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1966

Parasitic Copepods of Some Antarctic and New Zealand Fishes Parasitic Copepods of Some Antarctic and New Zealand Fishes

David E. Zwerner College of William and Mary - Virginia Institute of Marine Science

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Recommended Citation Recommended Citation Zwerner, David E., "Parasitic Copepods of Some Antarctic and New Zealand Fishes" (1966). Dissertations, Theses, and Masters Projects. Paper 1539617395. https://dx.doi.org/doi:10.25773/v5-cmwr-dy73

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PARASITIC COPEPODS OF SOME ANTARCTIC

AND NEW ZEALAND FISHES

A Thesis Presented to

The Faculty of the School of Marine Science The College of William and Mary in Virginia

In Partial Fulfillment Of the Requirements for the Degree of

Master of Arts

byDavid Eric Zwerner

1966

APPROVAL SHEET

This thesis is submitted in partial fulfillment of the requirements for the degree of

Master of Arts

David Eric Zwerner

Approved

William J. Hargis, Jr., Ph. yU

' Marvin L. Wass„ Ph.D.

Q t L)ceWillard A. Van Engel, Ph.M.

i i

ACKNOWLEDGMENTS

Appreciation is expressed to Dr. William J. Hargis, Jr., of the Virginia Institute of Marine Science, for support, guidance, and assistance received throughout the course of this study. I also wish to thank Mr. W. Stanley Wilson (formerly of the Institute) who collected and identified the New Zealand host material, and collected part of the Antarctic host specimens, Dr. Harry L. Holloway, Jr. of Roanoke College, Salem, Virginia who collected and identified the remaining Antarctic host material, and Dr. Hugh H. DeWitt of Stanford University, who checked or provided identifications of the difficult Antarctic fishes. I am obliged to the following members of the Virginia Institute of Marine Science for their aid: Messrs. William A. Dillon and Adrian R. Lawler who examined host material; Mrs. Sibyl R. Lawler who translated pertinent French literature and assisted in examining host material; and Mr. Robert S. Bailey for help with the photographic work.

I am grateful to Dr. George D. Grice of the Woods Hole Oceanographic Institution and especially to Dr. Harry C. Yeatman of the University of the South, Sewanee,

V

Tennessee for their assistance in the identification of the harpacticoid copepod described in this paper.

iii

I am further Indebted to Dr. Thomas E. Bowman and Mr. John C. McCain of the Division of Crustacea, U. S. National Museum, for their help in obtaining the literature necessary for this paper.

/

To my wife, Marjorie, who has been a mainstay of encouragement throughout the course of this study, I extend special thanks for her assistance in the preparation of the typescript.

This research was supported by Grant No. 13853 from the United States Antarctic Research Program of the National Science Foundation.

iv

TABLE OF CONTENTSPage

ACKNOWLEDGMENTS................................... iiiLIST OF TABLES............................. ...... viTEXT FIGURE...................................... viiLIST OF PLATES................................... viiiABSTRACT............................. ixINTRODUCTION. ......... ....................... .. 2MATERIALS AND METHODS. . ........................... 4RESULTS AND DISCUSSION............................ . 10

Neo s cute 11 id ium yeatmani........ • • • IINesippus incisus.................. ........ 16Eudactyiina sp......................... . • • • 22Kr^yeria mustell........ ................... 27Nothadichelidium polyprionum. ................ 34Neono thad iche1id ium leptocephali. .... 39Chondracanthus genvpteri •••••...*.. 42Clave I lodes intermedius. .... • •. .... 45N ob flnchis bie lloo JLeria............. ... 50

SUMMARY................... ................ *..... 54LITERATURE CITED... ............... 55VITA

v

LIST OF TABLESTable Page

1. Parasitic copepod check list for Antarctica,Australia, and New Zealand,........ . *. 60

2, Systematics and hosts of the parasiticcopepods treated................. 66

vi

TEXT FIGUREFigure Page

I. New Zealand.................... 6

vii

PlateLIST OF PLATES

Figures

I. Neoscutellidium veatmani gen. et sp* nov....... 1-11Nesjppus incisus Heegaard, 1962...... . 12-22

III. Eudactvlina sp............................ 23-34IV. Krdveria musteli sp. n o v . • 35-44

cV. No thad ichelid ium polyprionum gen. et sp. nov... 45-54Neono thad iche1id ium leptocephali gen. et

sp. nov................ 55-63VI. Chondracanthus genypteri Thomson* 1890...... 64-79VII. Clave1lodes intermedius (Quidor, 1906)

Wilson, 1915................ ........ 80-85Naobranchia helico lena sp. nov....... .... 86-91

viii

ABSTRACT

During March I960, 616 specimens of 35 species of fishes were collected from the marine waters of New Zealand and examined for parasites. Of „these, 258 individuals, or 38.6 per cent, representing 21 species were infected with parasitic copepods. From January 1961 to February 1962 and during November and December 1964 a total of 599 specimens of 13 species of fishes were collected from the waters adjacent to the Wilkes and McMurdo Sound stations in the Antarctic.Only 37 individuals, or 6.4 per cent, representing four species were infected with parasitic copepods.

