few miles but then had to turn back becausmiddote of fo9

The flight path was then eastward to the north side of Tigalda Island At Ugamak Island we circled it completely flying at about 300 to 500 feet We completed the survey at this point and returnt~d to King Salmon generally following the route we had taken on the flight down

SEA OTTER SIGHTINGS

Seven pods of sea otter were seen south of Amak Island These were photographed and later counted from middotthe photographs Beshycause of the density in the larger pods and the fact that some animals in the smaller pods were diving when the pictures were taken some individuals may have been missed

The number counted in each pod is as follows

1071 520 357 68 36 75 30

TOTAL 2157

Several hours later the pods had drifted four or five-miles northeastward (see map) bull In addition 1 one sea otter was seen at Amak Island and sixteen near Tigalda Island

171

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shy

SEA OTTER COUNT BARREN ISLANDS and SHUYAK - AFOGNAK ISLANDS

June 9 1970

On June 9 1970 an aerial survey of sea otter was flown in the Barren Islands and in the AfognakShuyak Islands area A Gruman Goose was used with Schneider serving as the only observer Offshore coverage was poor Conditions of visibility were as follows

Barren Islands- 1140 AM to 1225 PM- Water calm moderate surface glare Conditions verygood Count complete

Ban Island to Pernosa Bay- 1255 to 135PM- Conditions similar to Barren Islands but surface glare worse in spots Conditions generally good

Marmot Island to Latax Rocks - 255 to 345PM- Increased ripples on surface Glare Bad Conditions fair Count around Sea Otter Island complete although some otter may have been scattered offshore Remainder of count very superficial

Area

BARREN ISLANDS East Amatuli Island West Amatuli Island Sugarloaf Island Ushagat Island Rocks south of Ushacat Sud Island Nord Island

AFOGNAK-SHUYAK ISLANDS Ban Island-Shuyak Strait Pernosa Bay Marmot Island Sea Otter Island area East side of Shuyak Jest side of Shuyak Latax Rocks-Dark Island

TOTAL

TOTAL

Number of Sea Otters

0 2 0

76 200

29 0

307

18 6 3

121 0

33 26

207

Complete or Visibility Partial Count

Very good Complete ll If II

It II

II

II IJ

Good Complete II

Fair Partial If Complete II Partial II II

II

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64

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51

SEA OTTER COUNTS - KENAI June J970 to January 1971

Carl Divinyi flew aerial surveys of seals along the outside of the

Kenai Peninsula on a monthly basis He recorded all sea otters sighted

on these surveys The surveys did not cover the entire coast line and

the type of aircraft weather conditions and observers varied from one

survey to another Therefore the seperate counts cannot be compared

However when viewed in total they indicate the areas used by sea otters

and the relative abundance of otters in each area

The attached table presents the counts by broad areas

--

SEA

OTT

ERS

CO

UN

TED

ON

A

ER

IAL

SUR

VEYS

O

F K

EN

AI

PEN

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1

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1971

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125

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0

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243

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54

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55

152

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11

20

70

AERIAL SURVEY Prince William Sound

April 1970

Between April 15 and 18 1970 Carl Divinyi and Julius Reynolds fletr an aerial survey of most of Prince William Sound using a Cessna 185 The survey tras primarily to locate seals however sea otter were counted The following is a summary of the survey

I departed Anchorage on April 14 via Alaska Airlines and arrived in Cordova at 600 AM tJeather conditions prevented flying so Julius and I drove the local road network looking for anything in the way of wildlife

April 15 vleather was a little better than yesterday so we decided to try surveying along the east side of the Sound up to Valdez Arm We started the survey at Bomb Point and flew the coast and offshore islands up to Galena Bay

Observations Seal - Scattered small groups with the majority of the animals being in Port Fidalgo There were no noticeable concentrations (50 on up) of seals

Sea Otter - A total of 105 otter were counted with the majority (60) being located in St Matthews Bay The remainder of the animals were concentrated between Red Head and Knmvles Head outside of Port Gravina

Sea Lion- Small scattered groups in Port Fidalgo (60-80 total)

April 16 Weather inclement - no flying

April 17 We flew Hawkins Hinchinbrook Montague and Green Islands Very few seal- many otter and two large concentrations of sea lions (See maps for numbers and locations of seals sea otter and sea lion) We also counted 5 otter around Kayak Island

April 18 Flew those islands from Montague Strait to Icy Bay and from Port Bainbridge to Knight Island Passage (See maps for numbers and locations of seals sea otter and sea lions)

Julius Reynolds continued the survey along the mainland from Knight Island Passage to Esther Island and around Knight Ingot and Eleanor Islands

The area from Esther Passage to Valdez Arm and Culross Perry Lone Naked Peak and Storey Islands were not surveyed

SEA OTTER COUNTED Prince William Sound

April 1970

Number of Area Sea Otter Date

PORT BAINBRIDGE - ICY BAY Bainbridge Island 30 418 Evans Island 14 418 Elrington Island 4 418 Latouche Island 46 418 Whale Bay - Icy Bay 39 418

CHENEGA ISLk~D - MAIN BAY Chenega Island and Dangerous Passage 33 52 Crafton Island 2 52

PORT NELLIE JUAN - ESTHER ISLAND Blackstone Bay 1 52 Culross Island NS

ESTHER PASSAGE - VALDEZ ARM NS

GALENA BAY FISH BAY 103 415

FISH BAY - GRAVINA POINT 1 415

GRAVINA POINT - CORDOVA 1 415

HAWKINS ISLfu~D 1 4l7

HINCHINBROOK ISLfu~ 99 417

EGG ISLAND 2 417

MONTAGUE ISLAND 259 417

GREEN AND LITTLE GREEN ISLANDS 102 4J7

KNIGHT ISLAND 136 52

INGOT ISLAND 6 52

ELEANOR ISLAND 3 52

SMITH ISLAND 4 52

SEAL ISLAND 0 52

APPLEGATE ROCK 1 52

PERRY ISLAND NS

(continued) SEA OTTER COUNTED Prince William Sound

April 1970

Number of Area Sea Otter Date-

LONE ISLAND NS

NAKED ISLAND NS

PEAK ISLAND NS

STOREY ISLAND NS

KAYAK - WINGHAM ISLAND 5 417

TOTAL 892

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PRE-TRANSPLANT SURVEY OF SEA OTTERS Green 1 andMontague 1 -July 171970

On July 17 1970 an aerial survey of sea otters was made in the area

around Green Island and the adjacent coast of Montague Island The purpose

of the survey was to locate concentrations of animals prior to a capturing

operation which began the next day A Dornier twin-engine arrcraft was used

with Schneider Reynolds and Somerville acting as observers The sky was

overcast almost no wind and the sea calm However heavy rain showers intershy

fered with forward visibility and the counting conditions were only fair on

the average Conditions vmiddotere especially poor in Port Chalmers and Zaikof Bay

The count began at 210pm and ended at 345 pm

The numbers and locations of otters are shown on the map In addition

16 sea otters were seen on a superficial look at Constantine Harbor on Hinchshy

in brook Island and a pod of 35 were seen three miles east of Johnstone Pt

Most of the area included in the survey was traveled repeatedly in a

skiff over the next four days Durlng this time the weather became calm and

clear for the first time in several days and the bulk of the sea otters shifted

from the coast of Montague out into the shallow areas between there and Green 1

and to Green and Channel Islands It was obvious that many otters had been missed

Fn the aerial sur~ey and that the population in the area is quite dense A

total of 48 sea otters were captured in three days of netting

j

SEA OTTER SURVEY Salisbury Sound to Cape Spencer

August 23-25~ 1970

On August 23 1970 an aerial search for sea otters was made from Salisbury Sound to Torch Bay in the area north of Sitka A Helie Courier aircraft was used with Karl Schneider Alan Courtright and

middotWalt Cunningham serving as observers

The water on the outside coast was too rough for good visibility however it was calmer in the lee of rocks and islands and visibility was fair to good

On the initial survey only two otters were seen These were both in Surge Bay on Yakobi Island While returning after completing the count we located approximately 25 in the same area we saw the first two About 15 of these including at least one pup were in a single pod This illustrates how relatively large numbers of otters can be missed from the air The fact that none were seen in other areas does not mean that established populations do not exist elsewhere

On August 24 and 25 a search of the release area north of Khaz Bay was made by the skiff While flying into Kla) Bay for this seach two otters were seen near the old mine of Chichagof Two miners who have been working there reported that two otters have been there off and on since May

Rough weather and outboard problems restricted tl the search area~

and the effectiveness of the search in that area The following sightings were made

82470 I Adult West of Granite Islands 1 Adult South of Granite Islands

11 0A~dgtzl~4JVflup) middot1 In rocks SE of ranite Islands 82570 I Adult -n Southwest of Gran~te Islands

These sightings represent from four to seven animals

Reports over the last year indicate that sea otters regularly move in and out of Kla~ Bay and occasionally as far in as Sisters Lake bull

The fact that the count in the Khaz Bay area Has less than that made in 1969 does not mean much The animals appeared to be scattered during the present count the search did not include all of the nearby sea otter habitat and survey conditions were less than ideal

The population in Surge Bay appears to be separate and is probably well established Two otters were reported in the same location in January 1966 after only 23 otters had been released near Khaz Bay In 1969 two were seen in the same spot from the ait

While no estimate of the population of sea otters in the area can be made from the available information is evident that sea etters can be found along the entire west coast of Chichagof Island and that concentrations occur near Black Island and the Granite Islands north of Khaz Bay and in Surge Bay on Yakobi Island

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HARVEST AND TRANSPLANTS - 1970

Harvest

In May of 1970 a sea otter harvest was conducted on Tanaga Island the middotDelarof Islands and Amchitka Island The numbers to come from each island

and the distribution of the kill around each island were planned in advance Conservative estimates of the population tvere made and the total harvest numbers were based on these estimates The harvest level for Tanaga and the Delarof Islands was set at 15 percent assuming that the next harvest in these areas would be three years later so the rate of harvest would be 5 percent per year Amchitkas harvest level was set at 10 percent assuming an annual harvest and a harvest rate of 10 percent per year

The harvest plan was based on information collected on a June 1968 skiff survey for Tanaga on the 1965 USFWS aerial survey and the 1969 ADFampG aerial

survey for the Delarofs and on 1968 helicopter survey by the USFWS for Amchitka There is good evidence that all the population estimates tvere low especially for the Delarofs

Hunting was done from avo skiffs with two men each Both men shot when possible A 117-foot vessel was used for skinning and living quarters Table 1 shows the schedule of the hunt with the daily harvest numbers

Table 2 presents the projected and actual harvest numbers by area Table 3 presents the numbers killed pelts saved specimen and pelt numbers used and the sex ratio of the kill for each area Figures 1 2 and 3 show the actual numbers and locations in more detail tveather was the main factor causing deviations from the original harvest plan Possible male areas were identified on the 1968 skiff survey on Tanaga An attempt was made to concentrate more pressure on these areas to achieve a more balanced sex ratio This effort is reflected in the sex ratio of the kill A high degree of sexual segregation occurs at this time of year and without specific pressure on male areas we could expect 85 percent females to be taken as occurred on Amchitka With the effort the female percentage was reduced to 65 percent on Tanaga In the Delarofs a male area which had not been identified previously was hit and the effect was much the same

Transplants

Two transplant operations took place in 1970 The first was done in the same manner as the 1968 and 1969 transplants A total of 86 otters tvere captured at Amchitka One aircraft load was shipped out Twenty-nine were released in Oregon and 30 in vashington The animals were held in floating pens at the release sites for several days to recover from the trip Survival appeared to be excellent

The second operation took place in Prince William Sound The Canadian research vessel G B Reed came to the Green Island-Port Chalmers area with tanks built on deck Forty-six sea otters were captured Approximately 44 were alive when the vessel left Prince William Sound however only 14 survived to be released off Vancouver Island

Table 1

April 20

middotMay 1

May 2

May 3

May 4

May 5

May 6

May 7

May 8

May 9

May 10

May 11

May 12

May 13

Schedule of the 1970 harvest

1030 pm depart Homer

Midnight arrive Adak MV Active

Refuel and prepare boat

Arrive off Tanaga at noon

Complete skinning pack hides etc in pm

Delarofs

Spent am getting water at Amchitka

Clean up boat take down shelter pack gear

900 am arrive Amchitka fly to Anchorage

Killed 53

138

88

131

143

53

144

43

79

83

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(am)

(pm)

(by mid afternoon)

Table 2 Projected and actual harvest numbers May 1970

Projected Actual Tana~a Island Harvest Harvest

Bumpy Point - Hot Springs Bay 200 191

Hot Springs Bay - Twin Bays 50 50

Twin Bays - Cape Sasmik 50 ) ) 199

Cape Sasmik - Tower 120 )

Tower Tanaga Bay 180 166

TOTAL 600 606

Delarof Islands

Gareloi 20-25 0

Kavalga Ogliuga and Skagul 75-100 125

Ulak and Amatignak 35-55 19

TOTAL 150 144

Amchitka Island saved for

USFWS Counting Units 1 amp 2 100 transplant

3 30 82

4 50 36

5 120 87

TOTAL 300 205

Table 3 Details of sea otters harvested in May 1970

Tanaga Island

Total kill 606

Number of pelts saved 598

Number of small pup pelts discarded 8

Specimen numbers SO 70-4 to SO 70-609 Pelt numbers 3198-3769 and 3771-3796

(Seal 3770 void - damaged)

Approximate sex ratio - 220 males 386 females or approximately 64 percent females

Delarof Islands

Total kill 144

Number of pelts saved 138

Number of pups discarded 6

Specimen numbers SO 70-610 to SO 70-753 Pelt numbers 3797 to 3934

Sex ratio- 44 males 100 females or approximately 69 percent females

Amchitka Island

Total kill 205

Number of pelts saved 194

Number of pups discarded 11

Specimen numbers SO 70-754 to SO 70-958 Pelt numbers 3935 to 4128

Sex ratio- 27 males 178 females or approximately 867 percent females

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A combination of bad weather a lack of time to let the otters recover from their capture and problems with the holding facilities probably caused the high rate of mortality

Summary of Harvests and Transplants

Table 4 summarizes the sea otters removed from Alaskan populations in the middot last 20 years An attempt has been made to remove 300 otters from Amchitka each year since 196 7 Originally this number was set as 10 percent of the population which was conservatively estimated to toal 3000 Recent USFtfS helicopter counts of up to 3927 show that this estimate definitely tvas low In the past the removal of animals was from the southeas tern half of the island The 1970 harvest was purposely concentrated toward the northwestern end in order to spread out the pressure and to learn something about the population in that area

The numbers of animals released in all transplant attempts by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game are presented in Table 5

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Tab

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30

SEXUAL SEGREGATION IN SEA OTTER POPULATIONS

by Karl Schneider March 1971

Workers have recognized for some time that sea otters tend to segregate by sex Lensink (1962) described this segregation around the southeastern end of Amchitka Island and identified three male areasn and three female areas He speculated that females used areas of more favorable habitat and that younger males were excluded by territorial males scattered throughout the female areas The implication is that male areas contain younger or at least nonterritorial males

Marakov (1965) mentions sexual segregation around Medny Island in the USSRs Commander Islands

Kenyon (1969) gives a more complete description of the sexual segregation around the southeastern end of Amchitka and supports it with quantitative data from harvested animals

Both Lens ink (1962) and Kenyon (1969) w_ere primarily describing hauling grounds used by different sexes While Kenyons harvested animals included otters near shore neither study provided much information on the use of off-shore areas

A knmmiddotlledge of the degree of sexual segregation and the location of male and female areas is important to the management of sea otter populations Harvests must be regulated to avoid putting too much pressure on one segment of the population Then capturing animals for transplants it may be necessary to set nets in specific areas to obtain the desired sex ratio case of a localized kill of otters such as when oil spills occur it is important to know the distribution of sexes to evaluate the importance of kill to the population

In

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By the same token much can be learned about the distribution of sexes through harvest and capture operations The area from which each sea otter was taken during the harvests of 1967 1968 and 1970 was recorded Because a single hunting party might kill 30 otters over up to 10 miles of coast in a few hours it wasnt possible to record the precise locations Therefore the coast was broken up into areas and the numbers killed in each area ltJere totaled

Figs 1 - 5 show the locations of the areas letters correspond to those in Tables 1 - 4 which present the sex and age composition of animals harvested in each area The ages of the 1968 animals 1vere based on cementum deposition and reproductive condition The other samples were broken down using body size In general all males under 25 lb females under 20 lb and both sexes shorter than 100 em were considered dependent pups Females under 35 lb and 120 em and males under lb and 130 em were considered subadults These criteria probably underestimate the age of some animals Cementum deposition information for the 1967 and 1970 samples is not available yet

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35

Sexual Segregation - 2 - March 7 1971

Information from the 1967 Amchitka harvest and the 1968 1969 and 1970 Amchitka transplant capturing operations is not comparable and is not presented here However knmv-ledge gained from these operations has contributed to our understanding of sea otter distribution and this knowledge will be used in the following discussion In general this data confirms Kenyons (1969) observations for the Amchitka area although these summer and fall collections indicate a higher percentage of males in female areas than Kenyon reported from his winter and spring samples

Kenyon (1969) states that female areas are more numerous and less discrete than male areas He identified three male hauling grounds and 13 female hauling grounds on the southeastern end of Amchitka Island Recent studies involving netting and collecting animals off~hore indicate that male areas remain discrete off shore Usually these areas are off major points of land and pods of males often form in kelp beds directly off shore from the hauling ground

Female areas on the other hand are not discrete at all Any area that is not a male area can be considered a female area Some areas may be more attractive to females with pups or certain areas may be more favorable for hauling out but more females than males vill be found almost everyvthere except in the discrete male areas Therefore the main problem is to locate the male areas in et population He have no evidence of any shift in the location of male areas over a period of time so once an area is identified the information should remain useful for management purposes for many years Lensink (1962) and Kenyon (1969) correctly identified all of the male areas on southeastern Amchitka Extensive harves and netting have not turned up any new areas

Identification of Hale Areas by Pup Counts

In June 1968 a survey of most of Tanaga and Kanaga Islands was made by skiff to gather information to be used in planning harvests from those islands During this survey females with pups were counted separately from single animals No special effort was put into identifying pups fuen a female obviously had a pup it vas recorded Large pups often were separate from the females and some otters were seen only briefly in vaves kelp or at a distance Therefore many pups probably were missed and the counts should be considered minimal 6 and 7 present the counta by area A similar count was attempted by helicopter around Adak but for various reasons the results are not considered useable

From this skiff survey a number of areas were judged to be possible male areas Usually these were points or more exposed aTeas vhere relatively large numbers of single animals but no pups were seen

Shoal Point Kanaga Pass north of Eddy Rock and possibly Naga Point and Cape Tusik were suspected male areas on Kanaga Island Cape Sudak and Cape Amagalik were considered male areas on Tanaga Island and the area around Hazard Point and Lltnnoy Rock in Kanaga Pass was believed to blend with the Eddy Rock area

The best way to confirm the presence of male areas is to collect animals from these areas Harvests were conducted on Kanaga in October 1968 and on

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Tanaga in May 1970 The sex ratios of the animals harvested are compared with the skiff survey pup counts in Table 5 The harvest areas are fairly broad while male areas are usually only a few hundred yards wide As a result a harvest area may include both a male area and portions of female areas to either side

The hunting pressure is seldom even over an area Often a good portion of the kill in an area would come from one group of rocks Therefore the information in Table 5 must be tempered with some knmrledge of how the hunting pressure was distributed

Kanaga Island

Shoal Point - While a few more females than males were taken in this area the percentage of males is considerably higher than one would expect in a female area Because weather vas less than ideal much of the hunting pressure ~vas directed to areas other than Shoal Point Most of the females probably came from these areas Therefore Shoal Point is considered to be a male area$

Naga Poin~- There is no strong evidence that this is a male area While no pups were recorded on the survey the total number of otters counted was not high Fairly extesive hunting of the area under good conditions yielded a sex ratio that would be typical of a female area Therefore it is assumed that there is no male area near Naga Point

Cape Tusik - Cape Tusik was suspected as a male area more because of the nature of the area than arry counts The count included areas to both sides of the Cape which could contain females Very little of the hunting was done at the Cape Therefore judgement on this area will have to wait until more information is available Extensive hunting indicates that no male areas exist east of Cape Tusik to Cape Chlanak

Kanaga Pass - From the count it appears that a male area exists in the Pass however the harvest area extended to Hive Rock and included a large portion of female area (Fig 6) The area is too broad as is shown in Table 5 gtvhere the number of pups counted for the whole area is relatively high The results of the harvest confirm the conclusions drawn from the count While the sex ratio for the entire area is roughly equal most of the males came from Kanaga Pass Some females without pups came from that general area but the majority of the females came from the area northeast of the Pass

The Tanaga harvest showed the male areas more distinctly for several reasons The total harvest was higher the entire count area was covered in the harvest and the hunting pressure was recorded more precisely Also as will be shown later sexual segregation is more pronounced in the spring

Cape Sudak - Most of the hunting pressure was concentrated on the tip of the Cape Some pressure was exerted on the area back to~mrd Pendant Point which is a female area as shmvn by the count The high percentage of males harvested from this area clearly confirms that the tip of the Cape is a male area

Tab

le

5

Com

pari

son

of

Pup

s C

ou

nte

d w

ith

S

ex R

atio

o

f H

arv

est

Jun

e

1968

A

rea

Su

rvey

O

cto

ber

19

68 H

arv

est

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ps

10

0

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ps

10

0

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ult

s T

ota

l KA

NAGA

IS

LAN

D

Ind

ep

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er

Ind

ep

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er

M

M

F

Rou

nd

Hea

d-S

ho

al P

oin

t 0

85

3

45

6

55

4

71

5

29

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a P

oin

t-K

anag

a B

ay

66

1

64

middot

18

5

81

5

20

5

79

5

Cap

e C

hla

nak

-Cap

e T

usi

k

51

1

93

2

40

7

60

3

53

6

47

Edd

y R

ock

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e R

ock

61

3

4

35

7

64

3

47

1

52

9

Jun

e

1968

A

rea

Su

rvey

H

ay

1970

Har

ves

t

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ps

10

0

Pu

ps

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0

Ad

ult

s T

ota

l TA

NA

GA

IS

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Ind

ep

Ott

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F

M

F

Cap

e S

ud

ak-P

end

ant

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60

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6

Bar

nes

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run

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59

3

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83

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in B

ays

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77

9

23

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rin

Bay

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th B

ay

96

1

11

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3

92

7

14

3

85

7

So

uth

Bay

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em R

ock

14

8

31

6

7

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3

91

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09

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fern

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f 1

5

73

0

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0

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25

4

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em R

ock

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lak

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int

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igh

t 6

9

82

1

32

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68

1

51

8

49

Sexual Segregation - 4 - March~ 1971

Annoy RockHazard Point - The harvest did not confirm this area as a male area However the weather Vas marginal and this area was too rough to hunt Therefore the otters taken in the harvest ~vere from either side of the Point and almost none were taken from the portion that is suspected to be a male area Actually Kanaga Pass is narrow and shallmv Otters feed in all parts of the Pass when weather and tide rips permit This could be considered a single male a~ea that serves both islands

Cape Amagalik - Two harvest areas overlap this area Lash Bay-Inferno Reef and Harem Rock-Kulak Point The kill from both areas strongly reflects the presence of a male area Nost of the hunting pressure was concentrated on Inferno and this is probably the main male area

As shown above vle 1vere able to identify four major male areas from a relatively superficial pup count No male areas were located during the harvests that had not already been identified from the count ~mile more detailed observations including field identification of males would be desirable the pup counting technique appears to be an acceptable method

Identification of

Because male areas are usually on v1ell exposed points it is possible to pick likely areas from a chart and then confirm these areas by collecting animals This method avoids the expense time and danger involved in surface surveys of remote areas Actual collection of animals also provides the most concrete proof of the nature of an area

The follmving is a description of the sex composition of areas for which no skiff count data was available

Delarof Islands - No attempt 1vas made to predict the location of male areas in the Delarof Islands This is a difficult area to work in Ogliuga Skagul and a portion of Ulak Island v1ere hunted on Nay 9 1970 The hunting effort in the afternoon was concentrated around Tag Island and the rocks south of Skagul Island While the entire kill is lumped together most of the males bullJere taken in the afternoon The percentage of males taken for the day is higher than would be expected if only female areas were hunted Most likely there is a male area near Tag Island

Adak Island - Portions of Adak gtvere hunted in 196 7 and 1968 No attempt was made to identify male areas before or during either harvest The harvest figures indicate that no male areas have been hunted The Bay of Islands is a classical female area The results of extensive hunting in the Bay and around Eddy Island indicate that there is definitely no male area there The percentage of males taken between Mid Point and Blind Point vms higher than might normally be expected in a female area however the scarcity of subadult males indicates that no male area exists within this area A large congregation of otters is often seen near Finger Shoals This could be a male area but Ye have no information from there If this is a male area a few stray males from there could account for the slightly higher number of males in the kill

Sexual Segregation - 5 - March 1971

Amchitka Island - While much work has been done on Amchitka Island almost all of the effort has been concentrated on the southeastern end from Crmvn Reefer Point to a fegtv miles northvest of Rifle Range Point During the 1970 harvest an attempt was made to distribute the harvest over previously unstudied areas Two areas were suspected to be male areas These were the rocks off Chitka Point and either Bird Rock or Aleut Point Heather prevented us from hunting the area around Bird Rock and Aleut Point

The sex ratio of the kill does not indicate a male area near Chitka Point however heavy waves prevented hunting off the point Therefore a male area could be there but because no animals were killed there the kill figures would not reflect it

Composition of Female Areas

Kenyon (1969) found that 93 percent of the adult otters and 63 percent of the juvenile otters in female areas were females The May samples from the 1970 harvest agree with this The mean percent of adult females in female areas was 93 percent The mean percent of subadults not including pups was 73 percent Hmmiddot1ever in the September and October samples only 77 percent of the a-dults in female areas were female About 86 percent of the subadults were female

The seasonal difference in the sex ratio of adults is consistant for all samples These seasonal changes are most likely due to adult males moving into female areas to look for estrous females During late winter and spring breeding activity is at the lmmiddot1est point of the year and fev1er females are entering estrus In fall the number of estrous animals is at its peak and presumably more sexually mature males move into the female areas to breed It is interesting to note that there were about three times as many males in female areas in fall as in winter and spring Information from female reproductive tracts indicates that approximately three times as many females would be in estrus at any given time in fall It appears that the number of adult males in a female area is directly proportional to the number of estrous females From the data collected there appear to be 15 to 20 males for each female -vlith enlarged follicles at any given time hmmiddotrever hunter bias might influence this figure

Ages have been estimated for otters harvested in 1968 on the basis of cementum layering These age data indicate that virtually all of the males in discrete female areas are either pups (or very young subadults) or are 7 years old or older No males between the ages of 2 and 6 Here found This strongly implies some form of territorial behavior among actively breeding males Fighting among males has not been observed ofteci hmvever a mature male at the Tacoma Zoo became intolerant of a subadult male in the presence of females They frequently 11 fought 1 although neither seemed to attempt to injure the other Finally the young male had to be removed Under imiddotJild conditions the young male might have moved out of the area without repeated physical encounters

Kenyon (1969) described the breeding behavior of sea otters His description indicated that males patrolled an area and two to four males may pass a given point during a 3 or 4 hour period Vandevere (1970) observed some males

Sexual Segregation - 6 - March~ 1971

apparently defending a territory in California but from his observations t

it appeared that successful breeding males were more mobile Perhaps established breeding males are more tolerant of each other More likely they maintain a separation from each other which limits the number in an area In this way a number of males might be patrolling the sa~e area rather than each establishing a geographical territory This might be advantageous where female concentrations shift from place to place as weather conditions change

Why does the number of adult males in female areas decline when the number of estrous females declines The limiting factor may be competition for estrous females rather than territory size With fewer receptive females competition lvould be greater causing some animals to leave the area Another possible reason may be that males may have a seasonal fluctuation in fertility Lensink (1962) found that all adult males produced sperm at all seasons Limited studies since then support this however it is possible that most of these males were collected in or near female areas Also while a male may produce sperm at all seasons there may be less production at certain times of year

A contributing factor may be that males are capable of feeding in more exposed areas than pregnant females or females with pups With a reduced attraction to the female areas either middotbecause of fewer receptive females or a reduced fertility in the male the male may find it easier to obtain food away from the crowded female areas where competition for food is greatest

The fact that the number of males does appear proportional to the number of estrous females indicates that competition for these females may play an important part in regulating the number of males in female areas

The numbers of subadults in the samples is relatively low so fluctuations in sex ratio may be due to sampling errors However the greater number of subadult males found in female areas in winter and spring may be a true increase made possible by the reduction in the number of breeding males A subadul t male would have fewer encounters -vli th breeding males

There are areas within what would normally be considered female areas where subadult males tend to congregate particularly during the winter These are usually areas v7ith a ma-r-ginal food supply that are not heavily used by females Because there are fewmature females in these areas there are also very few breeding males The subadult males using these areas seem to be transients that find less pressure from breeding males there These areas are used less in summer probably because calrrer weather allows feeding farther off-shore reducing competition for food in the male areas

Examples of areas within female areas that are used by subadult males are Constantine Harbor on Amchitka and possibly Finger Bay on Adak

within Female

In the course of the harvest on Tanags in tIay of 1970 a large number of pregnant females tvith large fetuses were collected at one time Most of

Sexual Segregation - 7 - March 1971

these came from Tidgituk Island This raised the possibility that females may segregate according to reproductive condition Table 6 presents the numbers of females in each stage of the reproductive cycle by area on Tanaga Island More pregnant animals were collected in the South Bay area and within that area most of the females with fetuses weighing over 100 g were collected near Tidgiktuk Island This concentration was evident in the pup counts made in June 1968 when a very high percentage of pups was observed in the same area (Table 5) Host females ~vith large fetuses in May would pup by June

The data do not reveal any other areas of this type on Tanaga Harakov (1965) mentioned a bay on Hedny Island that tvas preferred by pregnant females and females with pups Obviously all females do not go to a specific area to pup but a female vi th a large fetus or small pup probably has more difficulty in exposed areas and seeks more protected areas This tendency can be seen by the casual observer Single animals are more likely to venture off-shore and remain in rougher waters In some areas such as Crown Reefer Point on Amchitka there is a male area near the tip of a point Males congregate directly off-shore However to either side of these male congregations there are females that are not accompanied by a pup and probab do not have large fetuses In this way we may find segregation in a single kelp bed off a point If we set a net near the outer margin of the kelp we will catch almost all males Ho~Vever if we set a net to the side of the bed and closer tmiddoto shore we will catch mostly single females Nets set in a nearby bay will catch more females with pups

Table 6 shoHs that an unusually high number of resorptions were found between Cape Sudak and THin Bays This is probably a reflection of poor physical condition of the otters in that area due to food Other data indicate a continuous decline in body condition from the east side of Adak to Kanaga Pass There is some evidence that condition improves west of Kanaga Pass The animals in the Kanaga Pass vicinity are probably in poorer condition than any of the other populations sampled Therefore the high incidence of resorptions in this area should not be considered an example of segregation within the population

ition of Male Areas

As mentioned earlier the harvest areas ivere broader than any male area Harvest areas containing male areas also contained portions of female areas Our identification of the location at rhich each animal vas collected is not precise enough to determine the exact composition of these rnale areas Kenyons (1969) data from winter and early spring harvests are more precise His data indicate that at that time of year 98 percent of the adults and 80 percent of the ju~veniles using male areas were males Of 128 males in male areas 100 gtvere adults and 28 were juveniles

Experience from netting in the summer and harvesting in fall indicates that the percentage of males remains very high all year While adult females may be very close to male areas it appears unlikely that a significant number regularly use these areas at any time year nile adult males enter female areas regularly and in predictable numbers for a specific purpose females probably ~nter male areas only occasionally in stray movements

le 6

R

epro

du

ctiv

e S

tag

e o

f S

exu

ally

Mat

ure

S

ea O

tters

Har

ves

ted

on

Tan

aga

Isla

nd

-by

Are

a -

May

1

97

0

Un

imp

lan

ted

Im

pla

nte

d

Pre

gn

ant

Pre

gp

ant

Fet

us

Wei

gh

t C

lass

A

rea

No

np

reg

nan

t N

o

No

1

2 3

4 5

Res

orp

tio

n

To

tal

A

Su

dak

-Pen

dan

t P

oin

t 12

7

26

8 30

0

2 0

3 2

2 27

( B

arn

es

Po

int-

Ho

t S

pri

ng

s B

ay

12

7 24

10

36

3

0 1

4 2

2 29

B

( (

Bar

nes

P

oin

t-T

run

k P

oin

t 1

1

6 25

7

29

0 3

1 3

0 2

24

( T

run

k P

oin

t-T

win

Bay

s 7

9 39

7

30

2 1

1 1

2 2

23

c ( (

Haz

ard

P

oin

t-T

win

Bay

s 4

3 27

4

36

0 0

1 1

2 11

( T

win

B

ays-

Her

d

Roc

k 1

2

8 28

9

31

1 1

1 Lf

2

29

D

( ( T

win

B

ays-

So

uth

Bay

2

9 45

9

45

3 0

1 1

Lf

1 20

( S

ou

th B

ay-L

ash

B

ay

6 9

29

16

51

1 0

0 9

6 1

31

E

( ((

So

uth

Bay

-Har

em R

ock

17

14

31

14

31

2 0

1 5

6 45

(

F ( (

L

ash

B

ay-I

nfe

rno

Ree

f 4

1 9

6 55

2

1 1

0 2

11

( G

(

Har

em R

ock

-Ku

lak

Po

int

12

6 29

3

14

1 0

1 1

0 21

H

Ku

lak

Po

int-

SE

B

igh

t 13

9

28

10

31

2 0

1 2

5 32

Bra

cket

s in

dic

ate

o

ver

lap

pin

g are

as

Sexual Segregation - 8 - March 1971

Juveniles of both sexes probably move more randomly than adults lihile definite sexual segregation exists among younger animals it is not as pronounced as in adults Because subadult males are tolerated less by breeding males they are more strictly confined to small areas than females of the same age This has been demonstrated in fall harvests where extensive hunting over a large area will turn up only an occasional subadult male Then ~v-hen the hunters move to a definite male area large numbers of subadult males are taken in a very short time and in a very small area

While Kenyons data indicate that 22 percent the males in male areas are subadults our experience indicates that this percentage may be quite a bit higher at least in fall Such a seasonal change in the age composition of males sounds reasonable We have demonstrated that adult males move into female areas in greater numbers in the fall perhaps tripling their numbers in these areas At the same time fewer subadult males are found in the female areas Presumably the adult males are coming from male areas and the subadults driven from the female areas move to the same male areas The result vmuld be a higher percentage of young males in male areas in fall than in spring

The total number of males of all ages in female areas doubles in fall About 13 percent of all animals taken in these areas in spring were ~ales while 25 percent ta-ken in fall were males This suggests that the total number of males in male areas declines in fall There are more adults leaving than subadults entering the area

Sex Ratio

With the sexes segregating and with the degree segregation changing seasonally and betmiddotween age groups it is extremely difficult to get a completely random harvest As a result we have not been able to determine the sex ratio of any population as a vhole Hith female areas being larger and in more protected areas where hunting is easier there is a strong tendency to take more females than males particularly in sp-ring ~Je have shown that through a concentrated effort the percentage of males harvested can be increased however even then the lowest percentage of females taken in recent harvests ivas 62 percent on Kanaga Island in October 1968 Similar results occur in capturing operations for transplants

On all islands studied there are fewer male areas thltm female areas The male areas are very small usually only a few hundred yards wide The male feeding areas off male areas are also very narrow although they may extend farther off-shore In short that portion of the total available sea otter habitat included in male areas is very smalL

All evidence indicates that there are more females in the population than males While the percentage of is probably greater than that indicated by most harvest statistics it appears that at least 60 perc2nt of the sea otters in the populations studied are females

There are probably several factors contributing to this uneven sex ratio First our reproductive data indicate that 56 percent of the pups are females at birth Secondly Kenyon (1969) has found a higher mortality rate among

Sexual Segregation - 9 - March 1971

newly weaned males There is also some evidence that males might not live as long as females These factors could easily account for the preponderance of females in the population

It is possible that the higher rate of mortality in juvenile males is in part due to the breeding behavior of adult males and the resulting sexual segregation Juvenile males tend to be restricted to male areas or areas of marginal feeding habitat where there is little competition from breeding males During the ltvinter the feeding activity of young males would tend to be confined to small areas close to shore in male areas Therefore they may be subjected to greater competition for food than females of the same age which have a broader area in ltvhich to feed Kenyon (1969) found that subadult males vere less hardy in captivity than subadult females Therefore we can not adequately evaluate the influence of segregation on mortality

The sex ratio probably varies from one population to the next Very little juvenile mortality occurs in expanding populations eliminating that source of sexual selection However there is some evidence that males are more numerous among the first animals to repopulate a new area While a high percentage of males might be found on a newly repopulated island the loss of these males tvould alter the sex ratio of the parent population even though juvenile mortality might not yet be a significant factor

If the nThuber of breeding males is regulated by the number of estrous females an even sex ratio might produce a surplus of sexually mature males This surplus would not be beneficial to the population and could be detrimental This would permit the situation of an unbalanced sex ratio to evolve

The segregation of sexes and ages and the composition of sea otter populations is complex Unless we can understand the situation we cannot fully evaluate the effects of mortality or habitat changes

REPRODUCTION IN THE FElIALE SEA OTTER

by Karl Schneider Narch 1971

A total of 1358 sea otter reproductive tracts collected between 1967 and 1970 have been examined Ovaries were sliced with a razor blade and examined macroscopically Uteri Here macroscopically examined both internally and Each tract vas classified as to its stage in the reproductive cycle

The follovJing criteria were used in classification

Nulliparou~- Horns of uterus small thin and smooth no corpora lutea or corpora albicantia although there is occasionally evidence of a past ovulation but no evidence of implantation

Nultiparous - Uterus thicker corpora lutea andor corpora albicantia present Includes primiparous animals (these are not readily separated from multiparous animals)

Anestrus - Largest follicle under 3 5 rnm no corpus luteum

Proestrus - Follicles over 35 mm

Es - Follicles approximately 10 rrm

Implan~elt_ Pregnant_ - Visible s~velling in uterine horn usually with fetal membranes embryo or fetus visible Corpus luteum usually over 10 mm

- Uterine horn sti11 noticeab enlarged fresh roughshy~=-=--=~

al scar newly formed corpus albicans with some luteal tissue remaining

The appearance of the uterus ~ras used to confirm the classification The thickness arrd of the horns thickness and consistency of the uterine walls coloration of the of the horns and condition of the rugae change with each stage In most cases it is possible to guess the condition of the tract by a superficial examination of its exterior This appearance was used only to confirm what was indicated by structures in the ovary hmvever

Table 1 smnmarizes the reproductive condition of the sexually mature females in the that are enough for comparison throughout the year In the following discussion reported by Sinha et (1966) and Kenyon (1969) are included 7here possible The January and Harch are from these studies

Tab

le

l Re

p iv

e co

nd

itio

n o

f se

xual

ly m

atur

e se

a o

tters

_

IF

etus

Wei

--------------+

__

_

ln~a

ctive

ov

arie

s 1

Act

ive ovari~

ht C

Jas

s

jmiddot

I P

ost

-I P

roes

tru

s U

nim

pl

lmpl

D

ates

L

ocat

ioQ

1A

nest

rus

Part

um

Res

orpt

ion

l +

E

stru

s P

reo

Pr

eQ

2 3

lJ

S

~

4-8

1970

T

anag

a 1

51

41

0

10

104

17

8 10

33

30

4 (1

68)

(1

35)

(3

3)

(3 3

) (2

89)

(34~

(5

6)

(27

) (3

3)

(11

4)(

10

9)

May

9

19

70

Del

arof

Is

18

14

1

4 13

27

10

0

3 6

8 77

4

) (1

82)

( l

3)

(5

(16

9)

( o

) (1

30

) (3

9)

(78

)(1

04

)

10-1

219

70 A

mch

ltka

I

34

18

3 8

27

48

5 6

2 23

12

13

8

( 0

(24

6)

(13

0)

(22

) (5

8)

(19

6)

(34

8)

(36

) (4

3)

(14

) (1

67)

(8

7)

~lune

23

middotmiddot ~middot~~middot~Amchitka

1

17

2 1

2 4

18

1 2

3 5

7 44

A

ugus

t 13

196

8 (3

86)

(4

5)

(23

) (4

5)

(90

) (4

09)

(2

3)

(45

) (6

8)

(11

3)(

15

8)

July

6-

~dgtAmchitka

1

17

1 0

4 14

10

0

1 1

2 6

46

Aug

ust

819

69

(36

9)

( 2

0 2

) (8

6)

(30

4)

(21

9)

(22

) (2

2)

(43

)(1

32

)

Sep

t 1

0-17

A

dak

72

5 0

16

31

40

6 6

8 9

11

164

1967

(4

39

) ( 3

1)

(98

) ( 1

89)

(

4)

(3 7

) (3

7)

(49

) (5

5)

(6

7)

Sep

t

25

-A

mch

itk

a l

55

6

0 18

19

17

4

0 0

0 3

115

OcL

6

19

67

(4

7~8)

(5

2)

(15

7)

(16

5)

(14

8)

(35

) (8

7)

(26

)

Oct

o 1

2-1

5

Kan

aga

I

58

6 1

13

31

31

4 4

7 11

5

140

1968

(4

13)

(4

3)

(o 7

) (9

3)

(22

2)

(22

2)

(29

) (2

9)

(50

) (7

8)

(36

)

Oct

18

-20

A

dak

l

46

6 0

1 24

13

2

3 0

4 4

100

1968

(L

16 0

) ( 6

0)

(11

0)

(24

0)

(13

0)

(20

) (3

0)

0)

(40

)

Num

bers

in

re

nth

esis

are

pe

rcen

t se

xu

ally

mat

ure

fem

ales

F

etus

wei

ght

clas

s 1

=

0-l

g

2 =

1-l

Og

3

= 1

0-lO

Og

4

= 1

00-lO

OO

g

5 =

Tra

nsp

lan

t m

ort

alit

ies

(all

o

ther

sam

ples

fr

om

har

ves

ts)

Reproduction - 2 - March 1971

There may be some problems associated with comparing samples taken from different islands and in different years but allowing for sampling errors the data for the most part fit a definite pattern indicating that the annual cycle remains fairly constant The one outstanding exception is the number of pregnant animals in the t1m summer samples In 1968 there were many more implanted pregnancies than unimplanted pregnancies and in 1969 the situation was reversed hmvever the total number pregnant was almost identical These are the t s being relatively small collected over a longer of time and consisting of transplant mortalities An average of the two samples fits the pattern established by the other samples Relatively large samples are necessary because all stages of reproduction are found at all times of the year The period from November to early January is not represented hmmiddotJever things are changing so rapidly during that period that any information from that time may be confusing unless a large sample was collected in a very short time

The information in Table 1 is shmvn graphically vJith Kenyons (1969) winter information in Figs 1 and 2 It has often been demonstrated that sea otters breed and pup at all times of the year A number of observers have noticed that roore pups are born in the spring aJd sumrrter than at other times and there have been conflicting reports about breeding times Kenyon (1969) was the first to demonstrate seasonal changes in pregnancy rates but only his January and Yarch sallples gtvere large enough to be at all reliable

Figs 1 and 2 demonstrate that there are cons le secsonal in the numbers of animals in each s reproductive cycle There is a definite period of increased bree activity and a definite period of increased pupping Hmmiddotlever ~ some anii1als are in each s at all times of the year and these periods of increased breeding and pupping do not have distinct boundaries

Each curve is influenced by two factors so it is difficult to isolate the magnitude of any single factoc For example the of implanted pregnancies vill increase ss blastocysts and decrease as births occur The birth rate may be increas but if the number of implantations increases at a greater rate the implanted curve will still rise The animal remains in the anestrus implanted pregnant~ and unimplanted pregnant categories for a relatively long time and there may be considerable carry-over from one sample to the next Because the animal a relatively short time in the proestrus-estrus and post partum categories there is little if any carry-middotover from one sample to the next and the changes betgtmiddotTeen are more likely to be absolute changes The mean fetus weight class prob does not mean much because it does not take the actual number of animals in each class into account ~hen there are many implanted pregnancies ard the mean lmiddotTeigh t class is lovJ there actually may be more Class 5 fetuses in the population than vhen there are fetv irr1planted pregnancies and the mean middotleight class is high Therefore the actual percentages of all sexually mature females middotlith Class 1 ald Class 5 fetuses have been plotted in 3 These curves represent the actual number of fetuses in the population Because tha sample sizes become quite small vhen the fetuses are divi(12d into classes and the surruner sample is believ2d

2

~ _s ~

~

c U)

~-1

r

~J

lt

c])

Reproduction - 3 - Narch 1971

to be biased tvo curves for each class are shmvn The solid curves approximate the data while the broken curves represent subjective adjustments to correct for sampling errors

cussion of the Annual

The number of animals entering estrus rises in the fall At this time the pregnancy rate is at its lowest point and a high percentage of anestrous (nonpregnant) animals are in the population Breeding activity increases and at this time the number of sexually mature males in 1female areas increases

The number of unimplanted pregnancies begins to rise sharply in October and November At the smne time the nlli~ber of anestrous animals drops as these animals enter estrus breed and become pregnant Thj_s is the peak breeding season

The birth rate is at its louest this period as is shovm by the lmv number of post-partum a1imals and the lovr number of large fetuses

By late November and December the pregnant rate is increas even though the birth rate is slightly This is due to ne7

blastocysts implanting as is shown the increase in Class 1 fetuses

By January the period of t breedLng activity has declined and the number of unimplanted animals ceaches a peak as the number of blastocysts implanting equals the number of The lanted pregnancy rate then begins to declLne as fevJer animals breed and more implant The greater rate of antation is reflected in a rise in the Class 1 fetuses as well as an overall increas in the er of lanted pregnancies However the rate of implantation is partially obscured by an increased birth rate removing animals from the total of lanted pregnancies This increase in the birth rate is shmm in the rise in post-partum females and a rise in Class 5 fetuses

This condition of a low rate of breeding high rate of implantation and increasing rate of birth lasts from February to Hay

At this point He come to the least reliable data The sumner are smaller gtJere collected over a period of seven weeks and were from animals that died as the result of capture and handling

Therefore tvhile the data shaH that the post-partum rate decreases sh indicating a reduction in the birth rate~ the number of Class 5 fetuses

a peak indicating a very high birth rate The true birth rate is probably some1middothere in betigtJeen Females 1-ith very young pups (hence are post-partum) are vlary and probably less likely to be caught in nets Females with fetuses appear to be more likely to from handling In several cases~ twisting of the reproductive tract by a fetus vJas the actual cause of death Anestrous females (including post partum) on the other hand are less likely to die

Reproduction - 4 - March 1971

There is no question that the birth rate remains high during June and July Kenyons (1969) sum11er sample is limited but shows a large number of Class 5 fetuses Again the sample is biased in favor of pregnampIt animals but not to large fetuses only His field counts as vJell as observations by most other observers indicate an increase in the nUIlber of dependent young througbout the surnmer The number of Lmted pregnancies declines throughout the suTimer tvhile the unimplanted pregnancy rate declines at a slmver rate and breeding remains fairly constant at a lm-v level This indicates that through June there are more implantations than conceptions hmvever the birth rate is substantially greater than the implantation rate By late July and August there are relatively fe1r implantations taking place as shmm by a leveling of the unimplanted pregnant rate and a lotr number of Class 1 fetuses The number of Class 5 fetuses declines as a result of births during June July and August

Therefore the season of greatest pupping may be quite broad running from April through August possibly with the peak occurring in late May or June Until a better summer sample is available tve cannot pin dmm the peak of this pupping season with certainty (See Fetal GrmmiddotJth Rate for further discussion on breeding and pupp peaks)

If the information were precise we could compare subtracting numbers of pups born and numbers of conceptions to deterrnine the actual percent of ferrales breeding implantLng ltmd pupping in each month Unfortunately the data are not Each curve is influenced by more than one factor and ue do not knmmiddotr for sure the magnitude of any of these factors Hmmiddotrever by comparing some of the major samples e may be able to get a rough idea of how much higher the birth rate is at one time of year than another

The combined Mty and the combined September and October samples provide the best comparison Both are large samples One occurs near the peak of pupp and a lmv point in breeding and the other is near the low point in pupping and the peak of b

The number of Height Class 5 fetuses~ post-partum femaless and proestrousshyestrous females come closest to represent a particular event The time spent in each category is comparatively short

There were approximately three times as many Class 5 fetuses in Hay thanmiddot in Septerqber-October There Here also about three times as wany postshypartum females in May In tember-October there were two and a half to three times as many proestrous-estrous animals as in Hay It ~middotwuld appear that the peaks of breeding and pupping are roughly three times as high as the lowest points

At the lmmiddot12st period of pupping 2 to 3 percent of the females have Class 5 fetuses and should pup middotJithin a month If vm assume the post-partum stage lasts L 5 to 2 months (see Recovery of the Uterus) it also appears that 2 to 3 percent pup in the ltwrest months of pupping

Reproduction - 5 - March 1971

Similarly about 3 percent of the females have enlarged follicles during the lmvest period of breeding He do not knmv how long this period lasts hovever

At the peak of pupping about 10 of the females have Class 5 fetuses and should pup in the next month About 15 are post partum indicating about 8 births per 100 females in the preceding month