Of the 18 different species of copepods found on the New Zealand fishes, seven are considered herein. They include Kr^veria mustell sp. nov., No thad iche1id ium polyprionum gen. et sp. nov., Neono thad iche1id ium leptocephali gen. et sp. nov., and Naobranchia helicolena sp. nov. A fifth, Eudactylina sp., is probably new. Two are redescribed; Nesippus incisus Heegaard, 1962, and Chondracanthus genypteri Thomson, 1890.One new host record is recorded and one new locality record established.

Only two species of copepods were found on the Antarctic fishes. Neoscutellidium yeatmani gen. et sp. nov., is described and Clave1lodes intermedium (Quidor, 1906) Wilson, 1915 is redescribed. One new locality record is established.

ix

The superfamily Dichelesthioidea, Yamaguti, 1963 is emended to accommodate the new family Nothadichelidae which is erected for No thad icheIid ium gen. nov. and Neono thad icheIid ium gen. nov.

x

PARASITIC COPEPODS OF SOME ANTARCTIC AND NEW ZEALAND FISHES

INTRODUCTION

This systematic study is concerned chiefly with parasitic copepods collected from the gills of marine fishes taken in New Zealand and Antarctic waters. A total of 18 copepod species was collected from New Zealand waters. These species represented seven families and nine genera and were collected from 18 families and 19 genera of fish hosts, a total of 621 copepods from 258 fishes. Seven of the New Zealand copepod species and the only two collected from Antarctic fishes are discussed. Table 1 lists those species of parasitic copepods reported in the literature for Antarctic, Australian, and New Zealand areas. Copepods are listed by family and the host(s), locality, and sources of information are given for each.

Although about 70 species of fishes are known to occur south of the Antarctic convergence (DeWitt and Tyler, 1960) only a dozen (including the new species described in this paper) parasitic copepods have been reported from this area.It is not known how many of the 70 fish species have been examined for parasitic copepods but out of 13 host species in our collection two species of copepods infected four species of fishes.

2

3

Investigations into the parasitic copepod fauna of New Zealand fishes have been rather infrequent (and the great number of new species that probably exist deserves more intensive pursuit). Thomson (1890 a,b) and Bassett-Smith (1898, 1899) were early workers in the New Zealand area, followed by Hutton (1904), Brady (1910), Quidor (1913), Phillips (1922), and more recently Heegaard (1940, 1943, 1962) who works on copepods from the entire southwestern Pacific Ocean. Most recent of the investigators is Hewitt (1963,1964 a, b, c, d) who is presently working on the caligids of New Zealand. At present 48 species representing 12 families of parasitic copepods have been reported from the New Zealand area and the same number of species in 14 families from Australian and Tasmanian waters. Because over 400 species of fishes are presently known from New Zealand waters and because parasitic copepods are relatively specific in host selection many more species should be present for discovery.

Over 80 per cent of the New Zealand and Antarctic fishes examined were infected with both monogenetic trematodes and parasitic copepods. Since the trematode data are already published or in preparation, an interesting examination of the theory of host-specificity might involve comparative study of copepod-host data and monogeneid-host data. Such studies might produce insight into the zoogeography and phylogeny of both host and parasite.

MATERIALS AND METHODS

New Zealand fishes were collected in March 1960 by Mr. W. S. Wilson from the commercial fisheries located in the following areas: Timaru, Dunedin, and Akaroa Harbor (Christ­church) on the east coast of the South Island, Cape Campbell in the Cook Strait area, and Auckland, North Island. Trawls, gill nets, and Danish seines were used to capture the fishes. Mr. Wilson identified the fishes using the keys and systematics of Waite (1923), Graham (1956), and Parrott (1957, 1958) and had the scientific names checked and updated by Miss M. K. McKenzie (Marine Department, Fisheries Research Station, Wingfield Street, Wellington, New Zealand).

Antarctic host material was collected from two areas: Mr. Wilson collected in the waters off Wilkes Station from January 1961 to February 19625 Dr. H. L. Holloway collected in McMurdo Sound waters in November and December 1964. Fishes were captured mostly in traps, a few by longline and trawl. Fishes from Mr. Wilson's collection were identified by Dr.Hugh H. DeWitt of Stanford University while those of Dr. Holloway's collection were identified at McMurdo by Dr. Donald E. Wohlschlag, also of Stanford University.