These percentages are prob2bly slightly high because of hunter bias avoiding females with pups It appeatmiddots that at least 2 to 3 percent of the mature females breed and 2 or 3 percent have pups in any month of the year In the peak breedingmonths such as September and October perhaps 7 to 10 percent breed and similarly in peak pupping months sud1 as Nay 7 to 10 percent of the mature females pup

Gestation Period

Barabash-Nikiforov (1947) listed the gestation period of 8 to 9 months This apparently was based on the case where a fentele mated in captivity as reported by i-Ialkovich (19 37) According to Barabash-Nikiforov a pup was born 8 months later although it died While it was fully formed~ it may have been premature

A number of births have occurred in captive animiddotmals held by the of Fish and Game in recent years In all cases the pu-ps ~-ere sc-middot1all usually less tha1 1600 g but still than the smallest pups found in the -rild All died after birth

The gestation period of 9 months has been repeated in the literature but apparently these statemsnts are based on Barabash~Nikiforol (19lf7)

Lensink (1962) also estimated a gestation period of 8-9 months based on field observations He felt that the peak of breeding was in August or September and the of pupping ~Alas in l-1arch or April

It has been demonstrated that the gestation of sea otters a relatively long period of tation (Sinha 1966 Kenyon 1969 and present study) Therefore any attempt to estimate the length of gestation by examinatioD of tracts must take both the unimp1anted and Janted periods into accour1t

Kenyon (1969) presented an estimate of the implanted period by assuming that the cube roots of fetus weights fall in a straight line after the embryo is established (Hugget and Hiddas 1951) and by that the European otter and Aluerican river otter would have similar fetal growth rates By plotting tenn fetus on a line established by the European otter he estimated an gestation of about 120 days not including the period for establishment of the ewbryo

Kenyon (1969) then an estimate made by Dr D G Chapman using the method of Hugget and ~-Jiddas (1951) v7here a regression line is fitted

Reproduction - 6 - March 1971

to the cube roots of the raeail of the mean weights of fetuses in each of five vreight clas3es plotted t the mean time of year

From the regression coefficient ob he estimated an implanted period of 154 days By taking the mean of the pregnant animals that are unimplanted pregnant estimated the tmimplanted period as about 75 months 1Ulmv-ing a half month for establishment the embryo he estimates a total gestation of 12-13 months

The data used by Chapman included a January-February sample of 26 a March-April sample of 49 and a Nay-August of only 9 The surtmler sample is more biased than the others and quite small yet the estimate relies heavily on this sa~ple when fetus size is the greatest

Table 2 presents a neTw- estimate of the length of the implanted period using the method used by Chapman but including data from the present study in addition to Chapmans data Table 3 presents a similar estimate of the length of the unimplanted period

The following is a comparison of the two estimates

Unimplanted Period 230 days 66 days Establishment of Embryo 15 15 days Implanted Period 154_~~Yrgt~ _73 d~s

Total Gestation Period 399 d2ys 154 days or about 13 months 5 months

Obviously there is some problem tiith the the technique or both A similar calculation mcde before the Hay sample had been collected produced an estimate of 109 for the implanted period and about 8 months for the total ation period

This method of calculation assur1es that there are definite peaks in breeding and pupping If breeding and pupping occurred evenly throughout the year the average fetus size 1vould remain constant thruughout the year and the technique would not tvork An ideal situation v10uld be when all individuals breed and pup 1vi thin very short periods Sea otters do have but are not well defined Very large s v-ould be required in such a case because there are ahvays fetuses of all sizes and the periods of maximum breeding are broad

Obviously Chapmans sample vas inadequate llthile the number of fetuses available for the latest estimate seems large and are well distributed throughout the year the a_ctual nu~ober in any cne fetus vJeight class at any one time of year is small A look at Table 1 shows that there is little consistency in many of the classes (Fig 3 shmvs that Class 1 and 5 fetuses do fit a pattern)

Table 2 Estimated Length of Period

Percent of Fetuses in each Height Class

Time of Collection After January

1 month 31 15 15 38 0

3 months 18 12 22 35 12

5 months 18 8 8 36 30

7 months 3 8 16 29 45

9 months 20 7 10 40 23

10 months

14 18 12 33 23

---~--

r1ean Time in Nonths After January 50 57 53 58 70

Cube Root of Umveighted Mean 063 17 36 7lf lllf

Specific Grmth Velocity 0169day EstirGated Implanted Period = 73 days

Table 3 Length of Unimplanted Period

Months After Unimplanted Unimplanted Percent January Pregnant Pregnant Unimplanted

1 month 35 26 58

3 months 49 49 so

5 months 128 179 42

7 months 22 37 37

9 months 50 57 47

10 months 55 44 56

Unweighted

48

Reproduction - 7 - March 1971

All of the estimates very heavily on su111mer samples 1vhen the fetuses are largest These sauples are knmm to be biased probably in favor of larger fetuses There are more fetuses at this time of year but it is exaggerated in these SCilllples

The latest estimate is not necess better than Chapmans even though it uses more and larger samples The velocity is twice as high as that calculated for fur seal and a 5 month gestation period is not consistent with other information available

This technique for estimating the lanted gestation period is not capable of providing a reliable estimate with the available data Therefore the 12 to 13 month estimate and any calculations based on it are meaningless

Much of the discrepancy lies in the estimate of the unimplanted period The present estimate that 48 percent of all pregnancies throughout the year are unimplanted based on an even distribution of relatively large samples and is probably more accurate than Chaprnan 1 s estimate of 60 percent

There are several other ways of guessing at the length of gestation Kenyons (1969) estillate of an lanted period of 4 months using the cube roots of term fetus weightscannot be relied upon entirely but it may be closer than the other estimates If gtmiddotJe estimate the uninplanted period from this using the 48 percent ted factor and allow for the establishment of the embryo we get a total gestation of about 85 months

Lensinks (1962) method of detennining the and peak pupping periods and taking the time between the peaks as the total gestation period has some merit TJnile Lens ink tried to determine these peaks by field observations we can determine them more accurately using the condition of reproductive tracts

Lensink felt that the peak of breeding middotlas in August or September The data in Fig 2 indicate that he may seen the beginning of the period of est breeding activity but that the probably in October or possibly November Lensink estimated period in larch or April Again the data indicate that he v1as s the beginning of the period of greatest pupping but the actual peak is probably in June or July At this time the number of Class 5 fetuses is greatest The nucJber of post-partum females probably and the number of implanted pregnant females is dropping from its If the gestation period of

11 11the average animal lasts from October or November to June or July we get a period of 8 or 9 months tThile Lensink 1 s was early 1 the duration agrees vith the present data

The peak of implantation when the greatest number of blastocysts are implanting can also be roughly estimated from 1 and 3 In February the number of unimplanted middot is dropping and the number of implanted pregnancies at this time th2 number of Class 1 fetuses probably reaches a

Reproduction - 8 - March 1971

Therefore it appears that the unimplanted period lasts 4 to 5 months and the implanted period from 4 to 5 months 1vi th a total gestation period of 8 to 9 months It should be recognized that these estimates are based on general trends and not distinct Therefore they are approximate However the relative length of the unimp and implanted p2riods agrees with the percentages noted previous and this estimate agrees vlith the one based on Kenyons (1969) comparison vdth other species of otters

If the gestation period ~Jere 12 months the percentage of pregnant animals would remain constant throughout the year If it vJere only slightly longer or slightly shorter there would be only a slight fluctuation unless there were a distinct breeding period Fig 4 indicates a substantial fluctuation indicating a gestation period substantially shorter or longer than 12 months

The peak of pregnancy occurs in February after the period of t breeding activi The luH point on the pregnancy curve occurs after those animals giving birth dur the period of t pupping have pupped That is the t point on the pregnancy curve occurs after the main breeding and the lmves ~ point is after the main pupping period These periods actually last for 4 or 5 months so these points are well after the peaks of breedtng and pupping

The time betbulleen the peak when the pregnant and the lmr point Hhen the fewest are the average gestation period This is about 85 months (Fig 4) The anestrus curve (Fig 1) shmvs the same thing betng a negative of the pregnancy curve

Hith three different methods we have estimated a gestation period of 8 to 9 months Hith the unimplanted period roughly equal to the implanted period This agreas bullmiddotlith the observed period in captivity (BarabashshyNikiforov 194 7) and with field estimates Any gestation period differirg greatly from this does not fit the data Conceivably one could apply a 20 to 21 month period to the SaTEpound information however if this 1vere the case females would have to breed vJhile accomp by a pup or be on a four year reproductive cycle This prospect stretches the imagination

There is a definite possibility that the gestatton period varies in sea otters as in land ot te~cs This most likely -middotwuld occur through variations in the unimplantec1 period The above discussion refers to the average gestation period Until better information to the contrary is obtained we must continue to assume that the average period is 8 to 9 months

Fetal Rate

the estimates of the gestation curve are corrects Kenyons (1969) Fig 4 should approximate the grouth curve of the fetus Fig 5 duplicates this and shmvs the actual Heights Fig 6 shous the same information more graphically It is possible to esttmgtte the of time in each v7eight classlt Fig 6 does not take the period for establishment of the embryo into account so Height Class 1 may be longer than shmm perhaps 15 days if we accept Chapcnans (Kenyon 1969) es e

--

3 0

~

shy

----- -shy --middotmiddot~middotmiddot

___ ___1__

_

(middot-)

_ ft -) J ~)___

~-

3

Reproduction - 9 - IYarch 1971

The short time span of Classes 2 and 3 explains the relatively small numbers and the lack of a consistent pattern found in those classes (Table 1)

Using this curve it should be possible to predict the time of birth and with less precision the time of Luplantation and conception of any particular fetus By plot the estimated birth dates of a large number of fetuses rve should be able to drav curves Ttlhich approximate the breeding implantation and pupping rates of the population throughout the year

Unfortunately there is overlap bet1veen samples collected at different times of year Some of the samples are too small to provide a comparison We vould have to give eight to each in order to combine them Therefore only the ttay September and October samples are used lfuere the September and October samples overlap t1lt10 separate points are used Fig 7 presents the estimated birth dates grouped by month Fig 8 separates the dates into half month groups February and Narch are not represented and April is an estimate based on post-partum females rather than fetus size (see Recovery of the Uterus after Parturition)

The peak in this curve may be exaggerated It has been shmvn previously that fetus size is less consistent bet-Jeen samples than the other categories (Table 1) because the sample size of each grouping is small Therefore the exact shape of the curve may not be correct If however we assume the approximate of the peak is correct we can construct a model of the based on our estimates of the gestation period besed on a fetal grmJth rate estimated from this estimated gestation Therefore our reasoning would be some~hat circular if -Je used thjs to prove the estimate Hmvever we can see how this model fits the rest of the data A good fit would tend to support the model and the estimates on which it is based

For this model He use the curve in 8 to represent births We assume a gestation period of 8 months -rith the unimplanted and implanted periods each lasting 4 months TheTefore v8 draw curves identical to the birth curve but moved backward 4 months for implantation of the blastocyst and 8 months for conception (Fig 9)

A comparison of the peaks in Fig 9 with the data in Figs 1 2 and 3 and the narrative b on those indicates a fairly close fit The data indicat that the peaks may ~ctually occur slightly later than indicated in 9 but in the timing and distances bet7een the peaks fit

The sharpness of the birth curve indicates that the breeding in1plantatimiddotm and pupping peaks mey be more distinct than indicated by the curves in Figs 1 and 2 It is possible to fit curves with more abrupt changes to the data Fig 10 shows the same data lith the curves drawn to more closely fit the model tfuether these curves are more accurate or not is open to debate The data on Hhich the birth curve is based is not that strong The numbers in each month are relatively snall If such distinct did occur it seems likely that field observers ~vould have seen the effects Nost field observations indicate broader peaks that are not gtvell defined

2

G

X

middot ~ -- (-- ) r~ ~ - - -- ~

yen~ r- ~--~

3

i__middot -~---

~

gt

- cshy

Reproduction - 10 - March 1971

~e of _Sexual Maturity

Tables 4 - 7 list the frequency of nmnbers of corpora albicantia and corpora lutea found in females of different areas It should be recognized that the ages may not al-1ays be exact Animals ATTichi tka to become sexually mature Hhen about 4 years old It appbull~ars that more animals may become mature earlier on Adak The information is too limited to dratmiddotJ any definite conclusions from this hmmiddotJever the food availabili ty is probably better at Adak and an earlier age of sexual 1naturity may be a response to better nutrition

Ovulation

From Tables 4 - 7 it can be seen that the maximum number of corpora albicantia for each age roughly the numb~r of years after sexual maturity that some animals ovulated once every year after maturity It can be assumed that many corpora albicantia are invisible macroscopically after several years particularly if they are not remnants of a corpus luteum of pregnancT Therefore it Ls possible that most females ovulate every year This poss ili ty is also supported by the high incidence of large follicles in lactating females (TabL~ 10)

faximum

Tables 4 ~ 7 indicate that at t some females continue to breed at a very old age The samples are teo small to determine 1rhether a t number become unproductive at any point

Ovulation

Many mustelids are induced ovulatoTs

On September 14 1967 a female vJas tsd Hhile copulating She appeared to have Oilnlated shortly before Because sea otters may copulate several times over a period of two or three (Kenyon 1969) ovulation may have been induced by a previous

ficance of

Delayed lon is a charact8ristic of the reproductive cycle of most mustelids Several theories on the ccdv of a delay have been advanced Most assume that spring is the most favorable time of year for survival of nemiddotJborn young but that either a wir1ter breeding season is not favorable or that the presence of a Ttlale vhich occurs only during the

season is for survival of the young of the previous pregnancy (1963) discusse the evolution of delayed implantation in mustelids and points out that there are exceptions even in arctic areas middotJhere time of birth be consLdered more may be born at any tin1e Scheduling the and made possible by delayed mey be advantageous but is not necessary

----

a - o lt bull y bull bull l w laquo bull _ ~ ~

Table 4 Frequency of occurance of Numbers of Corpora Albicantia Amchitkll - June-August 1968

N U M B E R 0 F C 0 R P 0 R A A L 8 I C A N T A

I

~I -AGE -~ 1 I 1084 6 92 5 7(Years) 1 middot shy

- 0-l - ~~ ~- middot~middot- middotmiddotmiddotmiddotmiddot

1-2

2

3

4 1 I

2 (1)

6

middot5

1 (1)1

2 (l) 17

1 ( 1 )

2 ( 1)

18

l (1) I ( 1 ) 9

10 1 ( 1) L ~j( 2 1 ( 1 )

2 ( l) 1

12

11

1 (1)

I I (I)

1

2 ( 1)

13

1 ( 1 )

2

114

115 I116

17

I 18

I I

I

119 I

Numbers inside parentheses =Number of individuals with corpus luteum

11

Table s Frequency of oc~urance of Numbers of Corpora A1bicantfa _Kanaga - October 1968

N U M B E R 0 F C 0 R P 0 R A Al B CANT IA - I I I

AGE 84 I I

5 I l 6 7z 3(Years) 1

l rbull ~middot--

--- shy

0-l

J l-2

~ 2

3

4 2 ( 1)

5 3

Ibullbull 6 6 (3)

4 (4)7

8 4

3 9 2 ( 1 )

j iI 10

I 1 (1)11

12

13

14

1 ( 1)

16

15

17

18

Numbers irisi

-

J

(Plus 2 with corpus 1 uteum but no corpora a 1 b i cant i a)4(1)

1

1 (1)

5 (1)

10(5)

3

3 1

3 (2)

2 ( 1) 3 (3)

1 ( 1)

2

1 ( 1 ) 5 ( 1)

1 ( 1 ) 3 ( 1)

1

3

2 ( 1 )

1 (l)

1 (1)4 ( 1)

2) 1 (1) 3 (3) 4 (1) 12 ( 1 ( 1 )

1 ( 1)

1

2

1 ( 1)

1 ( 1)

1

I I parentheses = Number of individuals with corpus luteum

11

Table 6 Frequency of occurance of Numbers of Corpora Albicantia Mid Pt-~Blind Pt~ Adak- October 1968

N U ~1 B E R 0 F c 0 R P 0 R A A L B I C A NT I A

-1AGE

~

9 1084 75 6Years) 1 2 3 - shy ----- --middot shy~--

- _ Q~L

1-2

(Plus one with_corpus luteum but no corpus a1bicans)2 1 l I I I I I ) (One with corpus luteum but no corpus albicans 3

4 2 (1)

l5

16 1 (1)

1 1)7

8

2 (1)9

10

111

12

I13

14

15 1 (1) 16

17

18

1 ( 1)

1 (l)

1

1

1

2

1

1

1

1(1 )

1

1 (1)

1 (1)

1

I

1

1

1

1

-I

Numbers Inside rentheses = Number of individuals with corpus luteum

1

Table 7- Frequency of occurance of Numbers of Corpora A1bicantia Three Arm Bay - Bay of Islands Adak - October 1968

AGE (Years)

Q-1

t-2

2

3

4

s 6

7

8

9

10

11

12

13

14

15

16

17

18

Numbers

N U ~ B E R 0 F C 0 R P 0 R A A l B C A N T I A

~L 3 4 slE - 1middotshy

7 1~8 __J~l_11 shy

1-

One -Ji th corpus 1uteum but no corpora _a 1bicantia I I I I I I I

One vJi th corpus luteum but no corpora al bicantia

12 I I (Plus one with corpumiddots luteum but no corpora albicantia)1 (1) 3

1 (1) 2 (1)

1 2 (2) l 1

1 (1) 4 (1)

2 (1)

2 1 )

1 ( 1 ) 1

2 (1)

2 (2)

l22 3

3 ( 1)2 21 ( 1)

2 ( 1)

2 ( 1)

l ( 1)1

1 1

1 I ll

I

I I

inside parentheses Nurnber of individuals with corpus luteurn

Reproduction - 11 - March 1971

The sea otter lives in an area of relatively uniform temperatures However storms tend to be more frequent and violent in fall and winter Survival may be better in the late spring and sunrmer Therefore there may be some advantage to birth in the late spring However there does not appear to be a great deal of advantage to a breeding season at any particular time of year

Wright (1963) pointed out that most musteHds lant after daylight begins to increase usually around February This is the period when sea otters implant at the t rate If the time of implantation is influenced by daylight the period when the greatest number of births occur could be shorter than the equivalent breeding period The data do not show this although the possibility cannot be ruled out

The fact that substantial numbers of sea otters are breeding implanting and pupping at all times of the year indicates that hile there may be some advantage to certain events taking place at a particular time of year that advantage is not great enough for a distinct breeding season to evolve

Sea otters exhibit certain characteristics that are comnon to most mustelid breeding cycles but these chfracteris tics are not as well developed as in most es

It is interesting to note that Hright (1963) correlated the molt of Mus with the implanted period TI1e molt began around the

ion and ended around parturi tioD

Sea otters molt all year and some are implanted pregnant at all times At this time e cannot say uhether any between molt and pregnancy exists

Fetal

Kenyon (1969) found that the number of caudally presented fetuses roughly equalled the nunber of cephali presented fetuses He suggests that this indicates a lack of tatim1 for birth occurring in water and infers that sea otters must give birth on land

Fetuses of all 1veights in the present study also appear to be randomly oriented however 97 Class 5 fetuses (1000 g and over) showed 60 percent cephalic ion while only 40 percent shovJed caudal presentation

The orientation of large fetuses should be a better indicator of orientation at birth Smaller fetuses may position Therefore it appears that more sea otters are born head first than tail first

If you accept Kenyons premise that caudal presentation is necessary or at least beneficial to birth in water this more recent information supports the supposition that birth occurs on land The little information available indicates that birth does occur on lando but there have been unconfirmed reports of birth in water

Reproduction - 12 - tltIarch 1971

At least some sirenians are born head first indicating that caudal presentation is not necessary North of Unimak Island a large population of sea otters lives off-shore There is little evidence that any numbers come ashore It seems likely that many of these animals are born in the water

Of 305 implant pregnancies 138 fetuses (45 percent) had implanted in the left horn and 167 (55 percent) in the right horn Most of the difference was in the 1970 Amchitka sample ~fithout this sarnple 48 percent implanted in the left horn and 52 percent in the right horn Kenyon (1969) found an equal number in each horn If a real difference exists it is probably exaggerated by the 1970 A~1chitka sample

Out of 305 implanted pregnencies the point of implantation was on the same side of the reproductive tract as the corpus luteum in all but three instances In one case the corpus lutaum was in the left ovary and the fetus in the right porn In the second case the corpus luteum in the right ovary but the fetus was in the left horn In the third case there was a corpus luteurn in each ovary and tHo fetuses (of different sexes) in the left horn

It appears that blastocysts rarely cross from one horn to the other

Because there is usually a long period of time parturition and the next est_rus is little if any inhibitory effect on ovulation caused by the corpus of the previous pregnancy The ovary producing the largest follicle appears to be random and there does not appear to be the tendency for a pregnancy to be from the opposite ovary from the previous pregnancy In many cases there are many more corpora albicantia in one ovary than in the other

When a loses a pup shortly after birth and enters estrus before the uterus has conpletely recovered and before the corpus luteum has completely degenerated there may be some inhibitory effect Of the ll+ such cases identified six had large follicles in the same ovary as the new corpus albicans and in the opposite ovary It appears that any inhibitory effect by the corpus luteum only lasts for a short time

A few cases bullv-ere found vJhere one ovary was not developed or othenvise not capable of producing ova Because one ovary may support repeated pregnancies such animals may be as productive as those ~vith both ovaries active

Kenyon (1969) reported that most implantations occur within the central third of the uterine horn This held true for the animals in the present study they may implant In one case with a near term fetus the placenta covered the at the base of the horn blocking the exit of the fetus

Reproductive - 13 - March 1971

Birth iltleigh t

Barabash-Nikiforov (1947) listed the vJeight of a ne-v1born sea otter as 2000 g Kenyon (1969) presents the best quantitative data published to this time He found a maximum fetus weight of 1869 g 1-Thi1e he found pups as small as 1020 g those under 3 lb (14 kg) had died shortly after birth He estimated that successful birth weights range from 3 to 5 1b dr 1 4 to 2 3 and for purposes of calculation assumes an average birth weight of 1850 g to 1900 g

The weights of Weight Class 5 fetuses the smallest pups found living in the wild and three pups born in captivity (all died within 24 hours) are presented in Fig 11 These data tend to support Kenyons (1969) estimate of the ~veight at birth Few living pups middotHeighing less than 1350 g or about 3 lb and fevJ fetuses over 2000 g ( 4 5 lb) are found Kenyons upper estimate appears to be slightly high although there are extreme cases in both directions One pup weighed only 2 lb (900 and one fetus could not be weighed accurately on available equipment but weighed over 2500 g

One female with a 4 lb (1816 g) pup still had the placenta attached to the uterine wall and must have given birth shortly before being collected

From 11 it appears that most pups are born when they weigh in the neighborhood of 1800 to 1900 g in most of the populations sanpled Hmvever the and Delarof s indicate a bith Jei of between 1600 and 1700 g The Delarapound sample is srEall but the Tanaga sample the largest containing over a third of the large fetuses collected and was collected at a time of year when the birth rate is very high

This may be a reflection of the phys condition of the adult females and indirectly a reflection of food availab ty There are other factors indicating that the population is declining because of overpopulation

This lov1er birth yeight raises the question of middotwhether the pups are born earlier or whether are in poorer condition when born In an effort to gain some insight into this the fetal of Class 4 and 5 fetuses from Tanaga were plotted against their total lengths (Fig 12) middot Fetuses from other eastern Adak which appears to be a very healthy population were checked this curve fonaed by the Tanaga fetuses In all cases they fell Jell Jithin the limits of the Tanaga data indicating that there is no difference in the of fetuses of a given length It seems unlikely that a reduction in the rate of weight gain vould be completely proportional to a reduction of linear growth It appears that the rate of growth of fetuses is relatively constant but that females in poor physical condition may not be able to support as large a fetus as those in good condition so parturition occurs earlier

This may have sone effect on information on this

Fig 12 demonstrates some other interesting points There is no difference in the ratio and the maximum fetus size of males and females

~

(

~

-s 2 0 C

U)

0

--

ez ()

~) I) middot _t t- (

j -~middot -~

Reproduction - 14 - March 1971

This infeTs although dolt2s not prove an identical grmgtlth rate afld birth weight for males and females The circles on Fig 12 show the ~eight-length ratio of pups These fall below the curve formed by fetus weight-length ratios indicating that pups of a given length are some-Jhat heavier than fetuses of the sarne length This indicates an increase in weight immediately after parturition

Sex Ratio at Birth

Kenyon (1969) found that of 58 fetuses for which sex could be determined~ 45 percent vJere male and percent were female He assumed an equal sex ratio until a larger sample could be obtained

In the present study 111 fetuses were male and 144 ~vere female When this sample is combined with Kenyonts the sex ratio of 313 fetuses is 44 percent male and 56 percent female

Sex Ratio of the

In the process of harvesting capturing animals for transplants and delineating male and femalen areas it has become apparent that there are more females in the population than males Our general ion is that perhaps 60 to 65 percent of the sea otters are females Kenyon (1969) demonstrated that more juvenile males than females are found dead on the beaches of Amchitka lhe age structure of harvested in 1968 indicates that males may not live as long as fetnales

The lower birth rate higher juvenile mortality rate and slightly shorter life expectancy of males could account for the unbalanced sex ratio in the population

The sex ratio may vary from one island to another Presumably there -vmuld be more females in a population that has reached the carrying capacity of the habitat where juvenile mortality is higher There is some indication that males are the first to expand into new areas of unoccupied habitat Therefore in recently populated anas one might find more males However this situation vould be

Lit

The normal litter size of the sea otter is one This has been recorded by almost every observer since Steller Barabash-Nikiforov (194 7) reported twins in utero and mentioned other reports of twin fetuses from the early days of SnoF (1910) recorded a set of neHborn tdns In more recent studies~ both Sinha et al (1966) and Kenyon (1969) found reproductive tracts with two corpora lutee but in all cases only one fetus las present In thousands of hours of observation by many observers no tbullvin pups have been reported

In the present study 24 instances of multiple ovulations were recorded Five of these resulted in hv-in and one in triplet fetuses The information is summarized in Table 8

Table 8 Multiple Corpora Lutea Fetuses

Total Pregnant Females 565

Implanted Pregnancies 316

Unimplanted with 2 CL (corpora lutea) 9

Implanted Pregnancies with ) L CL and 1 Fetus 8

Implanted Pregnancies Vlith 3 CL and 1 Fetus 1

Implanted Pregnancies gtvith 2 CL and 2 Fetuses 5

Implanted Pregnancies middotwith 3 CL and gt Fetuses 1

Total Nultiple Ovulations

I

Percent Nultiple Ovulations 42

Percent of Pregnancies tvith Nultiple Fetuses 19

Corpora Lutea per Pregnancy 105

Fetuses per 102

Reproduction - 15 - lvlarch 1971

From these data it appears that in over 4 percent of the estrus cycles more than one ovulation takes place Of these about half result in the development of more than one fetus In most cases the were of a relatively large size and probably would have been born normally

All multiple fetuses appeared to be the result separate ovulations All had separate placentas and in some cases were different sexes Four tracts with twins had both fetuses in the same horn one had one in each horn and the tract with triplets had one fetus in one horn and t~vo

in the other Another tract had skeletal remains of triplets in one horn that were being resorbed Another tract appeared to have had five fetuses in one horn but they were almost completely resorbed One might infer that physically there is not enough room in one horn to accomn10date more than two fetuses for any length of time

With the exception of the newborn tvins reported by Snow (1910) there are no docu_mented records of female sea otters with tvin pups There are occasional unconfirmed reports of twin pups but as Kenyon (1969) points out a female may tolerate a pup in addition to her ovm but it is unlikely that a female could care for more than one pup It is unlikely that more than one pup survi~es In any case orily 2 percent of the pregnancies produce more than one pup 1middot1ultiple births cannot be a significant factor in the productivity of sea otter populations

The presence of a corpus luteum without gross evidence of implantation t-7as assumed to be an unimplanted pregnancy No attempt ~vas made to locate unimplanted blastocysts As a result e have little information concerning the survival of the blastocyst Of 206 reproductive tracts classified as nulliparous on the basis of the appearance of the uterus 12 had one corpus albicens and one had two Some of these may have been large attritic follicles Others may have ovulated and even conceived but implantation did not occur All of these cases represent the failure of the antmals first estrus

A total of 16 resorptions or possible abortions were identified (Table 1) The follmJing is a brief descdption of these ~7i th possible causes of some

Des

Resorption of blastocyst or ovum (corpus luteum 4 degenerating no evidence of ion)

Failure at time of (possibly because 1 of growmiddotth inside horn adjacent to implantation site)

Resorbed or aborted fetus (implantation at end of horn) 3

Resorbed fetus (umbilicus tvTisted tightly) 1

Resorbed fetuses (three or more fetuses in one horn) 2

Resorbed fetus (cause unknmm) 5

Reproduction - 16 - March 1971

Six of these resorptions may have been caused by a lack of room for growth of the placenta because of crowding from other placentas a growth or the end of the uterine horn These six and the one vith the twisted umbilicus are probably the result of random events or accidents

The numbers of resorptions are too small to accurately measure the frequency of such occurrences although the relatively large number in the Tanaga sample (Table 1) may be a reflection of the physical condition of the animals Part of this number is probably due to the fact that there are more pregnant animals in the sample In general the samples with the most pregnancies also conta~~ned the most resorptions Half of the Tanaga resorptions can be attributed to random events but five vere still due to unknown causes Eight of the 10 resorptions fo1md on Tanaga came from a 20-mile stretch of shore about 25 percent of the harvest area

While concrete conclusions cannot be drawn from these data they do raise the possibiliy that under certain conditions 5 percent of the pregnancies may fail

near Parturition

Collecting methods were such that the presence of a pup with a female was often overlooked Therefore the lack of a pup with a post-partum or lactating female did not mean that the pup had died However 14 females shmving of recent pregnancy were estrus presumably having lost their pup shortly before or shortly parturition

Again the magnitude of this mortality cannot be determined but it appears to be of the same order as the in utero mortality

after

Reproductive tracts were subjectively classified as post-partum or anestrous on the basis of degeneration of the corpus luteum (or appearance of the corpus albicans) the size of the horn of pregnancy and appearance of the placent scar In an effort to get an idea of how long it takes for the tract to return to normal the size of pups accompanying females in each category is listed in Table 9

It appears that females pass the subjective boundary from post-middotpartum to anestrus Hhen the pup about 10 lb and is around 75 to 80 em long

We do not know a great deal about the early growth of a sea otter pup but judging from the cur1e e9tablished based on cementum deposition of older animals and from tooth eruption a 10 lb pup is probably in its second month of life

Table 9 Pups Accompanying Pos artum Females

Sex llleight

M 2 lb F 3 F 4 M 5 F 55 F 625 F 775 M 10 F 11

Total Length

47 em 52 56 60 65 65 70 81 (borderline post partum-anestrous) 82 (late post-partum)

Pups Accompanying Anestrous Females

F 10 lb 78 em M 11 79 M 12 8l F 13 83 M 13 92 H 16 83 M 17 91 1 19 87 M 19 90 F 21 107 F 22 96 1 22 101 M 2Llmiddot 97 1 26 100 F 28 103

Reproduction - 17 - Narch 1971

Frequency of Breeding

Steller (1751) reported female sea otters with ne~vborn pups also accompanied by another pup that seemed to be no more than a year old Nurie (1940) recorded an instance of copulation by a female accompanied by a pup and Jones (1952) caught a young pup simultaneously gtvith a copulating pair Lensink (1962) felt that few females breed until the pup is a year old but mentions females with small pups also accompanied by large pups

I believe that some of thes2 observations may be misinterpreations of the situation Females will occasionally leave small pups for a time often near other animals The pups Lensinkmentions would be 2 years old and probably veigh 30-35 lb Sea otters are gregarious and subadults may remain near other animals appearing to be accompanying their mother I suggest that most of Stellers and Lens inks large pups ere such cases

Females gtvi th pups probably do breed occasionally but the most recent evidence indicates that this rarely occurs Kenyon (1969) records no instances of females accompanied by pups copulating and states that he found no females- knm7n to be accompanied by a pup that were This holds true for the animals collected in the present study However the nature of the harvesting operation is such that many females accompanied by pups were not recorded as such Therefore we are forced to look at lactating and assume that they were accompanied by a pup at the time of collection or shortly before le 10 sm1marizes the reproductive condition of the lactat females In the 1967 and 1968 samples some animals vith enlarged marmary middotmre listed as lactating As a result some females with near tenrr fetuses Here included Because this condition is considered to be associated with the unborn pup rather than a previous pup these animals were with the anestrous animals in le 10 In 1969 and 1970 the presence of milk was used as the sole criterion and no term animals we-rmiddote in the sample

Of 246 lactating females only one had an ted fetus (excluding the near term animals from 1967-68) Several others had small corpora lutea indicating that they recently become or possibly had large

vhich had luteinized but tvere not actually pregnant The remainder had follicles

This information that accompanied by a pup rnay enter proestrus and even ovulate but that they only rarely mate That they enter estrus is subs iated by the f2ct that the number of corpora albicantia in many tracts equals the age of the amplimal minus the three years prior to nBturity (see les 4-7) This indicates that large follicles tend to develop year after sexual nl8turi ty is reached whether the female is accompanied by a pup or notlt

Some of those lactating females with s1all corpora lutea may have recently weaned or othenvise lost a pup and are already pregnant again Malkovich (1937) took a pup least 3 months old) away in cap She mated 12 days later here she had the male It appears that a female enters estrus shortly after Heaning

Table 10 Reproductive Condition of Lactating Females

Location

Bay of Is Adak I Sept 1967 9 1 100

Allchitka I Sept-Oct 1967 17 2 105

Mid Pt - Blind Pt Oct 1968 19 3 136 Adak I

Bay of Is Adak I Oct 1968 22 4 154 3

Kanaga I Oct 1968 32 s- 135

Al1chitka I July-Aug 1969 4 2 333

iTanaga I May 1970 56 10 152

Delarof Is May 1970 18 3 143

Amchitka I May 1970 36 3 77

---~--middot--middot----~-~----~----middot~-middot--~----~----- -------~-~ middotmiddot--middot-shy

TOTAL 213 33 134

Anestrous or pregnant with term fetus

]_ Large follicles or corpus luteum present

3 Includes one implanted pregnancy with small fetus

Reproduction - 18 - March 1971

The question of hml long it takes for a female to enter estrus again after losing a small pup arises Eleven tracts of the 1970 sample and ttvo of the 1968 Kanaga Sampnple shmJed signs of being pregnant recently) but were in proestrus These animals had an enlarged horn usually middotlith a slightly pigmented area but no actual placental scar a11d a recently fonned corpus albicans However they also had enlarged follicles and the tvalls of the uterus were typical of proes trous animals Another animal was similar but appeared to have already ovulated and a corpus luteum was present In these cases the female has lost a pup either through abortion or vithin the first veeks after parturition and has started into another estrus cycle Three or four of these had follicles vhich may have started to become attritic This may be because the uterus had not recovered sufficiently from the last pregnancy

The implication of this information is that with a normal pregnancy and successful rearing of a pup the female becomes pregnac1t only after the pup has been weaned This n~y be a year after parturition However if the pregnancy fails or if a pup dies even at birth the female may become pregnant again in a few weeks rather than _waiting the full year

Kenyon (1969) suggests that the period middotbetween and the next estrus may long because he found animals that tvere not lactating but had a faint placental scar and an indistinct but recent corpus albicans These animals ltJere cLtssi as anestrus in the present study They are undoubtedly bet1veen veaning of their last pup and the next estrus The number of suh animals is t durirg Sspte12~ cnd October At this time the rate of estrus increases a1d the number of anestrous animals decreases sharply the next feJ months Therefore many of these anestrous anirrals become pregnant 1lithin a month or two Kenyon (1969) suggests that if the female keeps the pup for about a year that the period between births may be smreltmiddotJhat greater than two years He is assuming a 12-13 month gestation period However the present study shows that the estimate of a 12-13 month tation period was based on an inadequate sample Estimates in the present study put this period at closer to 8 or 9 months Presumably the anestrous nonlactating females are in the 3 to 4 month period

This does not change Kenyons (1969) point that the and the next estrus may long It indicates that in a normal cycle it may be months Hmvever as it vvas shmvn above~ it is possible for a female to enter estrus very shortly after losing a pup

We have demonstrated that sea otters breed at all times of year and it is possible for a female to become pregnampJt again after losing a pup sooner than she vould have if the pup had survived HmmiddotJever we have also demonstrated that annual of b and pupping occur and that these are similar in different years and in different populations The assumption can be made that the average length of the entire reproductive cycle is some multiple of a year

Kenyon (1969) assumes that tbe pup remains with its mother for about a year ~-Ieights and measurements cementum deposition skull changes~

Reproduction - 19 - March 1971

observations in the wild and in captivity all support this assumption although it cannot be said with certainty that it averages 12 months It could be s tly ITDre or less

Assuming about 12 months for rearing the pup 8 to 9 months gestation plus a fev months rest bet~veen and the next estrus the average reproductive cycle takes two years

This is supported by Fig 4 which shows that slightly over half the mature females are pregnant in a year Kenyon (1969) pointed out that his January and Harch samples gtvere biased against females with pups a1d therefore in favor of pregnant females because of selective hunting This bias applies to the fall sa~ples as well The summer samples were biased similarly because pregnant females appeared to be less resistant to handling during capture Therefore the percent of pregnant females is shown to be higher than lt actually Has -rhile no correction factor can be applied to all samples the evidence indicates that the percent of pregnaJt females is usually around 10-middot15 lmrer than in the sample For example the 1970 Alnchitka sltLrnple t-ras less biased and was about 9 percent lower than the Tanaga sample vrhich is l)nm-rn to be more biased Also on a June skiff survey 7 percent of the animals counted had pups that were readily identified If ~re assune that this is the nunber of animals avoided it can be calculated that 15 of the sexually mature females all of vhich are t middotTen~ avoided Actually this will vary with the hunter weather time of year etc Hmvever it indicates the order of magnitude of the bias

Using this as a rough correctton for the data in 4 we find that about half or slightly fe-Jer of the sexually mature females are pregnant each year or as indicated ly the average female has a pup every two years

Reproduction - 20 - March 7 1971

Conclusions

1 While sea otters may breed or pup at any time of year there are seasonal fluctuations in the illliTlbers of females in each reproductive stage The pattern of these fluctuations remains the same from year to year and population to population

2 Breeding activity is highest in September October and November

3 More implantations occur during January February and March than at other times of year

4 The birth rate is highest during April Y~y and June

5 Tlvo and a half to three times as many females breed during the peak breeding months as during the months of lowest breeding activity

6 Similarly two and a half to three times as many females pup during peak pupping months

7 During the period of lovest b activity seually mature females breed at a rate of 2 to 3 percent per month During the peak of breeding 7 to 10 percent per month breed

8 During the period of lov1est pupping activity 2 or 3 percent per month pup and during the peak of pupping 7 to 10 percent per month pup

9 The gestation period about 8 to 9 months on the average About half of this time is ~~ ted period

10 Female sea otters become sexually mature ihen 3 to 4 years old

11 Females may ovulate on an average of once a year after sexual maturity is reached

12 Females continue to breed at Cn old age

13 Sea otters retain certain characteristics common to the reproductive cycles of many other mustelids however exceptions are common Scheduling of the reproductive cycle during the year may have some advantages or it may be a trait that has not been entirely lost through evolution

14 Sixty percent of the near term fetuses are cephalicly oriented and 40 percent are candllly oriented

15 Sea otters probably birth both on land and in the water

16 Blastocysts rarely cross from one uterine horn to the other

17 Consecutive pregnanctes may occur in the same horn The side of pregnancy appears random

Reproduction - 21 - March 1971

18 Implantation usually occurs in the central third of the horn but it may occur any~rhere in the horn

19 Birth weights may range from 900 g to over 2500 g however most pups weigh 1800 g to 1900 g at birth

20 Females from areas gtvhere the population is declining because of food shortages may pup earlier and as a result the pups 1 birth weight may average 200 g lmver than nonnal

21 Male and female fetuses gr01middot1 at the same rate ampJd are the same size at birth

22 Body condition of the female has little effect on the growth rate of the fetus

23 There is a rapid ~Jeight gain shortly after parturition

24 The sex ratio at birth is 44 percent male and 56 percent female

25 The normal litter size is one hmmiddotever in 2 percent of the implanted pregnancies two or fetuses survive to near term

26 Multiple ovulations occur in 4 percent of the estrus cycles

27 es do not appeErc- to be able to more than two fetuses in a single horn

28 Up to 5 percent of the p may fail dne to in after implantation

29 An unknown but probably significant number of first pregnancies fail before implantation

30 The uterus is usually considered recovered from a pregnancy when the pup vleighs about 10 lb and is probably in its second month of life

31 Sexually matu1middote females normally have one pup every two years

32 Females vrith a pup may ovulate but rarely mate

33 Slightly less than half of the sexually mature females are in any given year

34 A female may enter estrus within a few weeks of losing a pup Consequently she may become agaln gtvi thout waiting the same length of time as if the pup hacl survived

35 Hhen a pup is weaned normally there is a longer period perhaps 3 or 4 months between weaning and the next estrus

Reproduction - 22 - March 1971

CITED

Barabash-Nikiforov I I 1947 The sea otter (Kalan) Soviet Ministrov RSFSR Translated from Russian by Dr A Birron and Z S Cole Israel Program for Scientific Translation 1962 227 pp

Hugget A St G and W F Widdas 1951 The relationship betlveen marmnalian foetal -reight and conception age Jour Physiology 144306-317

Kenyon K W 1969 The sea otter in the eastern Pacific Ocean USFWS North American Fauna No 68

Malkovich T A 1937 The sea otter in captivity Priroda No 381-87 Translated by J T Naximovitch 1966 Fisheries Research Board of Canada Nanaimo BC Translation Series 657

Sinha A A C H Conavay and K t-J Kenyon 1966 Reproduction in the female sea otter J Hildl Ngrrit 39(1)121-130

Snow H J 1910 In forbidden seas Edward Arnold London 303 pp

Wright P L 1963 Variations in reproductive cycles in North Arrierican mustelids In Enders A C ed Delayed tation Univ of Chicago Press 318 pp

  • SEA OTTER STUDIES - 1970
    • Surveys
    • Harvests and Transplants
      • Sexual Segregation in Sea Otter Populations
      • Reproduction in the Female
        • AERIAL COUNT OF SEA OTTERS- SOUTHSIDE OF THE ALASKA PENINSULA
          • Description of Conditions During March 1970 Aerial Count of Sea Otters
          • Discussion of Results
          • Sanak Island-Sandman Reefs
          • Shumagin Islands
          • Sutwick Area
          • Augustine Island-Cape Douglas
            • SUMMARY OF SEA OTTER SURVEYS
            • Amagat 1 to False Pass Ikatan Peninisula Cape Lazaref
            • Sanak Is
            • Deer 1
            • Pavlof Is
            • Northern Shumagin Is
            • Cold Bay 1
            • Kupreanof Peninsula
            • Mitrofania 1 to Kupreanof Pennisula
Page 6: Sea otter studies - 1970 - Alaska

AERIAL COUNT OF SEA OTTERS MARCH 1970

Partial or Location Count Date Camp 1ete Count Visibility

Cape Lazaref - Cold lsecty_

Cape Lazaref 3 320 Complete Poor

lkatan Peninsula 71 320 Complete Poor

False Pass - Amagat I 66 320 Camp 1ete Poor

Cold Bay 321 Partial Poor

TOTAL 141

Sanak Islands 239 322 Camp 1ete Fair to Poor

Sandman Reefs

Clubbing Rocks 12 322 Complete Fair to Poor

Cherni I amp Rocks 495+ 322 Complete fair to Poor

Goose I 7 322 Complete Fair to Poor

Hay I 18 322 Camp lete Fair to Poor

Hunt I 2 322 Complete Fair to Poor

Deer I 322 Partial Fair to Poor~

TOTAL 568+

Pavlof Islands

Inner 11iasik 2 321 Camp 1ete Fair to Poor

Outer lliasik 16 321 Camp 1ete Fair to Poor

Goloi 2 321 Complete Fair to Poor

Dolgoi 67 321 Complete Fair to Poor

Poperechnoi 29 321 Complete Fair to Poor

Ukolnoi 2 321 Complete Fair to Poor

yenWosnesenski 4 321 Complete Fair to Poor

TOTAL 122

Partial or Location Count Date Comp 1ete Count vis ib j 1i ty

Northern Shumagin Islands

Unga 184 321 Complete Fair to Poor

Popof 52 321 Complete Fair to Poor

Korovin 46 321 Complete Fair to Poor

Karpa __2t 321 Complete Fair to Poor

TOTAL 286

Cold ~ to Beaver ~ 0 320-21 Partial Poor

Beaver Bay to Kupreanof Pen

Beaver Bay 2 321 Partial Fair to Poor

Cape A 1 iaks in 4 321 Partial Fair to Poor

Guillemot I 16 321 Partial Fair to Poor

__1Kupreanof Peninsula 321 Partial Fair to Poor

TOTAL 23

Kupreanof Pen to Castle Cape 0 321 Partial Fair to Poor

AntJrrCastle Cape to-1-Ji e ~

Castle Cape-Castle Bay 16 321 Partial Fair to Poor

Chignik Bay 118 320 Complete Fair

Nakchamik I 5 320 Complete Fair

Hook Bay 13 320 Complete Fair

Cape Kum 1i un 88 320 Complete Fair

Kujul ik Bay 1199 320 Complete Very Good

Univikshak I 62 320 Complete Fair

Cape Kuml ik 54 320 Complete Fair to Poor

Sutwick I 14 320 Complete Fair to Poor

Cape Agutka 323 Complete Fair~

TOTAL 1766

Partial or Location Count Date Comp 1ete Count vis i b i 1i ty

Cape Kunmik 13 323 Complete Fair

Naka 1i kok Bay amp offshore islands 16 323 Complete Fair

Chiginagak Bay 5 323 Complete Fair

Agripina amp lmuya Bay 105 323 Complete Good

Wide Bay to Puale Bay N 0 T S U R V E Y E D

Puale Bay to c Kubugakl i __]_ 327 Partial Fair

TOTAL 146

Kashvik Bay to L Chiniak 0 327 Partial Very Good to Excellent

Cape Chiniak to Shaw _I

Shakun Is amp Rocks 71 327 Partial Very Good to Exce 11 ent

Kiukpal ik J to Shaw 1 0 327 Partial Very Good to Excellent

TOTAL 71

In the present survey which was made under relatively poor conditions 239 were seen in the Sanak area and 568+ in the Sandman Reefs The latter count Was incomp 1ete and conditions were such that the number of otters was more a function of time spent counting rather than area covered Obviously many were missed As a result not much can be said about total numbers The main change evident is that 141 were seen along the mainland and eastern portion of Unimak Island and 122 were seen in the Pavlof Islands This indicates that a fairly extensive movement northward and along the Peninsula is occuring

A remnant population probably remained along the south side of Sanak Island in the early 1900bulls By the late l940 1 s they began spreading into the Sandman Reefs This northward expansion has continued to the present time There is still unoccupied or sparsely populated habitat along the mainland shore and in the Pavlof Islands There should be continued expansion of this population for a number of years although the numbers around Sanak Island and the western Sandman Reefs may have already reached a peak

With the arrival of substantial numbers in the Pavlof Islands this population is probably on the verge of mixing with the Shumagin Island populshyation No doubt some individuals have moved back and forth in the past but the two populations are almost continuous at the present time

As in past surveys few otters were seen around the north side of Sanak Island and Caton Island This is probably poor habitat for otters The main population is around the south and west sides The higher count on the west end is probably more a result of intensive searching rather than a higher population

Shumagin Is 1 ands

This has been considered the largest of the four populations in the Alaska Peninsula area The most recent counts have not been adequate enough to show any major increase in numbers in the last decade however major changes in distribution have occured which are very similar to the pattern mentioned in the Sanak-Sandman population

There were occasional reports of sea otters in the Shumagins in the 1930s By 1947 Victor Scheffer estimated that 500 1 ived around Simeonof Island In 1953 Hooper counted 633 around Simeonof and Little Koniuj i Island The 1957 survey showed 1829 Most of these were- in the southern islands Five individuals were scattered in the northern islands and one was on the mainland shore near Elephant Point The 1962 survey totaled only 1352 The animals were scattered well off shore and the count was low However the count on Nagai increased from 149 to 338 indicating a continued northward expansion

In 1969 1510 were counted in the southern islands under relatively poor conditions An additional 286 were counted in the northern islands in the present survey and 23 were seen along the mainland north of the Shumagins Again survey conditions were not good

There is a very clear expansion of the population from a center near Simeonof Substantial numbers now occur on all the islands and repopulation of the mainland is occuring The population in the northern islands should continue to increase and most of the habitat on the mainland is not occupied

There is no evidence of an increase in total numbers since 1957middot However the last two surveys have been less than ideal and the large shallow off shore areas have not been covered adequately Considering the survey conditions and the obvious extentions of range it is almost certain that the populations from San-ak Island to the Shumagins have continued to increase In general it appears that the southern islands and reefs have become fully populated and the northern areas are just developing significant populations The entire a rea may be comp 1ete 1 y repopu 1 a ted in the next 10 years This wi 11 depend largely on the status and potential of the off-shore areas