For the New Zealand species locality descriptions are given in the following format: nearest town or prominent

4

5

landmark followed by its coordinates in parentheses, province of landmark, and specific island location (North or South Island); next, approximate location of collection site with range in nautical miles from the landmark and bearing in degrees true; depth and bottom type given in parentheses.See Text Figure 1 for area involved. The position of each collecting station in the Antarctic is given by coordinates along with certain descriptive data as depth, bottom type, and nearest base.

Gill parasites were collected using the mass collecting technique for monogenetic trematodes as described by Hargis (1953). Under the procedure, the excised branchial basket or, preferably, the individual gill arches were immersed in a saturated solution of Chloretone (hydrous chlorobutano1, Parke- Davis) in filtered sea water for one or two hours and occasionally agitated. Killing, fixing, and preserving was accomplished by adding enough formalin or AFA (acetic-fcrcmalin-alcohol) to the anesthetic solution to bring the concentration of formalin to 10 per cent. This technique apparently did not harm the parasitic copepods although prolonged exposure to the un­buffered formalin or inherently acid AFA might have been expected to dissolve the exoskeleton. This technique permitted rapid processing of large numbers of fish. In the laboratory, parasites were separated from host material using a stereo- micro scope and stored in a solution of 70#per cent ethanol and 5 per cent glycerine.

6

NEW ZEALAND(STATUTE MILES)

HAURAKI

AUCKLA

NORTH ISLAND

6TON°* Sr

APE AMP8ELL

CHRISTCHURCHb a n k s p e n in s u l a

AKANOA

tvSOUTH ISLAND

ITA60 HARBOR

V?

Text Figure 1. New Zealand, (after Dillon and Hargis, 1965)

7

Many methods of staining and mounting the copepods were tried^ but none was found superior to lactic acid as a temporary mountant, methyl blue in lactic acid as a stain, and Hoyer's as a permanent mounting medium as recommended by Gooding (1957, 1960) and Lewis (1964a, b). Lactic acid (LA) when used as a temporary mountant cleared satisfactorily, softened, and gave good definition of hard parts in addition to giving good preservation.

Typical preparation procedure involved placing the specimen directly from 70 per cent ethanol into a LA wash for clearing, then mounting it in LA. Fairly chitinous copepods, such as caligids, were slowly boiled in LA for two or three minutes and then mounted. Boiling not only hastened the clearing process but it also broke down the soft tissues which often obstruct viewing. By adding small amounts of methyl blue in LA to the hot LA, staining and clearing occurred together. (Copepods which have been dry for years have been reclaimed into excellent mounts by use of the boiling LA technique.)Methyl blue was found to provide the contrast necessary for the microdissection of very small specimens. Copepods were easily disjoined in LA and either temporarily mounted in LA

Stains tested were; alum cochineal, borax carmine, Delafield's hematoxylin, eosin-Y (ethanol), fast green (ethanol), Harris' hematoxylin, iodine (ethanol), Mallory's triple, picric acid, picro-acid fuchsin, Reynolds.Mountants tried: ethanol, Euparal (Turtox). glycerine,, glycerine jelly, Piccolyte (Turtox), polyvinyl lactophenol.

8

or transferred directly to Hoyer’s. (During the course of the work it was found that washing an appendage in a saturated chloral hydrate solution before transferring it to the Hoyer’s minimized the initial and temporary blurring of the specimen due to the mixing of two solutions of different refractive indices.) Hoyer’s is slightly hygroscopic and for that reason coverslips were ringed with Permount (Fisher) to prevent the mount from becoming soft.

Where possible 10 good specimens of each species were examined (both male and female if available). Males were considered mature if they were attached to females. Maturity of non-ovigerous females and unattached males was judged on the basis of the development of the genital segment.

Measurements were taken of several copepods in each species and the average of the measurements determined.Inasmuch as measurements and ratios have not been commonly employed as an aid in the differentiation of species of copepods, no statistical analysis was made of the measurement data other than to report the mean and range of values. All measurements of curved structures were across the lines sub­tending the greatest arcs described by those structures. All measurements were made with a calibrated filar micrometer and are given in millimeters.

Drawings were made with the aid pf a microprojector (Bausch 5c Lomb Tri-Simplex) or camera lucida.

9

The morphological terminology follows that of Wilson (1915, 1922, 1932) and Yamaguti (1963). The overall taxonomical scheme employed in this work is that used by Yamaguti (1963).The individual descriptions are standardized and conform to the following format: 1. description of whole mount fromanterior to posterior including segmentation and body armament;2. antennae and mouth parts from anterior to posterior;3. thoracic legs from first through fifth. In this way the characters of identification with the higher taxa are discussed first. When both sexes are present, the female is described first.

Armature formulae used for the description of appendages in this work are of two types. The first, utilized for most species, is a simple formula employing arabic numerals for setae and Roman numerals for spines with the combination of numbers forming a couplet for each segment. The second armature formula, which was developed by Shiino (1952) and elaborated on by Lewis (1964a), employs capital and lower case letters to cryptically describe setation, spination, etc. of inner, middle, and outer margins of appendages. This formula was used only in the description and comparison of the more complicated structures on Neoscutellidium yeatmani gen. et/', sp. nov.

RESULTS AND DISCUSSION

The nine copepods treated, together with their classification and host species are listed in Table 2.

Class Copepoda (Latreille, 1831)Order Harpacticoida (G. 0. Sars, 1904)

Suborder OligoarthraFamily Tisbidae Stebbing, 1910

Subfamily Tisbinae Stebbing, 1910

Genus Neo s cute11id ium gen. nov.Generic diagnosis.-Harpacticoida, Oligoarthra, Tisbidae,

Tisbinae. Female: Body depressedj first thoracic segment fused with head; head and first thoracic segment covered dorsally with large cephalic shield; rostrum visible. Abdomen composed of five segments, the first two fused to form the genital segment. First antenna composed of eight segments. Second maxilla and maxilliped (characteristic appendages of the genus) 2-segmented, in form of simple hooks.First leg with 3-segments per ramus and adapted for grasping by presence of terminal geniculate setae; legs two through four natatory, with three segments per ramus; fifth leg reduced, lamellate, with one segment per ramus.

Type species: Neoscute11idium yeatmani gen. et sp. nov.

10

11

Discussion: Neoscutellidium gen, nov, clearly belongs to the subfamily Tisbinae and most closely agrees with Claus1s genus ScuteHidium from which it differs in the following ways:1. the rostrum in Neo s cute11id ium is visible in dorsal aspect;2, the cephalic shield is more or less orbicular in shape rather than conical as in Scutellidium: 3. the first antenna has eight segments instead of nine as in Scutellidium:4. both second maxillae and maxillipeds in Neo s cutel1id ium are modified into large, 2-segmented prehensile hooks; 5, Neo­scutellidium is a symbiont of fish while Scute1lidium in most cases is a free-living psammophil (Vervoort, 1964),

Neoscute1lidium most closely resembles Scutellidium,

Neo s cute11id ium yeatmani gen, et sp. nov.(Figures 1 to 11 )

Host and number examined: Rhigoohila dearborni DeWitt, 1962;family Zoarcidae. (92).

Location: Gills,Locality: McMurdo Sound area, Antarctica (77°50,S, 166°45IE).Number of copepods recovered: 7.Number of copepods examined: 7.

Description, Females Body depressed, tapering posteriorly, 0.793 (0.697 - 0.913)long. Large cephalicshield wider than long, 0.341 (0,314 -0.368) wide by 0,242 (0.206 - 0.280) long, covering dorsal aspect of head and fused first thoracic segment. Small triangular rostrum pro­truding from under anterior margin of shield. Thoracic

12

segments each bearing a pair of pereiopods, distinctly joined one to another with the exception of the first as

o -

noted. Large genital segment, 0.134 (0,132 - 0.139) long by 0.119 (0.107 - 0.128) wide, representing a fusion of first and second abdominal segments; segments three through five slightly telescoped. Elongate fureal rami approximately one- half length of latter three segments, slightly diverging.

First antenna moderately long (0.149), 8-segmented, with following number of naked setae from segment I to VIII:1-5-7-1-1-1-2-4*

Second antenna biramous. Exopod apparently un­segmented, bearing five naked setae. Endopod 2-segmented,bearing six naked setae, four being geniculate; segment IIwith one spine, segment I devoid of armament. Length as follows:

Basis: 0.076Exopod: 0.030Endopod, total: 0.064

Segment1: 0.039Segment II: 0.024

Mandible with well developed palp; cutting edge of praecoxa erescent-shaped, with fine hairs. Coxa and basis fused to form protopodite armed with two strong setae. Exopod

_ small with two naked setae at terminus. Endopod indistinctly2-segmented, with one naked seta on first segment and three on second segment. Length as follows:

13

Praecoxa: 0.065Protopodltd setae Exopod:Endopod:

0.0430.0220.066

First maxilla indistinct, apparently with well developed palp. Exopod (0.026 long) with two naked setae; endopod (0.030 long) with three shorter setae.