Sutwi ck Area

Reports of sea otters near Sutwick Island are available since 1936 This population was far removed from other populations and is probably a remnant left after hunting ceased In 1951 Jones counted 388 between Cape Kuml iun and Cape Kunmik Most were near Sutwick Island In 1957 889 were counted Nineteen of these were between Cape Kunmik and Cape Providence indicating some northeastward expansion In 1962 949 were seen with individuals straying as far as Cape lgvak and one between there and Cape Kuliak In 1965 a stray otter was seen at Kinak Bay A major shift in the population from Sutwick into Kujulik Bay occured This may be the result of time of year and weather

In the present survey 1766 were counted between Castle Cape and Cape ~~~gtf~aip the majority were in Kujulik Bay Large numbers have moved

~~ranging as far as Cape Kubugakl i The area from Wide Bay to Puale Bay was not surveyed because of weather

The total count for the population was 1912 even though conditions were less than ideal throughout the bcunt and we feel many were missed It is possible that the increase in numbers is entirely due to reproduction assuming a 10 percent annual increase However it is difficult to compare surveys

There is still room for expansion in both directions from this population The greater movement to the northeast is probably the result of better habitat The population around Kujulik Bay is very dense and a large movement of animals may occur if competition for food becomes serious Otherwise we should expect to see continued steady expansion into adjacent areas for a number of years

to come

Augustine Island-Cape Douqlas

For some time a population has existed around the Augustine Island area at the mouth of Cook In 1 et In 1948 approxImate I y 50 sea otters vvere reported from Augustine Island Reports of sightings have been made from Shaw Island to Tuxedni Bay In 1957 Spencer counted 40 at Augustine and one at Shaw Island Lensink counted 52 on Augustine in 1959 but he considered it a poor count In 1965 Kenyon counted 18 on Augustine and 101 in the Shaw Island-Cape Douglas area In March of 1969 Loren Flag saw 62 around Augustine and in May 1969 Jim Faro counted 130 another observer saw about 30 some of which were probably not tallied by Faro

On the present survey Augustine Island could not be surveyed because of weather No otters were seen around Cape Douglas but 71 were seen in Shakun Islands and rocks near the abandoned village of Kaguyak These were probably from the Cape Douglas group

It appears that the animals move about in the area To date no complete survey of the area has been made at one time Until this is done 1ittle can be said about the population other than that several hundred probably occur there

The following table is a summary of sightings and counts made by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game These data were collected by different individuals using different types of aircraft under varying conditions of weather A strict comparison of numbers should not be made from this table

SUM

MAR

Y OF

SE

A OT

TER

SURV

EYS

(Com

pile

d by

p

op

ula

tio

n f

rom

L

ensi

nk

(196

2 T

hes

is

K

enyo

n 0

96

2 amp

196

5 R

epor

ts

amp M

anu

scri

pt)

an

d AD

FampG

Dat

a)

Lo

cati

on

P

re

1951

19

51

1957

19

59

1962

19

65

1969

19

70

AU

GU

STIN

E -

C

DOUG

LAS

Aug

usti

ne

Isla

nd

A

ppro

xim

atel

y 50

(1

948)

Shaw

1

amp

Dou

glas

Are

a S

igh

tin

gs

from

Sh

aw

Is

to

Tux

edn

i B

ay

TOTA

L

40

___

_

41

52

-shy 52

18

_lQ

J

119

130+

-shy

130+

J

71

ALAS

KA

PEN

INSU

LA

c

Kul

iak

-

c

lgva

k

c

lgva

k -

c

Kun

mik

Ani

akch

ak

Bay

amp

Am

ber

Bay

Sutw

i ck

Are

a M

isce

llan

eous

R

epor

ts

Kuj

ul i

k B

ay

amp c

K

uml

ik

Sin

ce

1936

C

Kum

1 i u

n

Uni

viks

hak

Isla

nd

N

akch

amik

Is

lan

d

8

355 12

13

0 19 6

581

103

180

1

22

47

109

684 86

Not

S

urve

yed

7(+

)

139

197 14

1 2

53

101 62

5

I

Lo

cati

on

P

re

1951

19

51

1957

19

59

1962

19

65

1969

19

70

ALAS

KA

PEN

INSU

LA

(Co

nt

)

Chi

gnik

B

ay

Cas

tle

Bay

amp

Cap

e

TOTA

L

SHUM

AGIN

Mai

nlan

d S

ho

re

Kup

rean

of

Pen

to

P

avlo

f B

ay

Ung

a Is

lan

d

Popo

f Is

lan

d

Kor

ovin

Is

lan

d

Kar

pa

Isla

nd

And

roni

ca

amp t

he

Hay

stac

ks

Nag

ai

Sp

ecta

cle

Ben

del

Tu

rner

Tw

ins

Few

R

epor

ts

Bef

ore

19

40

0

-shy

-shy

388

889 1 2 2 1

149

26

8

~-

7

- 949 4

338 I105

-~-middot

118

_lsect

1 9

12

23

184 52

46 4

75

232 8 27 6

Lo

cati

on

P

re

1951

19

51

1957

19

59

1962

19

65

1969

19

70

SHUM

AGIN

(C

on

t)

Nea

r

Pen

insu

la

Big

Kon

i u

j i

Lit

tle K

oniu

j i

amp A

tkin

s

Sim

eono

f

Bir

d

Che

rnab

ura

TOTA

L

500

Est

imat

e (1

947)

l63

3 (

195 3

)

3 0

220

430

455

160

___L

ll

183

0

14 3

222

255

294 38

_J

l

1 35

2

150 15

296

290

329 76

_6

15

1 0

-shy 309

SANA

K -

SAND

MAN

Mai

nlan

d S

ho

re

Pav

lof

Bay

to

Un

imak

B

ay

Pav

lof

Isla

nd

s

Wos

nese

nski

Is

lan

d

Uko

l noi

Is

lan

d

Pop

erec

hnoi

Is

lan

d

Dol

goi

Isla

nd

2

141 4 2 29

67

Lo

cati

on

P

re

1951

19

51

1957

19

59

1962

19

65

1969

19

70

SAN

AK

-SA

NDM

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Pav

lof

Isla

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s

Gol

oi

Isla

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Out

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sik

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Sand

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and

(Co

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ng

Roc

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se

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r Is

lan

d

amp O

ther

R

eefs

San

ak

Isla

nd

TOTA

L

2 16 2

27

(194

8)

271

259

495+

No

sig

hti

ng

s b

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re

1942

~

2 12

76

3397

82

7

123

5429

5 -

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75

~-

MARINE MAMMALS SURVEY - BRISrOL BAY TO AKUTAN ISLAND June 2 1970

OBSERVER J Vania J Faro

PILOT George Tibbets Jr

AIRCRAFr Piper Azetex

middotWEATdER Partly cloudy throughout flight wind 10 miles per hour or less visibility generally excellent One bad area and that was at Akutan Island Air was turbulant there and we were unable to go completely around the island because of fog in Akutan Pass Water had slight ripple throughout flight and ocassionally there was glare but overall conditions for sightinmiddot=r animals was good

Departed King Salmon 1115 AM Returned to King Salmon 745 PM

FLIGHT PA~1 middotAfter leaving King Salmon we went over to the north side of the Alaska Peninsula and proceeded down the coast flying at 300 to 600 feet of altitude Generally we flew aboumiddott one mile off the beach At Port Heiden Moller and Cinder River we checked the outer sand bars for seals and some of the inner bars where I knew seal hauled out

We did not cover Izembeck Laroon very well Instead we flew offshore and checked Amak Island We then fueled up at Cold Bay Leaving Cold Bay we continued dmvn the north side of the peninsula (sea otter sighted) down to Cape Sharichef and crossed over to Akutan Island flying along the south side of it We were able to get around Cape Morgan and fly doNn the beach a few miles but then had to turn back becausmiddote of fo9

The flight path was then eastward to the north side of Tigalda Island At Ugamak Island we circled it completely flying at about 300 to 500 feet We completed the survey at this point and returnt~d to King Salmon generally following the route we had taken on the flight down

SEA OTTER SIGHTINGS

Seven pods of sea otter were seen south of Amak Island These were photographed and later counted from middotthe photographs Beshycause of the density in the larger pods and the fact that some animals in the smaller pods were diving when the pictures were taken some individuals may have been missed

The number counted in each pod is as follows

1071 520 357 68 36 75 30

TOTAL 2157

Several hours later the pods had drifted four or five-miles northeastward (see map) bull In addition 1 one sea otter was seen at Amak Island and sixteen near Tigalda Island

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June 9 1970

On June 9 1970 an aerial survey of sea otter was flown in the Barren Islands and in the AfognakShuyak Islands area A Gruman Goose was used with Schneider serving as the only observer Offshore coverage was poor Conditions of visibility were as follows

Barren Islands- 1140 AM to 1225 PM- Water calm moderate surface glare Conditions verygood Count complete

Ban Island to Pernosa Bay- 1255 to 135PM- Conditions similar to Barren Islands but surface glare worse in spots Conditions generally good

Marmot Island to Latax Rocks - 255 to 345PM- Increased ripples on surface Glare Bad Conditions fair Count around Sea Otter Island complete although some otter may have been scattered offshore Remainder of count very superficial

Area

BARREN ISLANDS East Amatuli Island West Amatuli Island Sugarloaf Island Ushagat Island Rocks south of Ushacat Sud Island Nord Island

AFOGNAK-SHUYAK ISLANDS Ban Island-Shuyak Strait Pernosa Bay Marmot Island Sea Otter Island area East side of Shuyak Jest side of Shuyak Latax Rocks-Dark Island

TOTAL

TOTAL

Number of Sea Otters

0 2 0

76 200

29 0

307

18 6 3

121 0

33 26

207

Complete or Visibility Partial Count

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51

SEA OTTER COUNTS - KENAI June J970 to January 1971

Carl Divinyi flew aerial surveys of seals along the outside of the

Kenai Peninsula on a monthly basis He recorded all sea otters sighted

on these surveys The surveys did not cover the entire coast line and

the type of aircraft weather conditions and observers varied from one

survey to another Therefore the seperate counts cannot be compared

However when viewed in total they indicate the areas used by sea otters

and the relative abundance of otters in each area

The attached table presents the counts by broad areas

--

SEA

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70

AERIAL SURVEY Prince William Sound

April 1970

Between April 15 and 18 1970 Carl Divinyi and Julius Reynolds fletr an aerial survey of most of Prince William Sound using a Cessna 185 The survey tras primarily to locate seals however sea otter were counted The following is a summary of the survey

I departed Anchorage on April 14 via Alaska Airlines and arrived in Cordova at 600 AM tJeather conditions prevented flying so Julius and I drove the local road network looking for anything in the way of wildlife

April 15 vleather was a little better than yesterday so we decided to try surveying along the east side of the Sound up to Valdez Arm We started the survey at Bomb Point and flew the coast and offshore islands up to Galena Bay

Observations Seal - Scattered small groups with the majority of the animals being in Port Fidalgo There were no noticeable concentrations (50 on up) of seals

Sea Otter - A total of 105 otter were counted with the majority (60) being located in St Matthews Bay The remainder of the animals were concentrated between Red Head and Knmvles Head outside of Port Gravina

Sea Lion- Small scattered groups in Port Fidalgo (60-80 total)

April 16 Weather inclement - no flying

April 17 We flew Hawkins Hinchinbrook Montague and Green Islands Very few seal- many otter and two large concentrations of sea lions (See maps for numbers and locations of seals sea otter and sea lion) We also counted 5 otter around Kayak Island

April 18 Flew those islands from Montague Strait to Icy Bay and from Port Bainbridge to Knight Island Passage (See maps for numbers and locations of seals sea otter and sea lions)

Julius Reynolds continued the survey along the mainland from Knight Island Passage to Esther Island and around Knight Ingot and Eleanor Islands

The area from Esther Passage to Valdez Arm and Culross Perry Lone Naked Peak and Storey Islands were not surveyed

SEA OTTER COUNTED Prince William Sound

April 1970

Number of Area Sea Otter Date

PORT BAINBRIDGE - ICY BAY Bainbridge Island 30 418 Evans Island 14 418 Elrington Island 4 418 Latouche Island 46 418 Whale Bay - Icy Bay 39 418

CHENEGA ISLk~D - MAIN BAY Chenega Island and Dangerous Passage 33 52 Crafton Island 2 52

PORT NELLIE JUAN - ESTHER ISLAND Blackstone Bay 1 52 Culross Island NS

ESTHER PASSAGE - VALDEZ ARM NS

GALENA BAY FISH BAY 103 415

FISH BAY - GRAVINA POINT 1 415

GRAVINA POINT - CORDOVA 1 415

HAWKINS ISLfu~D 1 4l7

HINCHINBROOK ISLfu~ 99 417

EGG ISLAND 2 417

MONTAGUE ISLAND 259 417

GREEN AND LITTLE GREEN ISLANDS 102 4J7

KNIGHT ISLAND 136 52

INGOT ISLAND 6 52

ELEANOR ISLAND 3 52

SMITH ISLAND 4 52

SEAL ISLAND 0 52

APPLEGATE ROCK 1 52

PERRY ISLAND NS

(continued) SEA OTTER COUNTED Prince William Sound

April 1970

Number of Area Sea Otter Date-

LONE ISLAND NS

NAKED ISLAND NS

PEAK ISLAND NS

STOREY ISLAND NS

KAYAK - WINGHAM ISLAND 5 417

TOTAL 892

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PRE-TRANSPLANT SURVEY OF SEA OTTERS Green 1 andMontague 1 -July 171970

On July 17 1970 an aerial survey of sea otters was made in the area

around Green Island and the adjacent coast of Montague Island The purpose

of the survey was to locate concentrations of animals prior to a capturing

operation which began the next day A Dornier twin-engine arrcraft was used

with Schneider Reynolds and Somerville acting as observers The sky was

overcast almost no wind and the sea calm However heavy rain showers intershy

fered with forward visibility and the counting conditions were only fair on

the average Conditions vmiddotere especially poor in Port Chalmers and Zaikof Bay

The count began at 210pm and ended at 345 pm

The numbers and locations of otters are shown on the map In addition

16 sea otters were seen on a superficial look at Constantine Harbor on Hinchshy

in brook Island and a pod of 35 were seen three miles east of Johnstone Pt

Most of the area included in the survey was traveled repeatedly in a

skiff over the next four days Durlng this time the weather became calm and

clear for the first time in several days and the bulk of the sea otters shifted

from the coast of Montague out into the shallow areas between there and Green 1

and to Green and Channel Islands It was obvious that many otters had been missed

Fn the aerial sur~ey and that the population in the area is quite dense A

total of 48 sea otters were captured in three days of netting

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SEA OTTER SURVEY Salisbury Sound to Cape Spencer

August 23-25~ 1970

On August 23 1970 an aerial search for sea otters was made from Salisbury Sound to Torch Bay in the area north of Sitka A Helie Courier aircraft was used with Karl Schneider Alan Courtright and

middotWalt Cunningham serving as observers

The water on the outside coast was too rough for good visibility however it was calmer in the lee of rocks and islands and visibility was fair to good

On the initial survey only two otters were seen These were both in Surge Bay on Yakobi Island While returning after completing the count we located approximately 25 in the same area we saw the first two About 15 of these including at least one pup were in a single pod This illustrates how relatively large numbers of otters can be missed from the air The fact that none were seen in other areas does not mean that established populations do not exist elsewhere

On August 24 and 25 a search of the release area north of Khaz Bay was made by the skiff While flying into Kla) Bay for this seach two otters were seen near the old mine of Chichagof Two miners who have been working there reported that two otters have been there off and on since May

Rough weather and outboard problems restricted tl the search area~

and the effectiveness of the search in that area The following sightings were made

82470 I Adult West of Granite Islands 1 Adult South of Granite Islands

11 0A~dgtzl~4JVflup) middot1 In rocks SE of ranite Islands 82570 I Adult -n Southwest of Gran~te Islands

These sightings represent from four to seven animals

Reports over the last year indicate that sea otters regularly move in and out of Kla~ Bay and occasionally as far in as Sisters Lake bull

The fact that the count in the Khaz Bay area Has less than that made in 1969 does not mean much The animals appeared to be scattered during the present count the search did not include all of the nearby sea otter habitat and survey conditions were less than ideal

The population in Surge Bay appears to be separate and is probably well established Two otters were reported in the same location in January 1966 after only 23 otters had been released near Khaz Bay In 1969 two were seen in the same spot from the ait

While no estimate of the population of sea otters in the area can be made from the available information is evident that sea etters can be found along the entire west coast of Chichagof Island and that concentrations occur near Black Island and the Granite Islands north of Khaz Bay and in Surge Bay on Yakobi Island

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Harvest

In May of 1970 a sea otter harvest was conducted on Tanaga Island the middotDelarof Islands and Amchitka Island The numbers to come from each island

and the distribution of the kill around each island were planned in advance Conservative estimates of the population tvere made and the total harvest numbers were based on these estimates The harvest level for Tanaga and the Delarof Islands was set at 15 percent assuming that the next harvest in these areas would be three years later so the rate of harvest would be 5 percent per year Amchitkas harvest level was set at 10 percent assuming an annual harvest and a harvest rate of 10 percent per year

The harvest plan was based on information collected on a June 1968 skiff survey for Tanaga on the 1965 USFWS aerial survey and the 1969 ADFampG aerial

survey for the Delarofs and on 1968 helicopter survey by the USFWS for Amchitka There is good evidence that all the population estimates tvere low especially for the Delarofs

Hunting was done from avo skiffs with two men each Both men shot when possible A 117-foot vessel was used for skinning and living quarters Table 1 shows the schedule of the hunt with the daily harvest numbers

Table 2 presents the projected and actual harvest numbers by area Table 3 presents the numbers killed pelts saved specimen and pelt numbers used and the sex ratio of the kill for each area Figures 1 2 and 3 show the actual numbers and locations in more detail tveather was the main factor causing deviations from the original harvest plan Possible male areas were identified on the 1968 skiff survey on Tanaga An attempt was made to concentrate more pressure on these areas to achieve a more balanced sex ratio This effort is reflected in the sex ratio of the kill A high degree of sexual segregation occurs at this time of year and without specific pressure on male areas we could expect 85 percent females to be taken as occurred on Amchitka With the effort the female percentage was reduced to 65 percent on Tanaga In the Delarofs a male area which had not been identified previously was hit and the effect was much the same

Transplants

Two transplant operations took place in 1970 The first was done in the same manner as the 1968 and 1969 transplants A total of 86 otters tvere captured at Amchitka One aircraft load was shipped out Twenty-nine were released in Oregon and 30 in vashington The animals were held in floating pens at the release sites for several days to recover from the trip Survival appeared to be excellent

The second operation took place in Prince William Sound The Canadian research vessel G B Reed came to the Green Island-Port Chalmers area with tanks built on deck Forty-six sea otters were captured Approximately 44 were alive when the vessel left Prince William Sound however only 14 survived to be released off Vancouver Island

Table 1

April 20

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May 2

May 3

May 4

May 5

May 6

May 7

May 8

May 9

May 10

May 11

May 12

May 13

Schedule of the 1970 harvest

1030 pm depart Homer

Midnight arrive Adak MV Active

Refuel and prepare boat

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Complete skinning pack hides etc in pm

Delarofs

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900 am arrive Amchitka fly to Anchorage

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88

131

143

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Table 2 Projected and actual harvest numbers May 1970

Projected Actual Tana~a Island Harvest Harvest

Bumpy Point - Hot Springs Bay 200 191

Hot Springs Bay - Twin Bays 50 50

Twin Bays - Cape Sasmik 50 ) ) 199

Cape Sasmik - Tower 120 )

Tower Tanaga Bay 180 166

TOTAL 600 606

Delarof Islands

Gareloi 20-25 0

Kavalga Ogliuga and Skagul 75-100 125

Ulak and Amatignak 35-55 19

TOTAL 150 144

Amchitka Island saved for

USFWS Counting Units 1 amp 2 100 transplant

3 30 82

4 50 36

5 120 87

TOTAL 300 205

Table 3 Details of sea otters harvested in May 1970

Tanaga Island

Total kill 606

Number of pelts saved 598

Number of small pup pelts discarded 8

Specimen numbers SO 70-4 to SO 70-609 Pelt numbers 3198-3769 and 3771-3796

(Seal 3770 void - damaged)

Approximate sex ratio - 220 males 386 females or approximately 64 percent females

Delarof Islands

Total kill 144

Number of pelts saved 138

Number of pups discarded 6

Specimen numbers SO 70-610 to SO 70-753 Pelt numbers 3797 to 3934

Sex ratio- 44 males 100 females or approximately 69 percent females

Amchitka Island

Total kill 205

Number of pelts saved 194

Number of pups discarded 11

Specimen numbers SO 70-754 to SO 70-958 Pelt numbers 3935 to 4128

Sex ratio- 27 males 178 females or approximately 867 percent females

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A combination of bad weather a lack of time to let the otters recover from their capture and problems with the holding facilities probably caused the high rate of mortality

Summary of Harvests and Transplants

Table 4 summarizes the sea otters removed from Alaskan populations in the middot last 20 years An attempt has been made to remove 300 otters from Amchitka each year since 196 7 Originally this number was set as 10 percent of the population which was conservatively estimated to toal 3000 Recent USFtfS helicopter counts of up to 3927 show that this estimate definitely tvas low In the past the removal of animals was from the southeas tern half of the island The 1970 harvest was purposely concentrated toward the northwestern end in order to spread out the pressure and to learn something about the population in that area

The numbers of animals released in all transplant attempts by the U S Fish and Wildlife Service and the Alaska Department of Fish and Game are presented in Table 5

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30

SEXUAL SEGREGATION IN SEA OTTER POPULATIONS

by Karl Schneider March 1971

Workers have recognized for some time that sea otters tend to segregate by sex Lensink (1962) described this segregation around the southeastern end of Amchitka Island and identified three male areasn and three female areas He speculated that females used areas of more favorable habitat and that younger males were excluded by territorial males scattered throughout the female areas The implication is that male areas contain younger or at least nonterritorial males

Marakov (1965) mentions sexual segregation around Medny Island in the USSRs Commander Islands

Kenyon (1969) gives a more complete description of the sexual segregation around the southeastern end of Amchitka and supports it with quantitative data from harvested animals

Both Lens ink (1962) and Kenyon (1969) w_ere primarily describing hauling grounds used by different sexes While Kenyons harvested animals included otters near shore neither study provided much information on the use of off-shore areas

A knmmiddotlledge of the degree of sexual segregation and the location of male and female areas is important to the management of sea otter populations Harvests must be regulated to avoid putting too much pressure on one segment of the population Then capturing animals for transplants it may be necessary to set nets in specific areas to obtain the desired sex ratio case of a localized kill of otters such as when oil spills occur it is important to know the distribution of sexes to evaluate the importance of kill to the population

In

the

By the same token much can be learned about the distribution of sexes through harvest and capture operations The area from which each sea otter was taken during the harvests of 1967 1968 and 1970 was recorded Because a single hunting party might kill 30 otters over up to 10 miles of coast in a few hours it wasnt possible to record the precise locations Therefore the coast was broken up into areas and the numbers killed in each area ltJere totaled

Figs 1 - 5 show the locations of the areas letters correspond to those in Tables 1 - 4 which present the sex and age composition of animals harvested in each area The ages of the 1968 animals 1vere based on cementum deposition and reproductive condition The other samples were broken down using body size In general all males under 25 lb females under 20 lb and both sexes shorter than 100 em were considered dependent pups Females under 35 lb and 120 em and males under lb and 130 em were considered subadults These criteria probably underestimate the age of some animals Cementum deposition information for the 1967 and 1970 samples is not available yet

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Sexual Segregation - 2 - March 7 1971

Information from the 1967 Amchitka harvest and the 1968 1969 and 1970 Amchitka transplant capturing operations is not comparable and is not presented here However knmv-ledge gained from these operations has contributed to our understanding of sea otter distribution and this knowledge will be used in the following discussion In general this data confirms Kenyons (1969) observations for the Amchitka area although these summer and fall collections indicate a higher percentage of males in female areas than Kenyon reported from his winter and spring samples

Kenyon (1969) states that female areas are more numerous and less discrete than male areas He identified three male hauling grounds and 13 female hauling grounds on the southeastern end of Amchitka Island Recent studies involving netting and collecting animals off~hore indicate that male areas remain discrete off shore Usually these areas are off major points of land and pods of males often form in kelp beds directly off shore from the hauling ground

Female areas on the other hand are not discrete at all Any area that is not a male area can be considered a female area Some areas may be more attractive to females with pups or certain areas may be more favorable for hauling out but more females than males vill be found almost everyvthere except in the discrete male areas Therefore the main problem is to locate the male areas in et population He have no evidence of any shift in the location of male areas over a period of time so once an area is identified the information should remain useful for management purposes for many years Lensink (1962) and Kenyon (1969) correctly identified all of the male areas on southeastern Amchitka Extensive harves and netting have not turned up any new areas