Second maxilla and maxilliped, both simple, unarmed,2-segmented; elongate hooks adapted for prehension and originating slightly posterior and medial to origin of first maxilla. Length of second maxilla exceeding that of cephalic shield (segment Is 0.120, segment II: 0.124, TL: 0.244); total length of maxilliped slightly less than cephalic shield (segment Is 0.111, segment II: 0.095, TL: 0.206). Appendages differing in shape and in amount of curvature expressed in terminal segment; second maxilla hooking very abruptly in the last distal fifth of the segment while the maxilliped makes a graceful curve for the entire length of the segment.

ramus except in reduced fifth leg; first leg prehensile; second through fourth natatory. Armament and measurements of thoracic legs as follows:

Thoracic legs all biramous with three segments per

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Eggs spherical, large (0*113 in diameter), carried ventrally in a loose egg sac*

Ma le: Unknown •

Discussion, This appears to be the first report of a tisbid copepod parasitic on a fish, although one has been reported from an octopus. This species is named for Dr, Harry C, Yeatman who kindly assisted in identifying this copepod.

Order Caligidea Stebbing, 1910Superfamily Caligoidea Dana, 1853

Family Pandaridae Wright, 1877 Genus Nesippus Heller, 1868

Nesippus incisus Heegaard, 1962 (Figures 12 to 22)

Hosts and number examined: Galeorhinus australis Macleay,Southern Tope or School Shark; family Carcharinidae,<i>.Mustelus antarcticus Guenther, Gummy Shark; familyCarcharinidae. (21).

Location: Gills.Localities: 1. Akaroa {43°48(S, 172°58,E), Canterbury

Province, South Island, N. Z.; Akaroa Harbor (4-10 fathoms; mud, sand, horse mussels),

2. Timaru (44°24'S, 171°15*E), Canterbury Province, South Island, N,*Z. ;a. 8.7 miles, 068° from Timaru (9 fathoms; sand)*

17

b. 20.0 miles, 068° from Timaru (16 fathoms; sand, mud).c. 45.2 miles, 068° from Timaru (28 fathoms; sand).

Previously reported host and locality: Mustelus antarcticusfrom New South Wales, Australia (Heegaard, 1962).

Number of copepods recovered: G. australis 3, M. antarcticus 26. Number of copepods examined: 10.

Description. Female: Typical caligid shape, total length 4.753 (4.416 - 4.985). Carapace elliptical, wider than long, 2.455 (2.260 - 2.618) wide by 2.022 (1.979 - 2.064) long. Frontal plates distinct, roughly equal in width for total span, with free ends covering all but terminal segments of first antennae as seen in dorsal view; fusion of frontal plates marked by deep incision. Lateral lobes semicircular, originating anteriorly from small nodule at base of first maxilla and continuing posteriorly to slightly underlap the lateral dorsal plate of the fused second and third thoracic segments. Cephalic area (over mouth cone) fanning out anteriorly to join frontal plates. Thoracic area (posterior to termination of lateral lobes) trapezoid in shape. Thin adhesion membrane rimming lateral and frontal lobes. First segment fused with head. Second and third thoracic segments fused, armed with dorsal plate with squared lateral margins. Fourth thoracic segment free with rounded dorsal plate (1.200 wide by 0.671 long) covering anterior margin of genital segment. Genital segment longer than wide, 1.936 (1.674 - 2.096) long by 1.537 (1.498 - 1.580) wide, tapering slightly anteriorly into

18

fourth segment and enlarging gradually posteriorly to form a central terminal lobe; abdomen completely obscured in dorsal view. Two large seminal receptacles with associated tubes easily visible in posterior third of genital segment; dark ovarian masses on either side of intestine. Abdomen triangular,1-segmented, attached to ventral surface of genital segment; anal cleft deep. Large palm-shaped caudal furcae(0.364 (0.305 - 0.429) long by 0.291 (0.244 - 0.328) wide), armed with four plumose setae and one small spine on lateral margin. Egg sacs long (1% times total length); eggs uniseriate, strongly compressed.

First antenna 3-segmented, fleshy; basal segment fused with frontal lobe and unarmed; second segment equal in width throughout length, terminated by over a dozen soft, short, naked setae, and one small plumose seta on posterior margin; terminal segment smaller than other two (% diameter of second segment), armed with about seven short, soft, setae. Second antenna apparently 4-segmented, uncinate; terminal segment sharply hooked distally and armed with two accessory spines on inner margin; third segment enlarged; first segment with adhesion pad.

First maxilla reduced to a nodular adhesion pad (see whole mount) just posterior to base of first antenna and at inception of lateral lobe.

19

Second maxilla reduced to conical swellings at base of and on either side of mouth cone; apex of maxilla armed with single stout spine.

First maxilliped 3-segmented, uncinate; first segment strongly developed, as long as segments two and three combined, and wider than either segment; segment two armed stout plumose seta on inner margin at union between segments two and three; terminal segment unarmed except for small serrations at tip.

Second maxilliped indistinctly 3-segmented, nodose; segments one and two stout, unarmed, with segment two one- half as long as segment one; segment three claw-like. Action of appendage subchelate, with flexhy disc on distal end of segment two apposing small accessory claw at base of segment three.

Four pairs of biramous thoracic legs; legs one through three with two segments per ramus; fourth leg with one segment per ramus. Armature of the legs consisting of long plumose setae, naked spines, marginal hairs, and marginal denticulations; armature according to formula as follows (see also Figures 19 to 22):

Leg 1: Exopod; 0-1, 4-I1I Endopod; 0-0, 3-0Leg 2: Exopod; 1-1, 5-IV Endopod; 1-0, 6-0Leg 3 : Exopod; 1-I, 4-IV

Endopod; 1-0, 4-0Leg 4: Exopod; 4-V

Endopod; 5-0

20

Male: Total length three-fourths that of female.Generally similar to female in antennae, mouth parts, segmentation and armament of thoracic legs, and body segmentation. Different from female as follows: 1. lateralplates on fourth segment not as greatly produced laterally;2. genital segment in male smaller proportionally and not overlapping abdomen as in female.

Discussion. The copepods of this population appear to be conspecific with Heegaardfs (1962) Nes ippus incisus despite several disagreements between my description and his. Conspecificity is established on the basis of the following similar characters: 1. measurements of total length and carapace length and width within acceptable limits; 2. antennae and mouth parts with the exception of the second maxillipeds agree in shape and basic armament; 3. coverage of the dorsal plates attached to fused second and third segments and the one attached to the fourth segment is similar.

The conflict between the descriptions is in the second maxillipeds and some of the armature of the thoracic legs. Heegaard8s drawing shows the second maxilliped of his population to be more elongate than mine and with a less distinct disc opposing the terminal segment. However,Heegaard1s appendage was drawn in situ while mine was first dissected out. If in fact the proximal end of the maxilliped is attached to the ventral surface of the head for a good part of the distance, as Heegaard states, then a dissected

21

appendage might be shorter than one in situ depending on where one disjoined the appendage. The above differences do not warrant the declaration of a new species.

The male is compared with the female for the first time for although the male was figured in the original description no mention of it was made in the text.

This description constitutes both a new host and locality record, as N. incisus was first reported only on Mustelus antarcticus taken in waters off New South Wales,Australia.

Superfamily Dichelesthioidea Yamaguti, 1963 diag. emend.

Superfamily diagnosis.-Caligidea. Females Body more or less distinctly segmented and usually cylindrical; head fused with first thoracic segment, occasionally separated; fourth segment as wide as third; genital segment elongated, usually of lesser width than rest of body, rarely enlarged as in Caligoidea. Egg sacs usually straight tubular, rarely coiled; eggs uniseriate. First antenna 2- to 15-segmented.Second antenna uncinate or chelate, commonly prehensile.Maxillary hook and furca absent. Maxilla setiferous, exceptionally uncinate. First and second maxillipeds pre­hensile; occasionally only one pair of maxillipeds present.Four pairs of legs present; all or first one or two biramous, others uniramous, rudimentary or lacking. In Anthosomatidae the legs may be fused into a single plate or represented by

22

paired wings, of which the anterior two are usually biramous.Male similar to female in general form. Parasitic on fishes.

Discussion. Yamaguti's super family is revised to in­clude the characteristics of the new family Nothadichelidae. Changes are made with respect to the elongate genital segment, the absence of one pair of maxillipeds, and the presence of coiled egg sacs. These amendments represent a broadening in the requirements for membership in the superfamily and Yamagut^s diagnosis is changed very little.

Family Eudactylinidae Yamaguti, 1963 Genus Eudactyllna van Beneden, 1853

Eudactyllna sp.(Figures 23 to 34)

Hosts and number examined: CephaloscyIlium IsabellaBonnaterre, Carpet Shark; family Scyliorhinidae.(Type host) (11).Mustelus antarcticus Guenther, Gummy Shark; family Carcharinidae. (7).Raja nasuta Muller and Henle, Skate; family Rajidae. (8). Torpedo fairchildi (Hutton), Electric Ray; family Torpedinidae. (2).

Location: Gills.Localities: 1. Akaroa (43°48'S, 172°58,E), Canterbury

Province, South Island, N. Z.; Akaroa Harbor (4-10 fathoms; mud, sand, horse mussels).