Identification of Hale Areas by Pup Counts

In June 1968 a survey of most of Tanaga and Kanaga Islands was made by skiff to gather information to be used in planning harvests from those islands During this survey females with pups were counted separately from single animals No special effort was put into identifying pups fuen a female obviously had a pup it vas recorded Large pups often were separate from the females and some otters were seen only briefly in vaves kelp or at a distance Therefore many pups probably were missed and the counts should be considered minimal 6 and 7 present the counta by area A similar count was attempted by helicopter around Adak but for various reasons the results are not considered useable

From this skiff survey a number of areas were judged to be possible male areas Usually these were points or more exposed aTeas vhere relatively large numbers of single animals but no pups were seen

Shoal Point Kanaga Pass north of Eddy Rock and possibly Naga Point and Cape Tusik were suspected male areas on Kanaga Island Cape Sudak and Cape Amagalik were considered male areas on Tanaga Island and the area around Hazard Point and Lltnnoy Rock in Kanaga Pass was believed to blend with the Eddy Rock area

The best way to confirm the presence of male areas is to collect animals from these areas Harvests were conducted on Kanaga in October 1968 and on

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7A

Sexual Segregation - 3 - March 1 1971

Tanaga in May 1970 The sex ratios of the animals harvested are compared with the skiff survey pup counts in Table 5 The harvest areas are fairly broad while male areas are usually only a few hundred yards wide As a result a harvest area may include both a male area and portions of female areas to either side

The hunting pressure is seldom even over an area Often a good portion of the kill in an area would come from one group of rocks Therefore the information in Table 5 must be tempered with some knmrledge of how the hunting pressure was distributed

Kanaga Island

Shoal Point - While a few more females than males were taken in this area the percentage of males is considerably higher than one would expect in a female area Because weather vas less than ideal much of the hunting pressure ~vas directed to areas other than Shoal Point Most of the females probably came from these areas Therefore Shoal Point is considered to be a male area$

Naga Poin~- There is no strong evidence that this is a male area While no pups were recorded on the survey the total number of otters counted was not high Fairly extesive hunting of the area under good conditions yielded a sex ratio that would be typical of a female area Therefore it is assumed that there is no male area near Naga Point

Cape Tusik - Cape Tusik was suspected as a male area more because of the nature of the area than arry counts The count included areas to both sides of the Cape which could contain females Very little of the hunting was done at the Cape Therefore judgement on this area will have to wait until more information is available Extensive hunting indicates that no male areas exist east of Cape Tusik to Cape Chlanak

Kanaga Pass - From the count it appears that a male area exists in the Pass however the harvest area extended to Hive Rock and included a large portion of female area (Fig 6) The area is too broad as is shown in Table 5 gtvhere the number of pups counted for the whole area is relatively high The results of the harvest confirm the conclusions drawn from the count While the sex ratio for the entire area is roughly equal most of the males came from Kanaga Pass Some females without pups came from that general area but the majority of the females came from the area northeast of the Pass

The Tanaga harvest showed the male areas more distinctly for several reasons The total harvest was higher the entire count area was covered in the harvest and the hunting pressure was recorded more precisely Also as will be shown later sexual segregation is more pronounced in the spring

Cape Sudak - Most of the hunting pressure was concentrated on the tip of the Cape Some pressure was exerted on the area back to~mrd Pendant Point which is a female area as shmvn by the count The high percentage of males harvested from this area clearly confirms that the tip of the Cape is a male area

Tab

le

5

Com

pari

son

of

Pup

s C

ou

nte

d w

ith

S

ex R

atio

o

f H

arv

est

Jun

e

1968

A

rea

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rvey

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ber

19

68 H

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est

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ps

10

0

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ps

10

0

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ult

s T

ota

l KA

NAGA

IS

LAN

D

Ind

ep

Ott

er

Ind

ep

Ott

er

M

M

F

Rou

nd

Hea

d-S

ho

al P

oin

t 0

85

3

45

6

55

4

71

5

29

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a P

oin

t-K

anag

a B

ay

66

1

64

middot

18

5

81

5

20

5

79

5

Cap

e C

hla

nak

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e T

usi

k

51

1

93

2

40

7

60

3

53

6

47

Edd

y R

ock

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e R

ock

61

3

4

35

7

64

3

47

1

52

9

Jun

e

1968

A

rea

Su

rvey

H

ay

1970

Har

ves

t

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ps

10

0

Pu

ps

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0

Ad

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l TA

NA

GA

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LAN

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Ind

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Ott

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F

M

F

Cap

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ak-P

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6

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run

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92

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8

31

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t 6

9

82

1

32

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68

1

51

8

49

Sexual Segregation - 4 - March~ 1971

Annoy RockHazard Point - The harvest did not confirm this area as a male area However the weather Vas marginal and this area was too rough to hunt Therefore the otters taken in the harvest ~vere from either side of the Point and almost none were taken from the portion that is suspected to be a male area Actually Kanaga Pass is narrow and shallmv Otters feed in all parts of the Pass when weather and tide rips permit This could be considered a single male a~ea that serves both islands

Cape Amagalik - Two harvest areas overlap this area Lash Bay-Inferno Reef and Harem Rock-Kulak Point The kill from both areas strongly reflects the presence of a male area Nost of the hunting pressure was concentrated on Inferno and this is probably the main male area

As shown above vle 1vere able to identify four major male areas from a relatively superficial pup count No male areas were located during the harvests that had not already been identified from the count ~mile more detailed observations including field identification of males would be desirable the pup counting technique appears to be an acceptable method

Identification of

Because male areas are usually on v1ell exposed points it is possible to pick likely areas from a chart and then confirm these areas by collecting animals This method avoids the expense time and danger involved in surface surveys of remote areas Actual collection of animals also provides the most concrete proof of the nature of an area

The follmving is a description of the sex composition of areas for which no skiff count data was available

Delarof Islands - No attempt 1vas made to predict the location of male areas in the Delarof Islands This is a difficult area to work in Ogliuga Skagul and a portion of Ulak Island v1ere hunted on Nay 9 1970 The hunting effort in the afternoon was concentrated around Tag Island and the rocks south of Skagul Island While the entire kill is lumped together most of the males bullJere taken in the afternoon The percentage of males taken for the day is higher than would be expected if only female areas were hunted Most likely there is a male area near Tag Island

Adak Island - Portions of Adak gtvere hunted in 196 7 and 1968 No attempt was made to identify male areas before or during either harvest The harvest figures indicate that no male areas have been hunted The Bay of Islands is a classical female area The results of extensive hunting in the Bay and around Eddy Island indicate that there is definitely no male area there The percentage of males taken between Mid Point and Blind Point vms higher than might normally be expected in a female area however the scarcity of subadult males indicates that no male area exists within this area A large congregation of otters is often seen near Finger Shoals This could be a male area but Ye have no information from there If this is a male area a few stray males from there could account for the slightly higher number of males in the kill

Sexual Segregation - 5 - March 1971

Amchitka Island - While much work has been done on Amchitka Island almost all of the effort has been concentrated on the southeastern end from Crmvn Reefer Point to a fegtv miles northvest of Rifle Range Point During the 1970 harvest an attempt was made to distribute the harvest over previously unstudied areas Two areas were suspected to be male areas These were the rocks off Chitka Point and either Bird Rock or Aleut Point Heather prevented us from hunting the area around Bird Rock and Aleut Point

The sex ratio of the kill does not indicate a male area near Chitka Point however heavy waves prevented hunting off the point Therefore a male area could be there but because no animals were killed there the kill figures would not reflect it

Composition of Female Areas

Kenyon (1969) found that 93 percent of the adult otters and 63 percent of the juvenile otters in female areas were females The May samples from the 1970 harvest agree with this The mean percent of adult females in female areas was 93 percent The mean percent of subadults not including pups was 73 percent Hmmiddot1ever in the September and October samples only 77 percent of the a-dults in female areas were female About 86 percent of the subadults were female

The seasonal difference in the sex ratio of adults is consistant for all samples These seasonal changes are most likely due to adult males moving into female areas to look for estrous females During late winter and spring breeding activity is at the lmmiddot1est point of the year and fev1er females are entering estrus In fall the number of estrous animals is at its peak and presumably more sexually mature males move into the female areas to breed It is interesting to note that there were about three times as many males in female areas in fall as in winter and spring Information from female reproductive tracts indicates that approximately three times as many females would be in estrus at any given time in fall It appears that the number of adult males in a female area is directly proportional to the number of estrous females From the data collected there appear to be 15 to 20 males for each female -vlith enlarged follicles at any given time hmmiddotrever hunter bias might influence this figure

Ages have been estimated for otters harvested in 1968 on the basis of cementum layering These age data indicate that virtually all of the males in discrete female areas are either pups (or very young subadults) or are 7 years old or older No males between the ages of 2 and 6 Here found This strongly implies some form of territorial behavior among actively breeding males Fighting among males has not been observed ofteci hmvever a mature male at the Tacoma Zoo became intolerant of a subadult male in the presence of females They frequently 11 fought 1 although neither seemed to attempt to injure the other Finally the young male had to be removed Under imiddotJild conditions the young male might have moved out of the area without repeated physical encounters

Kenyon (1969) described the breeding behavior of sea otters His description indicated that males patrolled an area and two to four males may pass a given point during a 3 or 4 hour period Vandevere (1970) observed some males

Sexual Segregation - 6 - March~ 1971

apparently defending a territory in California but from his observations t

it appeared that successful breeding males were more mobile Perhaps established breeding males are more tolerant of each other More likely they maintain a separation from each other which limits the number in an area In this way a number of males might be patrolling the sa~e area rather than each establishing a geographical territory This might be advantageous where female concentrations shift from place to place as weather conditions change

Why does the number of adult males in female areas decline when the number of estrous females declines The limiting factor may be competition for estrous females rather than territory size With fewer receptive females competition lvould be greater causing some animals to leave the area Another possible reason may be that males may have a seasonal fluctuation in fertility Lensink (1962) found that all adult males produced sperm at all seasons Limited studies since then support this however it is possible that most of these males were collected in or near female areas Also while a male may produce sperm at all seasons there may be less production at certain times of year

A contributing factor may be that males are capable of feeding in more exposed areas than pregnant females or females with pups With a reduced attraction to the female areas either middotbecause of fewer receptive females or a reduced fertility in the male the male may find it easier to obtain food away from the crowded female areas where competition for food is greatest

The fact that the number of males does appear proportional to the number of estrous females indicates that competition for these females may play an important part in regulating the number of males in female areas

The numbers of subadults in the samples is relatively low so fluctuations in sex ratio may be due to sampling errors However the greater number of subadult males found in female areas in winter and spring may be a true increase made possible by the reduction in the number of breeding males A subadul t male would have fewer encounters -vli th breeding males

There are areas within what would normally be considered female areas where subadult males tend to congregate particularly during the winter These are usually areas v7ith a ma-r-ginal food supply that are not heavily used by females Because there are fewmature females in these areas there are also very few breeding males The subadult males using these areas seem to be transients that find less pressure from breeding males there These areas are used less in summer probably because calrrer weather allows feeding farther off-shore reducing competition for food in the male areas

Examples of areas within female areas that are used by subadult males are Constantine Harbor on Amchitka and possibly Finger Bay on Adak

within Female

In the course of the harvest on Tanags in tIay of 1970 a large number of pregnant females tvith large fetuses were collected at one time Most of

Sexual Segregation - 7 - March 1971

these came from Tidgituk Island This raised the possibility that females may segregate according to reproductive condition Table 6 presents the numbers of females in each stage of the reproductive cycle by area on Tanaga Island More pregnant animals were collected in the South Bay area and within that area most of the females with fetuses weighing over 100 g were collected near Tidgiktuk Island This concentration was evident in the pup counts made in June 1968 when a very high percentage of pups was observed in the same area (Table 5) Host females ~vith large fetuses in May would pup by June

The data do not reveal any other areas of this type on Tanaga Harakov (1965) mentioned a bay on Hedny Island that tvas preferred by pregnant females and females with pups Obviously all females do not go to a specific area to pup but a female vi th a large fetus or small pup probably has more difficulty in exposed areas and seeks more protected areas This tendency can be seen by the casual observer Single animals are more likely to venture off-shore and remain in rougher waters In some areas such as Crown Reefer Point on Amchitka there is a male area near the tip of a point Males congregate directly off-shore However to either side of these male congregations there are females that are not accompanied by a pup and probab do not have large fetuses In this way we may find segregation in a single kelp bed off a point If we set a net near the outer margin of the kelp we will catch almost all males Ho~Vever if we set a net to the side of the bed and closer tmiddoto shore we will catch mostly single females Nets set in a nearby bay will catch more females with pups

Table 6 shoHs that an unusually high number of resorptions were found between Cape Sudak and THin Bays This is probably a reflection of poor physical condition of the otters in that area due to food Other data indicate a continuous decline in body condition from the east side of Adak to Kanaga Pass There is some evidence that condition improves west of Kanaga Pass The animals in the Kanaga Pass vicinity are probably in poorer condition than any of the other populations sampled Therefore the high incidence of resorptions in this area should not be considered an example of segregation within the population

ition of Male Areas

As mentioned earlier the harvest areas ivere broader than any male area Harvest areas containing male areas also contained portions of female areas Our identification of the location at rhich each animal vas collected is not precise enough to determine the exact composition of these rnale areas Kenyons (1969) data from winter and early spring harvests are more precise His data indicate that at that time of year 98 percent of the adults and 80 percent of the ju~veniles using male areas were males Of 128 males in male areas 100 gtvere adults and 28 were juveniles

Experience from netting in the summer and harvesting in fall indicates that the percentage of males remains very high all year While adult females may be very close to male areas it appears unlikely that a significant number regularly use these areas at any time year nile adult males enter female areas regularly and in predictable numbers for a specific purpose females probably ~nter male areas only occasionally in stray movements

le 6

R

epro

du

ctiv

e S

tag

e o

f S

exu

ally

Mat

ure

S

ea O

tters

Har

ves

ted

on

Tan

aga

Isla

nd

-by

Are

a -

May

1

97

0

Un

imp

lan

ted

Im

pla

nte

d

Pre

gn

ant

Pre

gp

ant

Fet

us

Wei

gh

t C

lass

A

rea

No

np

reg

nan

t N

o

No

1

2 3

4 5

Res

orp

tio

n

To

tal

A

Su

dak

-Pen

dan

t P

oin

t 12

7

26

8 30

0

2 0

3 2

2 27

( B

arn

es

Po

int-

Ho

t S

pri

ng

s B

ay

12

7 24

10

36

3

0 1

4 2

2 29

B

( (

Bar

nes

P

oin

t-T

run

k P

oin

t 1

1

6 25

7

29

0 3

1 3

0 2

24

( T

run

k P

oin

t-T

win

Bay

s 7

9 39

7

30

2 1

1 1

2 2

23

c ( (

Haz

ard

P

oin

t-T

win

Bay

s 4

3 27

4

36

0 0

1 1

2 11

( T

win

B

ays-

Her

d

Roc

k 1

2

8 28

9

31

1 1

1 Lf

2

29

D

( ( T

win

B

ays-

So

uth

Bay

2

9 45

9

45

3 0

1 1

Lf

1 20

( S

ou

th B

ay-L

ash

B

ay

6 9

29

16

51

1 0

0 9

6 1

31

E

( ((

So

uth

Bay

-Har

em R

ock

17

14

31

14

31

2 0

1 5

6 45

(

F ( (

L

ash

B

ay-I

nfe

rno

Ree

f 4

1 9

6 55

2

1 1

0 2

11

( G

(

Har

em R

ock

-Ku

lak

Po

int

12

6 29

3

14

1 0

1 1

0 21

H

Ku

lak

Po

int-

SE

B

igh

t 13

9

28

10

31

2 0

1 2

5 32

Bra

cket

s in

dic

ate

o

ver

lap

pin

g are

as

Sexual Segregation - 8 - March 1971

Juveniles of both sexes probably move more randomly than adults lihile definite sexual segregation exists among younger animals it is not as pronounced as in adults Because subadult males are tolerated less by breeding males they are more strictly confined to small areas than females of the same age This has been demonstrated in fall harvests where extensive hunting over a large area will turn up only an occasional subadult male Then ~v-hen the hunters move to a definite male area large numbers of subadult males are taken in a very short time and in a very small area

While Kenyons data indicate that 22 percent the males in male areas are subadults our experience indicates that this percentage may be quite a bit higher at least in fall Such a seasonal change in the age composition of males sounds reasonable We have demonstrated that adult males move into female areas in greater numbers in the fall perhaps tripling their numbers in these areas At the same time fewer subadult males are found in the female areas Presumably the adult males are coming from male areas and the subadults driven from the female areas move to the same male areas The result vmuld be a higher percentage of young males in male areas in fall than in spring

The total number of males of all ages in female areas doubles in fall About 13 percent of all animals taken in these areas in spring were ~ales while 25 percent ta-ken in fall were males This suggests that the total number of males in male areas declines in fall There are more adults leaving than subadults entering the area

Sex Ratio

With the sexes segregating and with the degree segregation changing seasonally and betmiddotween age groups it is extremely difficult to get a completely random harvest As a result we have not been able to determine the sex ratio of any population as a vhole Hith female areas being larger and in more protected areas where hunting is easier there is a strong tendency to take more females than males particularly in sp-ring ~Je have shown that through a concentrated effort the percentage of males harvested can be increased however even then the lowest percentage of females taken in recent harvests ivas 62 percent on Kanaga Island in October 1968 Similar results occur in capturing operations for transplants

On all islands studied there are fewer male areas thltm female areas The male areas are very small usually only a few hundred yards wide The male feeding areas off male areas are also very narrow although they may extend farther off-shore In short that portion of the total available sea otter habitat included in male areas is very smalL

All evidence indicates that there are more females in the population than males While the percentage of is probably greater than that indicated by most harvest statistics it appears that at least 60 perc2nt of the sea otters in the populations studied are females

There are probably several factors contributing to this uneven sex ratio First our reproductive data indicate that 56 percent of the pups are females at birth Secondly Kenyon (1969) has found a higher mortality rate among

Sexual Segregation - 9 - March 1971

newly weaned males There is also some evidence that males might not live as long as females These factors could easily account for the preponderance of females in the population

It is possible that the higher rate of mortality in juvenile males is in part due to the breeding behavior of adult males and the resulting sexual segregation Juvenile males tend to be restricted to male areas or areas of marginal feeding habitat where there is little competition from breeding males During the ltvinter the feeding activity of young males would tend to be confined to small areas close to shore in male areas Therefore they may be subjected to greater competition for food than females of the same age which have a broader area in ltvhich to feed Kenyon (1969) found that subadult males vere less hardy in captivity than subadult females Therefore we can not adequately evaluate the influence of segregation on mortality

The sex ratio probably varies from one population to the next Very little juvenile mortality occurs in expanding populations eliminating that source of sexual selection However there is some evidence that males are more numerous among the first animals to repopulate a new area While a high percentage of males might be found on a newly repopulated island the loss of these males tvould alter the sex ratio of the parent population even though juvenile mortality might not yet be a significant factor

If the nThuber of breeding males is regulated by the number of estrous females an even sex ratio might produce a surplus of sexually mature males This surplus would not be beneficial to the population and could be detrimental This would permit the situation of an unbalanced sex ratio to evolve

The segregation of sexes and ages and the composition of sea otter populations is complex Unless we can understand the situation we cannot fully evaluate the effects of mortality or habitat changes

REPRODUCTION IN THE FElIALE SEA OTTER

by Karl Schneider Narch 1971

A total of 1358 sea otter reproductive tracts collected between 1967 and 1970 have been examined Ovaries were sliced with a razor blade and examined macroscopically Uteri Here macroscopically examined both internally and Each tract vas classified as to its stage in the reproductive cycle

The follovJing criteria were used in classification

Nulliparou~- Horns of uterus small thin and smooth no corpora lutea or corpora albicantia although there is occasionally evidence of a past ovulation but no evidence of implantation

Nultiparous - Uterus thicker corpora lutea andor corpora albicantia present Includes primiparous animals (these are not readily separated from multiparous animals)

Anestrus - Largest follicle under 3 5 rnm no corpus luteum

Proestrus - Follicles over 35 mm

Es - Follicles approximately 10 rrm

Implan~elt_ Pregnant_ - Visible s~velling in uterine horn usually with fetal membranes embryo or fetus visible Corpus luteum usually over 10 mm

- Uterine horn sti11 noticeab enlarged fresh roughshy~=-=--=~

al scar newly formed corpus albicans with some luteal tissue remaining

The appearance of the uterus ~ras used to confirm the classification The thickness arrd of the horns thickness and consistency of the uterine walls coloration of the of the horns and condition of the rugae change with each stage In most cases it is possible to guess the condition of the tract by a superficial examination of its exterior This appearance was used only to confirm what was indicated by structures in the ovary hmvever

Table 1 smnmarizes the reproductive condition of the sexually mature females in the that are enough for comparison throughout the year In the following discussion reported by Sinha et (1966) and Kenyon (1969) are included 7here possible The January and Harch are from these studies

Tab

le

l Re

p iv

e co

nd

itio

n o

f se

xual

ly m

atur

e se

a o

tters

_

IF

etus

Wei

--------------+

__

_

ln~a

ctive

ov

arie

s 1

Act

ive ovari~

ht C

Jas

s

jmiddot

I P

ost

-I P

roes

tru

s U

nim

pl

lmpl

D

ates

L

ocat

ioQ

1A

nest

rus

Part

um

Res

orpt

ion

l +

E

stru

s P

reo

Pr

eQ

2 3

lJ

S

~

4-8

1970

T

anag

a 1

51

41

0

10

104

17

8 10

33

30

4 (1

68)

(1

35)

(3

3)

(3 3

) (2

89)

(34~

(5

6)

(27

) (3

3)

(11

4)(

10

9)

May

9

19

70

Del

arof

Is

18

14

1

4 13

27

10

0

3 6

8 77

4

) (1

82)

( l

3)

(5

(16

9)

( o

) (1

30

) (3

9)

(78

)(1

04

)

10-1

219

70 A

mch

ltka

I

34

18

3 8

27

48

5 6

2 23

12

13

8

( 0

(24

6)

(13

0)

(22

) (5

8)

(19

6)

(34

8)

(36

) (4

3)

(14

) (1

67)

(8

7)

~lune

23

middotmiddot ~middot~~middot~Amchitka

1

17

2 1

2 4

18

1 2

3 5

7 44

A

ugus

t 13

196

8 (3

86)

(4

5)

(23

) (4

5)

(90

) (4

09)

(2

3)

(45

) (6

8)

(11

3)(

15

8)

July

6-

~dgtAmchitka

1

17

1 0

4 14

10

0

1 1

2 6

46

Aug

ust

819

69

(36

9)

( 2

0 2

) (8

6)

(30

4)

(21

9)

(22

) (2

2)

(43

)(1

32

)

Sep

t 1

0-17

A

dak

72

5 0

16

31

40

6 6

8 9

11

164

1967

(4

39

) ( 3

1)

(98

) ( 1

89)

(

4)

(3 7

) (3

7)

(49

) (5

5)

(6

7)

Sep

t

25

-A

mch

itk

a l

55

6

0 18

19

17

4

0 0

0 3

115

OcL

6

19

67

(4

7~8)

(5

2)

(15

7)

(16

5)

(14

8)

(35

) (8

7)

(26

)

Oct

o 1

2-1

5

Kan

aga

I

58

6 1

13

31

31

4 4

7 11

5

140

1968

(4

13)

(4

3)

(o 7

) (9

3)

(22

2)

(22

2)

(29

) (2

9)

(50

) (7

8)

(36

)

Oct

18

-20

A

dak

l

46

6 0

1 24

13

2

3 0

4 4

100

1968

(L

16 0

) ( 6

0)

(11

0)

(24

0)

(13

0)

(20

) (3

0)

0)

(40

)

Num

bers

in

re

nth

esis

are

pe

rcen

t se

xu

ally

mat

ure

fem

ales

F

etus

wei

ght

clas

s 1

=

0-l

g

2 =

1-l

Og

3

= 1

0-lO

Og

4

= 1

00-lO

OO

g

5 =

Tra

nsp

lan

t m

ort

alit

ies

(all

o

ther

sam

ples

fr

om

har

ves

ts)

Reproduction - 2 - March 1971

There may be some problems associated with comparing samples taken from different islands and in different years but allowing for sampling errors the data for the most part fit a definite pattern indicating that the annual cycle remains fairly constant The one outstanding exception is the number of pregnant animals in the t1m summer samples In 1968 there were many more implanted pregnancies than unimplanted pregnancies and in 1969 the situation was reversed hmvever the total number pregnant was almost identical These are the t s being relatively small collected over a longer of time and consisting of transplant mortalities An average of the two samples fits the pattern established by the other samples Relatively large samples are necessary because all stages of reproduction are found at all times of the year The period from November to early January is not represented hmmiddotJever things are changing so rapidly during that period that any information from that time may be confusing unless a large sample was collected in a very short time

The information in Table 1 is shmvn graphically vJith Kenyons (1969) winter information in Figs 1 and 2 It has often been demonstrated that sea otters breed and pup at all times of the year A number of observers have noticed that roore pups are born in the spring aJd sumrrter than at other times and there have been conflicting reports about breeding times Kenyon (1969) was the first to demonstrate seasonal changes in pregnancy rates but only his January and Yarch sallples gtvere large enough to be at all reliable

Figs 1 and 2 demonstrate that there are cons le secsonal in the numbers of animals in each s reproductive cycle There is a definite period of increased bree activity and a definite period of increased pupping Hmmiddotlever ~ some anii1als are in each s at all times of the year and these periods of increased breeding and pupping do not have distinct boundaries

Each curve is influenced by two factors so it is difficult to isolate the magnitude of any single factoc For example the of implanted pregnancies vill increase ss blastocysts and decrease as births occur The birth rate may be increas but if the number of implantations increases at a greater rate the implanted curve will still rise The animal remains in the anestrus implanted pregnant~ and unimplanted pregnant categories for a relatively long time and there may be considerable carry-over from one sample to the next Because the animal a relatively short time in the proestrus-estrus and post partum categories there is little if any carry-middotover from one sample to the next and the changes betgtmiddotTeen are more likely to be absolute changes The mean fetus weight class prob does not mean much because it does not take the actual number of animals in each class into account ~hen there are many implanted pregnancies ard the mean lmiddotTeigh t class is lovJ there actually may be more Class 5 fetuses in the population than vhen there are fetv irr1planted pregnancies and the mean middotleight class is high Therefore the actual percentages of all sexually mature females middotlith Class 1 ald Class 5 fetuses have been plotted in 3 These curves represent the actual number of fetuses in the population Because tha sample sizes become quite small vhen the fetuses are divi(12d into classes and the surruner sample is believ2d

2

~ _s ~

~

c U)

~-1

r

~J

lt

c])

Reproduction - 3 - Narch 1971

to be biased tvo curves for each class are shmvn The solid curves approximate the data while the broken curves represent subjective adjustments to correct for sampling errors

cussion of the Annual

The number of animals entering estrus rises in the fall At this time the pregnancy rate is at its lowest point and a high percentage of anestrous (nonpregnant) animals are in the population Breeding activity increases and at this time the number of sexually mature males in 1female areas increases

The number of unimplanted pregnancies begins to rise sharply in October and November At the smne time the nlli~ber of anestrous animals drops as these animals enter estrus breed and become pregnant Thj_s is the peak breeding season

The birth rate is at its louest this period as is shovm by the lmv number of post-partum a1imals and the lovr number of large fetuses

By late November and December the pregnant rate is increas even though the birth rate is slightly This is due to ne7

blastocysts implanting as is shown the increase in Class 1 fetuses

By January the period of t breedLng activity has declined and the number of unimplanted animals ceaches a peak as the number of blastocysts implanting equals the number of The lanted pregnancy rate then begins to declLne as fevJer animals breed and more implant The greater rate of antation is reflected in a rise in the Class 1 fetuses as well as an overall increas in the er of lanted pregnancies However the rate of implantation is partially obscured by an increased birth rate removing animals from the total of lanted pregnancies This increase in the birth rate is shmm in the rise in post-partum females and a rise in Class 5 fetuses

This condition of a low rate of breeding high rate of implantation and increasing rate of birth lasts from February to Hay

At this point He come to the least reliable data The sumner are smaller gtJere collected over a period of seven weeks and were from animals that died as the result of capture and handling

Therefore tvhile the data shaH that the post-partum rate decreases sh indicating a reduction in the birth rate~ the number of Class 5 fetuses

a peak indicating a very high birth rate The true birth rate is probably some1middothere in betigtJeen Females 1-ith very young pups (hence are post-partum) are vlary and probably less likely to be caught in nets Females with fetuses appear to be more likely to from handling In several cases~ twisting of the reproductive tract by a fetus vJas the actual cause of death Anestrous females (including post partum) on the other hand are less likely to die

Reproduction - 4 - March 1971

There is no question that the birth rate remains high during June and July Kenyons (1969) sum11er sample is limited but shows a large number of Class 5 fetuses Again the sample is biased in favor of pregnampIt animals but not to large fetuses only His field counts as vJell as observations by most other observers indicate an increase in the nUIlber of dependent young througbout the surnmer The number of Lmted pregnancies declines throughout the suTimer tvhile the unimplanted pregnancy rate declines at a slmver rate and breeding remains fairly constant at a lm-v level This indicates that through June there are more implantations than conceptions hmvever the birth rate is substantially greater than the implantation rate By late July and August there are relatively fe1r implantations taking place as shmm by a leveling of the unimplanted pregnant rate and a lotr number of Class 1 fetuses The number of Class 5 fetuses declines as a result of births during June July and August

Therefore the season of greatest pupping may be quite broad running from April through August possibly with the peak occurring in late May or June Until a better summer sample is available tve cannot pin dmm the peak of this pupping season with certainty (See Fetal GrmmiddotJth Rate for further discussion on breeding and pupp peaks)

If the information were precise we could compare subtracting numbers of pups born and numbers of conceptions to deterrnine the actual percent of ferrales breeding implantLng ltmd pupping in each month Unfortunately the data are not Each curve is influenced by more than one factor and ue do not knmmiddotr for sure the magnitude of any of these factors Hmmiddotrever by comparing some of the major samples e may be able to get a rough idea of how much higher the birth rate is at one time of year than another

The combined Mty and the combined September and October samples provide the best comparison Both are large samples One occurs near the peak of pupp and a lmv point in breeding and the other is near the low point in pupping and the peak of b

The number of Height Class 5 fetuses~ post-partum femaless and proestrousshyestrous females come closest to represent a particular event The time spent in each category is comparatively short

There were approximately three times as many Class 5 fetuses in Hay thanmiddot in Septerqber-October There Here also about three times as wany postshypartum females in May In tember-October there were two and a half to three times as many proestrous-estrous animals as in Hay It ~middotwuld appear that the peaks of breeding and pupping are roughly three times as high as the lowest points

At the lmmiddot12st period of pupping 2 to 3 percent of the females have Class 5 fetuses and should pup middotJithin a month If vm assume the post-partum stage lasts L 5 to 2 months (see Recovery of the Uterus) it also appears that 2 to 3 percent pup in the ltwrest months of pupping

Reproduction - 5 - March 1971

Similarly about 3 percent of the females have enlarged follicles during the lmvest period of breeding He do not knmv how long this period lasts hovever

At the peak of pupping about 10 of the females have Class 5 fetuses and should pup in the next month About 15 are post partum indicating about 8 births per 100 females in the preceding month

These percentages are prob2bly slightly high because of hunter bias avoiding females with pups It appeatmiddots that at least 2 to 3 percent of the mature females breed and 2 or 3 percent have pups in any month of the year In the peak breedingmonths such as September and October perhaps 7 to 10 percent breed and similarly in peak pupping months sud1 as Nay 7 to 10 percent of the mature females pup

Gestation Period

Barabash-Nikiforov (1947) listed the gestation period of 8 to 9 months This apparently was based on the case where a fentele mated in captivity as reported by i-Ialkovich (19 37) According to Barabash-Nikiforov a pup was born 8 months later although it died While it was fully formed~ it may have been premature

A number of births have occurred in captive animiddotmals held by the of Fish and Game in recent years In all cases the pu-ps ~-ere sc-middot1all usually less tha1 1600 g but still than the smallest pups found in the -rild All died after birth

The gestation period of 9 months has been repeated in the literature but apparently these statemsnts are based on Barabash~Nikiforol (19lf7)

Lensink (1962) also estimated a gestation period of 8-9 months based on field observations He felt that the peak of breeding was in August or September and the of pupping ~Alas in l-1arch or April

It has been demonstrated that the gestation of sea otters a relatively long period of tation (Sinha 1966 Kenyon 1969 and present study) Therefore any attempt to estimate the length of gestation by examinatioD of tracts must take both the unimp1anted and Janted periods into accour1t

Kenyon (1969) presented an estimate of the implanted period by assuming that the cube roots of fetus weights fall in a straight line after the embryo is established (Hugget and Hiddas 1951) and by that the European otter and Aluerican river otter would have similar fetal growth rates By plotting tenn fetus on a line established by the European otter he estimated an gestation of about 120 days not including the period for establishment of the ewbryo

Kenyon (1969) then an estimate made by Dr D G Chapman using the method of Hugget and ~-Jiddas (1951) v7here a regression line is fitted

Reproduction - 6 - March 1971

to the cube roots of the raeail of the mean weights of fetuses in each of five vreight clas3es plotted t the mean time of year

From the regression coefficient ob he estimated an implanted period of 154 days By taking the mean of the pregnant animals that are unimplanted pregnant estimated the tmimplanted period as about 75 months 1Ulmv-ing a half month for establishment the embryo he estimates a total gestation of 12-13 months

The data used by Chapman included a January-February sample of 26 a March-April sample of 49 and a Nay-August of only 9 The surtmler sample is more biased than the others and quite small yet the estimate relies heavily on this sa~ple when fetus size is the greatest

Table 2 presents a neTw- estimate of the length of the implanted period using the method used by Chapman but including data from the present study in addition to Chapmans data Table 3 presents a similar estimate of the length of the unimplanted period

The following is a comparison of the two estimates

Unimplanted Period 230 days 66 days Establishment of Embryo 15 15 days Implanted Period 154_~~Yrgt~ _73 d~s

Total Gestation Period 399 d2ys 154 days or about 13 months 5 months

Obviously there is some problem tiith the the technique or both A similar calculation mcde before the Hay sample had been collected produced an estimate of 109 for the implanted period and about 8 months for the total ation period

This method of calculation assur1es that there are definite peaks in breeding and pupping If breeding and pupping occurred evenly throughout the year the average fetus size 1vould remain constant thruughout the year and the technique would not tvork An ideal situation v10uld be when all individuals breed and pup 1vi thin very short periods Sea otters do have but are not well defined Very large s v-ould be required in such a case because there are ahvays fetuses of all sizes and the periods of maximum breeding are broad

Obviously Chapmans sample vas inadequate llthile the number of fetuses available for the latest estimate seems large and are well distributed throughout the year the a_ctual nu~ober in any cne fetus vJeight class at any one time of year is small A look at Table 1 shows that there is little consistency in many of the classes (Fig 3 shmvs that Class 1 and 5 fetuses do fit a pattern)

Table 2 Estimated Length of Period

Percent of Fetuses in each Height Class

Time of Collection After January

1 month 31 15 15 38 0

3 months 18 12 22 35 12

5 months 18 8 8 36 30

7 months 3 8 16 29 45

9 months 20 7 10 40 23

10 months

14 18 12 33 23

---~--

r1ean Time in Nonths After January 50 57 53 58 70

Cube Root of Umveighted Mean 063 17 36 7lf lllf

Specific Grmth Velocity 0169day EstirGated Implanted Period = 73 days

Table 3 Length of Unimplanted Period

Months After Unimplanted Unimplanted Percent January Pregnant Pregnant Unimplanted

1 month 35 26 58

3 months 49 49 so

5 months 128 179 42

7 months 22 37 37

9 months 50 57 47

10 months 55 44 56

Unweighted

48

Reproduction - 7 - March 1971

All of the estimates very heavily on su111mer samples 1vhen the fetuses are largest These sauples are knmm to be biased probably in favor of larger fetuses There are more fetuses at this time of year but it is exaggerated in these SCilllples

The latest estimate is not necess better than Chapmans even though it uses more and larger samples The velocity is twice as high as that calculated for fur seal and a 5 month gestation period is not consistent with other information available

This technique for estimating the lanted gestation period is not capable of providing a reliable estimate with the available data Therefore the 12 to 13 month estimate and any calculations based on it are meaningless

Much of the discrepancy lies in the estimate of the unimplanted period The present estimate that 48 percent of all pregnancies throughout the year are unimplanted based on an even distribution of relatively large samples and is probably more accurate than Chaprnan 1 s estimate of 60 percent

There are several other ways of guessing at the length of gestation Kenyons (1969) estillate of an lanted period of 4 months using the cube roots of term fetus weightscannot be relied upon entirely but it may be closer than the other estimates If gtmiddotJe estimate the uninplanted period from this using the 48 percent ted factor and allow for the establishment of the embryo we get a total gestation of about 85 months

Lensinks (1962) method of detennining the and peak pupping periods and taking the time between the peaks as the total gestation period has some merit TJnile Lens ink tried to determine these peaks by field observations we can determine them more accurately using the condition of reproductive tracts

Lensink felt that the peak of breeding middotlas in August or September The data in Fig 2 indicate that he may seen the beginning of the period of est breeding activity but that the probably in October or possibly November Lensink estimated period in larch or April Again the data indicate that he v1as s the beginning of the period of greatest pupping but the actual peak is probably in June or July At this time the number of Class 5 fetuses is greatest The nucJber of post-partum females probably and the number of implanted pregnant females is dropping from its If the gestation period of

11 11the average animal lasts from October or November to June or July we get a period of 8 or 9 months tThile Lensink 1 s was early 1 the duration agrees vith the present data

The peak of implantation when the greatest number of blastocysts are implanting can also be roughly estimated from 1 and 3 In February the number of unimplanted middot is dropping and the number of implanted pregnancies at this time th2 number of Class 1 fetuses probably reaches a

Reproduction - 8 - March 1971

Therefore it appears that the unimplanted period lasts 4 to 5 months and the implanted period from 4 to 5 months 1vi th a total gestation period of 8 to 9 months It should be recognized that these estimates are based on general trends and not distinct Therefore they are approximate However the relative length of the unimp and implanted p2riods agrees with the percentages noted previous and this estimate agrees vlith the one based on Kenyons (1969) comparison vdth other species of otters

If the gestation period ~Jere 12 months the percentage of pregnant animals would remain constant throughout the year If it vJere only slightly longer or slightly shorter there would be only a slight fluctuation unless there were a distinct breeding period Fig 4 indicates a substantial fluctuation indicating a gestation period substantially shorter or longer than 12 months

The peak of pregnancy occurs in February after the period of t breeding activi The luH point on the pregnancy curve occurs after those animals giving birth dur the period of t pupping have pupped That is the t point on the pregnancy curve occurs after the main breeding and the lmves ~ point is after the main pupping period These periods actually last for 4 or 5 months so these points are well after the peaks of breedtng and pupping

The time betbulleen the peak when the pregnant and the lmr point Hhen the fewest are the average gestation period This is about 85 months (Fig 4) The anestrus curve (Fig 1) shmvs the same thing betng a negative of the pregnancy curve

Hith three different methods we have estimated a gestation period of 8 to 9 months Hith the unimplanted period roughly equal to the implanted period This agreas bullmiddotlith the observed period in captivity (BarabashshyNikiforov 194 7) and with field estimates Any gestation period differirg greatly from this does not fit the data Conceivably one could apply a 20 to 21 month period to the SaTEpound information however if this 1vere the case females would have to breed vJhile accomp by a pup or be on a four year reproductive cycle This prospect stretches the imagination

There is a definite possibility that the gestatton period varies in sea otters as in land ot te~cs This most likely -middotwuld occur through variations in the unimplantec1 period The above discussion refers to the average gestation period Until better information to the contrary is obtained we must continue to assume that the average period is 8 to 9 months

Fetal Rate

the estimates of the gestation curve are corrects Kenyons (1969) Fig 4 should approximate the grouth curve of the fetus Fig 5 duplicates this and shmvs the actual Heights Fig 6 shous the same information more graphically It is possible to esttmgtte the of time in each v7eight classlt Fig 6 does not take the period for establishment of the embryo into account so Height Class 1 may be longer than shmm perhaps 15 days if we accept Chapcnans (Kenyon 1969) es e

--

3 0

~

shy

----- -shy --middotmiddot~middotmiddot

___ ___1__

_

(middot-)

_ ft -) J ~)___

~-

3

Reproduction - 9 - IYarch 1971

The short time span of Classes 2 and 3 explains the relatively small numbers and the lack of a consistent pattern found in those classes (Table 1)

Using this curve it should be possible to predict the time of birth and with less precision the time of Luplantation and conception of any particular fetus By plot the estimated birth dates of a large number of fetuses rve should be able to drav curves Ttlhich approximate the breeding implantation and pupping rates of the population throughout the year

Unfortunately there is overlap bet1veen samples collected at different times of year Some of the samples are too small to provide a comparison We vould have to give eight to each in order to combine them Therefore only the ttay September and October samples are used lfuere the September and October samples overlap t1lt10 separate points are used Fig 7 presents the estimated birth dates grouped by month Fig 8 separates the dates into half month groups February and Narch are not represented and April is an estimate based on post-partum females rather than fetus size (see Recovery of the Uterus after Parturition)

The peak in this curve may be exaggerated It has been shmvn previously that fetus size is less consistent bet-Jeen samples than the other categories (Table 1) because the sample size of each grouping is small Therefore the exact shape of the curve may not be correct If however we assume the approximate of the peak is correct we can construct a model of the based on our estimates of the gestation period besed on a fetal grmJth rate estimated from this estimated gestation Therefore our reasoning would be some~hat circular if -Je used thjs to prove the estimate Hmvever we can see how this model fits the rest of the data A good fit would tend to support the model and the estimates on which it is based

For this model He use the curve in 8 to represent births We assume a gestation period of 8 months -rith the unimplanted and implanted periods each lasting 4 months TheTefore v8 draw curves identical to the birth curve but moved backward 4 months for implantation of the blastocyst and 8 months for conception (Fig 9)

A comparison of the peaks in Fig 9 with the data in Figs 1 2 and 3 and the narrative b on those indicates a fairly close fit The data indicat that the peaks may ~ctually occur slightly later than indicated in 9 but in the timing and distances bet7een the peaks fit

The sharpness of the birth curve indicates that the breeding in1plantatimiddotm and pupping peaks mey be more distinct than indicated by the curves in Figs 1 and 2 It is possible to fit curves with more abrupt changes to the data Fig 10 shows the same data lith the curves drawn to more closely fit the model tfuether these curves are more accurate or not is open to debate The data on Hhich the birth curve is based is not that strong The numbers in each month are relatively snall If such distinct did occur it seems likely that field observers ~vould have seen the effects Nost field observations indicate broader peaks that are not gtvell defined

2

G

X

middot ~ -- (-- ) r~ ~ - - -- ~

yen~ r- ~--~

3

i__middot -~---

~

gt

- cshy

Reproduction - 10 - March 1971

~e of _Sexual Maturity

Tables 4 - 7 list the frequency of nmnbers of corpora albicantia and corpora lutea found in females of different areas It should be recognized that the ages may not al-1ays be exact Animals ATTichi tka to become sexually mature Hhen about 4 years old It appbull~ars that more animals may become mature earlier on Adak The information is too limited to dratmiddotJ any definite conclusions from this hmmiddotJever the food availabili ty is probably better at Adak and an earlier age of sexual 1naturity may be a response to better nutrition

Ovulation

From Tables 4 - 7 it can be seen that the maximum number of corpora albicantia for each age roughly the numb~r of years after sexual maturity that some animals ovulated once every year after maturity It can be assumed that many corpora albicantia are invisible macroscopically after several years particularly if they are not remnants of a corpus luteum of pregnancT Therefore it Ls possible that most females ovulate every year This poss ili ty is also supported by the high incidence of large follicles in lactating females (TabL~ 10)

faximum

Tables 4 ~ 7 indicate that at t some females continue to breed at a very old age The samples are teo small to determine 1rhether a t number become unproductive at any point

Ovulation

Many mustelids are induced ovulatoTs

On September 14 1967 a female vJas tsd Hhile copulating She appeared to have Oilnlated shortly before Because sea otters may copulate several times over a period of two or three (Kenyon 1969) ovulation may have been induced by a previous

ficance of

Delayed lon is a charact8ristic of the reproductive cycle of most mustelids Several theories on the ccdv of a delay have been advanced Most assume that spring is the most favorable time of year for survival of nemiddotJborn young but that either a wir1ter breeding season is not favorable or that the presence of a Ttlale vhich occurs only during the

season is for survival of the young of the previous pregnancy (1963) discusse the evolution of delayed implantation in mustelids and points out that there are exceptions even in arctic areas middotJhere time of birth be consLdered more may be born at any tin1e Scheduling the and made possible by delayed mey be advantageous but is not necessary

----

a - o lt bull y bull bull l w laquo bull _ ~ ~

Table 4 Frequency of occurance of Numbers of Corpora Albicantia Amchitkll - June-August 1968

N U M B E R 0 F C 0 R P 0 R A A L 8 I C A N T A

I

~I -AGE -~ 1 I 1084 6 92 5 7(Years) 1 middot shy

- 0-l - ~~ ~- middot~middot- middotmiddotmiddotmiddotmiddot

1-2

2

3

4 1 I

2 (1)

6

middot5

1 (1)1

2 (l) 17

1 ( 1 )

2 ( 1)

18

l (1) I ( 1 ) 9

10 1 ( 1) L ~j( 2 1 ( 1 )

2 ( l) 1

12

11

1 (1)

I I (I)

1

2 ( 1)

13

1 ( 1 )

2

114

115 I116

17

I 18

I I

I

119 I

Numbers inside parentheses =Number of individuals with corpus luteum

11

Table s Frequency of oc~urance of Numbers of Corpora A1bicantfa _Kanaga - October 1968

N U M B E R 0 F C 0 R P 0 R A Al B CANT IA - I I I

AGE 84 I I

5 I l 6 7z 3(Years) 1

l rbull ~middot--

--- shy

0-l

J l-2

~ 2

3

4 2 ( 1)

5 3

Ibullbull 6 6 (3)

4 (4)7

8 4

3 9 2 ( 1 )

j iI 10

I 1 (1)11

12

13

14

1 ( 1)

16

15

17

18

Numbers irisi

-

J

(Plus 2 with corpus 1 uteum but no corpora a 1 b i cant i a)4(1)

1

1 (1)

5 (1)

10(5)

3

3 1

3 (2)

2 ( 1) 3 (3)

1 ( 1)

2

1 ( 1 ) 5 ( 1)

1 ( 1 ) 3 ( 1)

1

3

2 ( 1 )

1 (l)

1 (1)4 ( 1)

2) 1 (1) 3 (3) 4 (1) 12 ( 1 ( 1 )

1 ( 1)

1

2

1 ( 1)

1 ( 1)

1

I I parentheses = Number of individuals with corpus luteum

11

Table 6 Frequency of occurance of Numbers of Corpora Albicantia Mid Pt-~Blind Pt~ Adak- October 1968

N U ~1 B E R 0 F c 0 R P 0 R A A L B I C A NT I A

-1AGE

~

9 1084 75 6Years) 1 2 3 - shy ----- --middot shy~--

- _ Q~L

1-2

(Plus one with_corpus luteum but no corpus a1bicans)2 1 l I I I I I ) (One with corpus luteum but no corpus albicans 3

4 2 (1)

l5

16 1 (1)

1 1)7

8

2 (1)9

10

111

12

I13

14

15 1 (1) 16

17

18

1 ( 1)

1 (l)

1

1

1

2

1

1

1

1(1 )

1

1 (1)

1 (1)

1

I

1

1

1

1

-I

Numbers Inside rentheses = Number of individuals with corpus luteum

1

Table 7- Frequency of occurance of Numbers of Corpora A1bicantia Three Arm Bay - Bay of Islands Adak - October 1968

AGE (Years)

Q-1

t-2

2

3

4

s 6

7

8

9

10

11

12

13

14

15

16

17

18

Numbers

N U ~ B E R 0 F C 0 R P 0 R A A l B C A N T I A

~L 3 4 slE - 1middotshy

7 1~8 __J~l_11 shy

1-

One -Ji th corpus 1uteum but no corpora _a 1bicantia I I I I I I I

One vJi th corpus luteum but no corpora al bicantia

12 I I (Plus one with corpumiddots luteum but no corpora albicantia)1 (1) 3

1 (1) 2 (1)

1 2 (2) l 1

1 (1) 4 (1)

2 (1)

2 1 )

1 ( 1 ) 1

2 (1)

2 (2)

l22 3

3 ( 1)2 21 ( 1)

2 ( 1)

2 ( 1)

l ( 1)1

1 1

1 I ll

I

I I

inside parentheses Nurnber of individuals with corpus luteurn

Reproduction - 11 - March 1971

The sea otter lives in an area of relatively uniform temperatures However storms tend to be more frequent and violent in fall and winter Survival may be better in the late spring and sunrmer Therefore there may be some advantage to birth in the late spring However there does not appear to be a great deal of advantage to a breeding season at any particular time of year

Wright (1963) pointed out that most musteHds lant after daylight begins to increase usually around February This is the period when sea otters implant at the t rate If the time of implantation is influenced by daylight the period when the greatest number of births occur could be shorter than the equivalent breeding period The data do not show this although the possibility cannot be ruled out

The fact that substantial numbers of sea otters are breeding implanting and pupping at all times of the year indicates that hile there may be some advantage to certain events taking place at a particular time of year that advantage is not great enough for a distinct breeding season to evolve