23

2. Dunedin (45°53,S, 170°31,E)> Otago Province,South Island, N. Z, :a. 2.6 miles, 090° from Taiaroa Heads (45°

47*S, 170°45,E) (17 fathoms; fine sand).b. 4.3 miles, 135° from Taiaroa Heads (45°

47*S, 170°45,E) (8 fathoms; hard sand).c. 4.3 miles, 045° from Karitane (45°34,S,

170°51'E).3. Timaru (44°24'S, 17l015*E), Canterbury

Province, South Island, N. Z.%a. 8.7 miles, 068° from Timaru (9 fathoms;

sand).b. 20.0 miles, 068° from Timaru (16 fathoms;

sand, mud).c. 45.2 miles, 068° from Timaru (28 fathoms;

sand).Number of copepods recovered; C. Isabella 53, M. antarcticus 5,

R. nasuta 22, T. fairchildi 76.Number of copepods examined; 15.

Description. Female; Body elongate, 2.295 (1.854 «2.845) long (including caudal furcae). Head covered with a carapace and fused to first thoracic segment; carapace slightly longer than wide, 0.442 (0,356 - 0,507) long by 0.419 (0.336 - 0,504) wide, narrowing anteriorly; dorsal surface smooth, slightly convex with lateral margin turning ventrally for a short distance; first antennae visible in dorsal aspect; no carapace sutures apparent. Thoracic

24

segments two through five free, covered with very small spines, lightly arched dorsal plates without lateral lobes. Length of segments as follows in decreasing order: fourth, third, fifth, second. Lamellate fifth legs easily seen in dorsal view. Genital segment (0.230 long, 0.258 wide) three-fourths the length and width of the fifth segment; egg sacs attached ventrally. Abdomen 2-segmented, narrowing posteriorly,0.222 long by 0.200 wide; anal segment half as long as first abdominal segment. Caudal furcae 0.098 long by 0.057 wide, lamellate, 1-segmented, setiferous. Egg sacs approximately five-sixths of total length, uniseriate; about 22 slightly flattened eggs per sac.

First antenna 4-segmented, strongly developed though non-prehensile; segments one and two at right angles to segments three and four. Segments armed with many spiny claws, the largest affixed to distal end of segment two; armament: X-II-V-X.

Second antenna 3-segmented, uncinate; first and third segments equal in length and almost three times longer than the second segment. Many small spines on inner margin (concave side) of segment one; one large spine on inner margin of second segment, and two small spines at base of hooked portion of third segment.

Mouth cone small, bluntly rounded, enclosing slender, slightly curved mandibles; mandibles with three tubercles on the grinding edges.

25

Maxilla biramous, with exopod shorter than endopod; exopod tipped with two spiniform setae, the shorter one- fourth as long as the other; endopod also with two setae,

t.the shorter a little more than one-half the size of the other.

First maxilliped 3-segmented; first segment as large as second, rounded and rectangular in shape, with several denticles on inner distal margin; second segment teardrop­shaped (larger end distal), with denticles lining the inner margin; third segment two-fifths as long as second segment, claw-shaped, with small denticles on inner margin.

Second maxilliped strong, 2-segmented, chelate. Segment one large, unarmed. Segment two chelate, equal in length to segment one; movable upper section with small spine on inner margin; stationary section with denticles on central surface and terminating in a ribbed canal housing tip of upper movable section.

Five pairs of thoracic legs present; first four pairs biramous with three segments per ramus; fifth pair uniramous with one segment (see whole mount, Figure 23), Surface of both rami of legs three and four and segments two and three of endo­pod two covered with denticles. Segment one of endopod two greatly enlarged, accounting for 90 per cent of the bulk of the ramus. Bases of legs two, three and four with marginal denticles. Fifth pair of legs lamellate, covered with hairs, terminating in two setae. Armament of thoracic legs consisting

26

of spines; major spine formula for first four pairs of legs as follows:

Leg 1: Exopod; Endopod; I-I-IIO-O-IILeg 2: Exopod; Endopod; 0-0-00-0-11Leg 3: Exopod; Endopod; I-I-ILI0-0-1Leg 4: Exopod; Endopod; I-I-III 0-0-1

Ma le2 Unknown.

Discussion. Comparison of the copepods of my population with the type genus, Eudactylina van Beneden, 1853, clearly indicates that they belong to the family Eudactylinidae because: 1. neither wings nor plates are attached to any ofthe body regions; 2. the first four pairs of thoracic legs are present and are equally well developed; 3. the second antenna is prehensile; and 4. two pairs of prehensile maxillipeds are present. Affinity with the genus is based on the following characters: 1. the second antenna is uncinate; 2. the genitalsegment is not appreciably elongated; 3. the rami of the first four legs each contain three segments; and 4. the second maxilliped is equipped with a stout chela.