Sea otters exhibit certain characteristics that are comnon to most mustelid breeding cycles but these chfracteris tics are not as well developed as in most es

It is interesting to note that Hright (1963) correlated the molt of Mus with the implanted period TI1e molt began around the

ion and ended around parturi tioD

Sea otters molt all year and some are implanted pregnant at all times At this time e cannot say uhether any between molt and pregnancy exists

Fetal

Kenyon (1969) found that the number of caudally presented fetuses roughly equalled the nunber of cephali presented fetuses He suggests that this indicates a lack of tatim1 for birth occurring in water and infers that sea otters must give birth on land

Fetuses of all 1veights in the present study also appear to be randomly oriented however 97 Class 5 fetuses (1000 g and over) showed 60 percent cephalic ion while only 40 percent shovJed caudal presentation

The orientation of large fetuses should be a better indicator of orientation at birth Smaller fetuses may position Therefore it appears that more sea otters are born head first than tail first

If you accept Kenyons premise that caudal presentation is necessary or at least beneficial to birth in water this more recent information supports the supposition that birth occurs on land The little information available indicates that birth does occur on lando but there have been unconfirmed reports of birth in water

Reproduction - 12 - tltIarch 1971

At least some sirenians are born head first indicating that caudal presentation is not necessary North of Unimak Island a large population of sea otters lives off-shore There is little evidence that any numbers come ashore It seems likely that many of these animals are born in the water

Of 305 implant pregnancies 138 fetuses (45 percent) had implanted in the left horn and 167 (55 percent) in the right horn Most of the difference was in the 1970 Amchitka sample ~fithout this sarnple 48 percent implanted in the left horn and 52 percent in the right horn Kenyon (1969) found an equal number in each horn If a real difference exists it is probably exaggerated by the 1970 A~1chitka sample

Out of 305 implanted pregnencies the point of implantation was on the same side of the reproductive tract as the corpus luteum in all but three instances In one case the corpus lutaum was in the left ovary and the fetus in the right porn In the second case the corpus luteum in the right ovary but the fetus was in the left horn In the third case there was a corpus luteurn in each ovary and tHo fetuses (of different sexes) in the left horn

It appears that blastocysts rarely cross from one horn to the other

Because there is usually a long period of time parturition and the next est_rus is little if any inhibitory effect on ovulation caused by the corpus of the previous pregnancy The ovary producing the largest follicle appears to be random and there does not appear to be the tendency for a pregnancy to be from the opposite ovary from the previous pregnancy In many cases there are many more corpora albicantia in one ovary than in the other

When a loses a pup shortly after birth and enters estrus before the uterus has conpletely recovered and before the corpus luteum has completely degenerated there may be some inhibitory effect Of the ll+ such cases identified six had large follicles in the same ovary as the new corpus albicans and in the opposite ovary It appears that any inhibitory effect by the corpus luteum only lasts for a short time

A few cases bullv-ere found vJhere one ovary was not developed or othenvise not capable of producing ova Because one ovary may support repeated pregnancies such animals may be as productive as those ~vith both ovaries active

Kenyon (1969) reported that most implantations occur within the central third of the uterine horn This held true for the animals in the present study they may implant In one case with a near term fetus the placenta covered the at the base of the horn blocking the exit of the fetus

Reproductive - 13 - March 1971

Birth iltleigh t

Barabash-Nikiforov (1947) listed the vJeight of a ne-v1born sea otter as 2000 g Kenyon (1969) presents the best quantitative data published to this time He found a maximum fetus weight of 1869 g 1-Thi1e he found pups as small as 1020 g those under 3 lb (14 kg) had died shortly after birth He estimated that successful birth weights range from 3 to 5 1b dr 1 4 to 2 3 and for purposes of calculation assumes an average birth weight of 1850 g to 1900 g

The weights of Weight Class 5 fetuses the smallest pups found living in the wild and three pups born in captivity (all died within 24 hours) are presented in Fig 11 These data tend to support Kenyons (1969) estimate of the ~veight at birth Few living pups middotHeighing less than 1350 g or about 3 lb and fevJ fetuses over 2000 g ( 4 5 lb) are found Kenyons upper estimate appears to be slightly high although there are extreme cases in both directions One pup weighed only 2 lb (900 and one fetus could not be weighed accurately on available equipment but weighed over 2500 g

One female with a 4 lb (1816 g) pup still had the placenta attached to the uterine wall and must have given birth shortly before being collected

From 11 it appears that most pups are born when they weigh in the neighborhood of 1800 to 1900 g in most of the populations sanpled Hmvever the and Delarof s indicate a bith Jei of between 1600 and 1700 g The Delarapound sample is srEall but the Tanaga sample the largest containing over a third of the large fetuses collected and was collected at a time of year when the birth rate is very high

This may be a reflection of the phys condition of the adult females and indirectly a reflection of food availab ty There are other factors indicating that the population is declining because of overpopulation

This lov1er birth yeight raises the question of middotwhether the pups are born earlier or whether are in poorer condition when born In an effort to gain some insight into this the fetal of Class 4 and 5 fetuses from Tanaga were plotted against their total lengths (Fig 12) middot Fetuses from other eastern Adak which appears to be a very healthy population were checked this curve fonaed by the Tanaga fetuses In all cases they fell Jell Jithin the limits of the Tanaga data indicating that there is no difference in the of fetuses of a given length It seems unlikely that a reduction in the rate of weight gain vould be completely proportional to a reduction of linear growth It appears that the rate of growth of fetuses is relatively constant but that females in poor physical condition may not be able to support as large a fetus as those in good condition so parturition occurs earlier

This may have sone effect on information on this

Fig 12 demonstrates some other interesting points There is no difference in the ratio and the maximum fetus size of males and females

~

(

~

-s 2 0 C

U)

0

--

ez ()

~) I) middot _t t- (

j -~middot -~

Reproduction - 14 - March 1971

This infeTs although dolt2s not prove an identical grmgtlth rate afld birth weight for males and females The circles on Fig 12 show the ~eight-length ratio of pups These fall below the curve formed by fetus weight-length ratios indicating that pups of a given length are some-Jhat heavier than fetuses of the sarne length This indicates an increase in weight immediately after parturition

Sex Ratio at Birth

Kenyon (1969) found that of 58 fetuses for which sex could be determined~ 45 percent vJere male and percent were female He assumed an equal sex ratio until a larger sample could be obtained

In the present study 111 fetuses were male and 144 ~vere female When this sample is combined with Kenyonts the sex ratio of 313 fetuses is 44 percent male and 56 percent female

Sex Ratio of the

In the process of harvesting capturing animals for transplants and delineating male and femalen areas it has become apparent that there are more females in the population than males Our general ion is that perhaps 60 to 65 percent of the sea otters are females Kenyon (1969) demonstrated that more juvenile males than females are found dead on the beaches of Amchitka lhe age structure of harvested in 1968 indicates that males may not live as long as fetnales

The lower birth rate higher juvenile mortality rate and slightly shorter life expectancy of males could account for the unbalanced sex ratio in the population

The sex ratio may vary from one island to another Presumably there -vmuld be more females in a population that has reached the carrying capacity of the habitat where juvenile mortality is higher There is some indication that males are the first to expand into new areas of unoccupied habitat Therefore in recently populated anas one might find more males However this situation vould be

Lit

The normal litter size of the sea otter is one This has been recorded by almost every observer since Steller Barabash-Nikiforov (194 7) reported twins in utero and mentioned other reports of twin fetuses from the early days of SnoF (1910) recorded a set of neHborn tdns In more recent studies~ both Sinha et al (1966) and Kenyon (1969) found reproductive tracts with two corpora lutee but in all cases only one fetus las present In thousands of hours of observation by many observers no tbullvin pups have been reported

In the present study 24 instances of multiple ovulations were recorded Five of these resulted in hv-in and one in triplet fetuses The information is summarized in Table 8

Table 8 Multiple Corpora Lutea Fetuses

Total Pregnant Females 565

Implanted Pregnancies 316

Unimplanted with 2 CL (corpora lutea) 9

Implanted Pregnancies with ) L CL and 1 Fetus 8

Implanted Pregnancies Vlith 3 CL and 1 Fetus 1

Implanted Pregnancies gtvith 2 CL and 2 Fetuses 5

Implanted Pregnancies middotwith 3 CL and gt Fetuses 1

Total Nultiple Ovulations

I

Percent Nultiple Ovulations 42

Percent of Pregnancies tvith Nultiple Fetuses 19

Corpora Lutea per Pregnancy 105

Fetuses per 102

Reproduction - 15 - lvlarch 1971

From these data it appears that in over 4 percent of the estrus cycles more than one ovulation takes place Of these about half result in the development of more than one fetus In most cases the were of a relatively large size and probably would have been born normally

All multiple fetuses appeared to be the result separate ovulations All had separate placentas and in some cases were different sexes Four tracts with twins had both fetuses in the same horn one had one in each horn and the tract with triplets had one fetus in one horn and t~vo

in the other Another tract had skeletal remains of triplets in one horn that were being resorbed Another tract appeared to have had five fetuses in one horn but they were almost completely resorbed One might infer that physically there is not enough room in one horn to accomn10date more than two fetuses for any length of time

With the exception of the newborn tvins reported by Snow (1910) there are no docu_mented records of female sea otters with tvin pups There are occasional unconfirmed reports of twin pups but as Kenyon (1969) points out a female may tolerate a pup in addition to her ovm but it is unlikely that a female could care for more than one pup It is unlikely that more than one pup survi~es In any case orily 2 percent of the pregnancies produce more than one pup 1middot1ultiple births cannot be a significant factor in the productivity of sea otter populations

The presence of a corpus luteum without gross evidence of implantation t-7as assumed to be an unimplanted pregnancy No attempt ~vas made to locate unimplanted blastocysts As a result e have little information concerning the survival of the blastocyst Of 206 reproductive tracts classified as nulliparous on the basis of the appearance of the uterus 12 had one corpus albicens and one had two Some of these may have been large attritic follicles Others may have ovulated and even conceived but implantation did not occur All of these cases represent the failure of the antmals first estrus

A total of 16 resorptions or possible abortions were identified (Table 1) The follmJing is a brief descdption of these ~7i th possible causes of some

Des

Resorption of blastocyst or ovum (corpus luteum 4 degenerating no evidence of ion)

Failure at time of (possibly because 1 of growmiddotth inside horn adjacent to implantation site)

Resorbed or aborted fetus (implantation at end of horn) 3

Resorbed fetus (umbilicus tvTisted tightly) 1

Resorbed fetuses (three or more fetuses in one horn) 2

Resorbed fetus (cause unknmm) 5

Reproduction - 16 - March 1971

Six of these resorptions may have been caused by a lack of room for growth of the placenta because of crowding from other placentas a growth or the end of the uterine horn These six and the one vith the twisted umbilicus are probably the result of random events or accidents

The numbers of resorptions are too small to accurately measure the frequency of such occurrences although the relatively large number in the Tanaga sample (Table 1) may be a reflection of the physical condition of the animals Part of this number is probably due to the fact that there are more pregnant animals in the sample In general the samples with the most pregnancies also conta~~ned the most resorptions Half of the Tanaga resorptions can be attributed to random events but five vere still due to unknown causes Eight of the 10 resorptions fo1md on Tanaga came from a 20-mile stretch of shore about 25 percent of the harvest area

While concrete conclusions cannot be drawn from these data they do raise the possibiliy that under certain conditions 5 percent of the pregnancies may fail

near Parturition

Collecting methods were such that the presence of a pup with a female was often overlooked Therefore the lack of a pup with a post-partum or lactating female did not mean that the pup had died However 14 females shmving of recent pregnancy were estrus presumably having lost their pup shortly before or shortly parturition

Again the magnitude of this mortality cannot be determined but it appears to be of the same order as the in utero mortality

after

Reproductive tracts were subjectively classified as post-partum or anestrous on the basis of degeneration of the corpus luteum (or appearance of the corpus albicans) the size of the horn of pregnancy and appearance of the placent scar In an effort to get an idea of how long it takes for the tract to return to normal the size of pups accompanying females in each category is listed in Table 9

It appears that females pass the subjective boundary from post-middotpartum to anestrus Hhen the pup about 10 lb and is around 75 to 80 em long

We do not know a great deal about the early growth of a sea otter pup but judging from the cur1e e9tablished based on cementum deposition of older animals and from tooth eruption a 10 lb pup is probably in its second month of life

Table 9 Pups Accompanying Pos artum Females

Sex llleight

M 2 lb F 3 F 4 M 5 F 55 F 625 F 775 M 10 F 11

Total Length

47 em 52 56 60 65 65 70 81 (borderline post partum-anestrous) 82 (late post-partum)

Pups Accompanying Anestrous Females

F 10 lb 78 em M 11 79 M 12 8l F 13 83 M 13 92 H 16 83 M 17 91 1 19 87 M 19 90 F 21 107 F 22 96 1 22 101 M 2Llmiddot 97 1 26 100 F 28 103

Reproduction - 17 - Narch 1971

Frequency of Breeding

Steller (1751) reported female sea otters with ne~vborn pups also accompanied by another pup that seemed to be no more than a year old Nurie (1940) recorded an instance of copulation by a female accompanied by a pup and Jones (1952) caught a young pup simultaneously gtvith a copulating pair Lensink (1962) felt that few females breed until the pup is a year old but mentions females with small pups also accompanied by large pups

I believe that some of thes2 observations may be misinterpreations of the situation Females will occasionally leave small pups for a time often near other animals The pups Lensinkmentions would be 2 years old and probably veigh 30-35 lb Sea otters are gregarious and subadults may remain near other animals appearing to be accompanying their mother I suggest that most of Stellers and Lens inks large pups ere such cases

Females gtvi th pups probably do breed occasionally but the most recent evidence indicates that this rarely occurs Kenyon (1969) records no instances of females accompanied by pups copulating and states that he found no females- knm7n to be accompanied by a pup that were This holds true for the animals collected in the present study However the nature of the harvesting operation is such that many females accompanied by pups were not recorded as such Therefore we are forced to look at lactating and assume that they were accompanied by a pup at the time of collection or shortly before le 10 sm1marizes the reproductive condition of the lactat females In the 1967 and 1968 samples some animals vith enlarged marmary middotmre listed as lactating As a result some females with near tenrr fetuses Here included Because this condition is considered to be associated with the unborn pup rather than a previous pup these animals were with the anestrous animals in le 10 In 1969 and 1970 the presence of milk was used as the sole criterion and no term animals we-rmiddote in the sample

Of 246 lactating females only one had an ted fetus (excluding the near term animals from 1967-68) Several others had small corpora lutea indicating that they recently become or possibly had large

vhich had luteinized but tvere not actually pregnant The remainder had follicles

This information that accompanied by a pup rnay enter proestrus and even ovulate but that they only rarely mate That they enter estrus is subs iated by the f2ct that the number of corpora albicantia in many tracts equals the age of the amplimal minus the three years prior to nBturity (see les 4-7) This indicates that large follicles tend to develop year after sexual nl8turi ty is reached whether the female is accompanied by a pup or notlt

Some of those lactating females with s1all corpora lutea may have recently weaned or othenvise lost a pup and are already pregnant again Malkovich (1937) took a pup least 3 months old) away in cap She mated 12 days later here she had the male It appears that a female enters estrus shortly after Heaning

Table 10 Reproductive Condition of Lactating Females

Location

Bay of Is Adak I Sept 1967 9 1 100

Allchitka I Sept-Oct 1967 17 2 105

Mid Pt - Blind Pt Oct 1968 19 3 136 Adak I

Bay of Is Adak I Oct 1968 22 4 154 3

Kanaga I Oct 1968 32 s- 135

Al1chitka I July-Aug 1969 4 2 333

iTanaga I May 1970 56 10 152

Delarof Is May 1970 18 3 143

Amchitka I May 1970 36 3 77

---~--middot--middot----~-~----~----middot~-middot--~----~----- -------~-~ middotmiddot--middot-shy

TOTAL 213 33 134

Anestrous or pregnant with term fetus

]_ Large follicles or corpus luteum present

3 Includes one implanted pregnancy with small fetus

Reproduction - 18 - March 1971

The question of hml long it takes for a female to enter estrus again after losing a small pup arises Eleven tracts of the 1970 sample and ttvo of the 1968 Kanaga Sampnple shmJed signs of being pregnant recently) but were in proestrus These animals had an enlarged horn usually middotlith a slightly pigmented area but no actual placental scar a11d a recently fonned corpus albicans However they also had enlarged follicles and the tvalls of the uterus were typical of proes trous animals Another animal was similar but appeared to have already ovulated and a corpus luteum was present In these cases the female has lost a pup either through abortion or vithin the first veeks after parturition and has started into another estrus cycle Three or four of these had follicles vhich may have started to become attritic This may be because the uterus had not recovered sufficiently from the last pregnancy

The implication of this information is that with a normal pregnancy and successful rearing of a pup the female becomes pregnac1t only after the pup has been weaned This n~y be a year after parturition However if the pregnancy fails or if a pup dies even at birth the female may become pregnant again in a few weeks rather than _waiting the full year

Kenyon (1969) suggests that the period middotbetween and the next estrus may long because he found animals that tvere not lactating but had a faint placental scar and an indistinct but recent corpus albicans These animals ltJere cLtssi as anestrus in the present study They are undoubtedly bet1veen veaning of their last pup and the next estrus The number of suh animals is t durirg Sspte12~ cnd October At this time the rate of estrus increases a1d the number of anestrous animals decreases sharply the next feJ months Therefore many of these anestrous anirrals become pregnant 1lithin a month or two Kenyon (1969) suggests that if the female keeps the pup for about a year that the period between births may be smreltmiddotJhat greater than two years He is assuming a 12-13 month gestation period However the present study shows that the estimate of a 12-13 month tation period was based on an inadequate sample Estimates in the present study put this period at closer to 8 or 9 months Presumably the anestrous nonlactating females are in the 3 to 4 month period

This does not change Kenyons (1969) point that the and the next estrus may long It indicates that in a normal cycle it may be months Hmvever as it vvas shmvn above~ it is possible for a female to enter estrus very shortly after losing a pup

We have demonstrated that sea otters breed at all times of year and it is possible for a female to become pregnampJt again after losing a pup sooner than she vould have if the pup had survived HmmiddotJever we have also demonstrated that annual of b and pupping occur and that these are similar in different years and in different populations The assumption can be made that the average length of the entire reproductive cycle is some multiple of a year

Kenyon (1969) assumes that tbe pup remains with its mother for about a year ~-Ieights and measurements cementum deposition skull changes~

Reproduction - 19 - March 1971

observations in the wild and in captivity all support this assumption although it cannot be said with certainty that it averages 12 months It could be s tly ITDre or less

Assuming about 12 months for rearing the pup 8 to 9 months gestation plus a fev months rest bet~veen and the next estrus the average reproductive cycle takes two years

This is supported by Fig 4 which shows that slightly over half the mature females are pregnant in a year Kenyon (1969) pointed out that his January and Harch samples gtvere biased against females with pups a1d therefore in favor of pregnant females because of selective hunting This bias applies to the fall sa~ples as well The summer samples were biased similarly because pregnant females appeared to be less resistant to handling during capture Therefore the percent of pregnant females is shown to be higher than lt actually Has -rhile no correction factor can be applied to all samples the evidence indicates that the percent of pregnaJt females is usually around 10-middot15 lmrer than in the sample For example the 1970 Alnchitka sltLrnple t-ras less biased and was about 9 percent lower than the Tanaga sample vrhich is l)nm-rn to be more biased Also on a June skiff survey 7 percent of the animals counted had pups that were readily identified If ~re assune that this is the nunber of animals avoided it can be calculated that 15 of the sexually mature females all of vhich are t middotTen~ avoided Actually this will vary with the hunter weather time of year etc Hmvever it indicates the order of magnitude of the bias

Using this as a rough correctton for the data in 4 we find that about half or slightly fe-Jer of the sexually mature females are pregnant each year or as indicated ly the average female has a pup every two years

Reproduction - 20 - March 7 1971

Conclusions

1 While sea otters may breed or pup at any time of year there are seasonal fluctuations in the illliTlbers of females in each reproductive stage The pattern of these fluctuations remains the same from year to year and population to population

2 Breeding activity is highest in September October and November

3 More implantations occur during January February and March than at other times of year

4 The birth rate is highest during April Y~y and June

5 Tlvo and a half to three times as many females breed during the peak breeding months as during the months of lowest breeding activity

6 Similarly two and a half to three times as many females pup during peak pupping months

7 During the period of lovest b activity seually mature females breed at a rate of 2 to 3 percent per month During the peak of breeding 7 to 10 percent per month breed

8 During the period of lov1est pupping activity 2 or 3 percent per month pup and during the peak of pupping 7 to 10 percent per month pup

9 The gestation period about 8 to 9 months on the average About half of this time is ~~ ted period

10 Female sea otters become sexually mature ihen 3 to 4 years old

11 Females may ovulate on an average of once a year after sexual maturity is reached

12 Females continue to breed at Cn old age

13 Sea otters retain certain characteristics common to the reproductive cycles of many other mustelids however exceptions are common Scheduling of the reproductive cycle during the year may have some advantages or it may be a trait that has not been entirely lost through evolution

14 Sixty percent of the near term fetuses are cephalicly oriented and 40 percent are candllly oriented

15 Sea otters probably birth both on land and in the water

16 Blastocysts rarely cross from one uterine horn to the other

17 Consecutive pregnanctes may occur in the same horn The side of pregnancy appears random

Reproduction - 21 - March 1971

18 Implantation usually occurs in the central third of the horn but it may occur any~rhere in the horn

19 Birth weights may range from 900 g to over 2500 g however most pups weigh 1800 g to 1900 g at birth

20 Females from areas gtvhere the population is declining because of food shortages may pup earlier and as a result the pups 1 birth weight may average 200 g lmver than nonnal

21 Male and female fetuses gr01middot1 at the same rate ampJd are the same size at birth

22 Body condition of the female has little effect on the growth rate of the fetus

23 There is a rapid ~Jeight gain shortly after parturition

24 The sex ratio at birth is 44 percent male and 56 percent female

25 The normal litter size is one hmmiddotever in 2 percent of the implanted pregnancies two or fetuses survive to near term

26 Multiple ovulations occur in 4 percent of the estrus cycles

27 es do not appeErc- to be able to more than two fetuses in a single horn

28 Up to 5 percent of the p may fail dne to in after implantation

29 An unknown but probably significant number of first pregnancies fail before implantation

30 The uterus is usually considered recovered from a pregnancy when the pup vleighs about 10 lb and is probably in its second month of life

31 Sexually matu1middote females normally have one pup every two years

32 Females vrith a pup may ovulate but rarely mate

33 Slightly less than half of the sexually mature females are in any given year

34 A female may enter estrus within a few weeks of losing a pup Consequently she may become agaln gtvi thout waiting the same length of time as if the pup hacl survived

35 Hhen a pup is weaned normally there is a longer period perhaps 3 or 4 months between weaning and the next estrus

Reproduction - 22 - March 1971

CITED

Barabash-Nikiforov I I 1947 The sea otter (Kalan) Soviet Ministrov RSFSR Translated from Russian by Dr A Birron and Z S Cole Israel Program for Scientific Translation 1962 227 pp

Hugget A St G and W F Widdas 1951 The relationship betlveen marmnalian foetal -reight and conception age Jour Physiology 144306-317

Kenyon K W 1969 The sea otter in the eastern Pacific Ocean USFWS North American Fauna No 68

Malkovich T A 1937 The sea otter in captivity Priroda No 381-87 Translated by J T Naximovitch 1966 Fisheries Research Board of Canada Nanaimo BC Translation Series 657

Sinha A A C H Conavay and K t-J Kenyon 1966 Reproduction in the female sea otter J Hildl Ngrrit 39(1)121-130

Snow H J 1910 In forbidden seas Edward Arnold London 303 pp

Wright P L 1963 Variations in reproductive cycles in North Arrierican mustelids In Enders A C ed Delayed tation Univ of Chicago Press 318 pp

  • SEA OTTER STUDIES - 1970
    • Surveys
    • Harvests and Transplants
      • Sexual Segregation in Sea Otter Populations
      • Reproduction in the Female
        • AERIAL COUNT OF SEA OTTERS- SOUTHSIDE OF THE ALASKA PENINSULA
          • Description of Conditions During March 1970 Aerial Count of Sea Otters
          • Discussion of Results
          • Sanak Island-Sandman Reefs
          • Shumagin Islands
          • Sutwick Area
          • Augustine Island-Cape Douglas
            • SUMMARY OF SEA OTTER SURVEYS
            • Amagat 1 to False Pass Ikatan Peninisula Cape Lazaref
            • Sanak Is
            • Deer 1
            • Pavlof Is
            • Northern Shumagin Is
            • Cold Bay 1
            • Kupreanof Peninsula
            • Mitrofania 1 to Kupreanof Pennisula
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