A study of the literature at hand indicates that the copepods of my population appear to be most similar (though not conspecific) to Eudactylina rachelae Green, 1958, taken from Torpedo nobiliana captured in waters off Plymouth,England. Of the 22 known species of Eudactylina. I have been

27

able to check the descriptions and/or figures of 18 and have eliminated them all from conspecificity. However, the des­criptions of four species (E. carchariae-glauci Hesse, 1883, li* corrugata Bere, 1930, E. musteli-laevis Hesse, 1883, and E. valei Nunes-Ruivo. 1956) are not available to me and the determination of whether my population is a new species must await the acquisition of necessary literature. None of the outstanding four species were taken from the same host species as the copepods of my population nor do the localities of the four approach closer than 9000 nautical miles to New Zealand.

Genus Kr^yeria van Beneden, 1853

Kr^yeria musteli nov. sp.(Figures 35 to 44)

Hosts and number examined: Galeorhinus australis Macleay,Southern Tope or School Shark; family Carcharinidae. (9).Mustelus antarcticus Guenther, Gummy Shark; family Carcharinidae. (Type host) (24).

Location: Gills.Localities: 1. Akaroa (43°48*S, 172°58IE), Canterbury

Province, South Island, N.Z.; Akaroa Harbor (4-10 fathoms; mud, sand, horse mussels).

2. Timaru (44°24'S, 171°15»E), Canterbury Province, South Island, N. Z.;a. 8.7 miles, 068° from Timaru (9 fathoms;

sand).

28

b. 20.0 miles, 068° from Timaru (16 fathoms; sand, mud).

Number of copepods recovered: G. australis 27, M. antarcticus 27. Number of copepods examined: 10.

Description. Female: Elongated, narrow, 5.51 long (measurement exclusive of caudal furcae). Head fused with first segment, covered with a carapace; carapace wider than long, 0.60 wide by 0.58 long, rounded laterally and posteriorly, tapering anteriorly to form a slight lobe. Movable styliform process articulating with posterior skeletal support of carapace, reaching to posterior margin of second thoracic segment; tips of processes sharp, slightly curved. Thoracic segments two, three, and four free, each with an incon­spicuous dorsal plate without lateral lobes; segments two and three of equal length, segment four one-half times longer than segment three. Fifth and genital segments fused, 3.72 long by 0.32 wide. Abdomen narrow, 0.58 long by 0.14 wide,1-segmented, 2-pseudosegmented; moderately deep anal cleft.Caudal furcae cylindrical, 0.22 long, with inner margins fringed with minute hairs; furcae terminating in three plumose setae, the longest seta situated medially. Egg sacs one-half total length of body*uniseriate; approximately 18 slightly flattened eggs per sac.

First antenna 6-segmented, non-prehensile, tapering distally, armed with naked setae; setation: 7-5-3-1-0-9.

29

Second antenna strongly chelate, apparently 2-segmented; shorter movable portion of chela resting in a deep channel in terminus of larger stationary member.

Rounded mouth cone enclosing slender, slightly curving mandibles bearing six major serrations on grinding edges.

Maxilla not observed in the many specimens prepared though standard bipartite structure assumed to be present

First maxilliped 3-segmented; second segment enlarged, covered with fringing hairs on distal inner margin and a pad on distal outer margin bearing denticles. Third segment smaller than second, in form of a stout hook with twb terminal setae.

Second maxilliped 2-segmented, both segments of approximately equal length; first segment strong, second segment slender with sharp hook distally.

Four pairs of well developed biramous thoracic legs with three segments per ramus. Ramus armature consisting of long plumose setae and short, weakly developed spines.Single outwardly directed spines on bases of legs two and four. Setation formula as followss

Leg Is Exopod; 1-0, 1-0, 4-IIEndopod; 1-0, 0-0, 6-0Leg 2s Exopod; 1-1, 1-0, 5-IIEndopod; 1-0, 0-0, 6-0Leg 3s Exopod; 1-0, 1-0, 5-1Endopod; 1-0, 0-0, 4-0

30

Leg 4: Exopod; 1-1, 1-1, 5-IIEndopod; 1-0, 1-0, 3-0

Male: Unknown.

Discussion: Kr^yeria mustell nov. sp. is most closelyrelated to both K. lineata as described by van Beneden (1853), and K. sublineata as described by Yamaguti and Yamasu (1959), and may very well form the third member of a subspecies triad.

rousteli resembles K. lineata in body length, structure of the second antenna, mandible, first maxilliped and second maxilliped. It resembles K. sublineata in thoracic leg armature (it appears that Yamaguti and Yamasu have counted spines as plumose setae), and in the structure of the abdomen.

musteli differs from both K. lineata and K. sublineata in the lesser number of segments and armature of the first antenna, in the shorter extension of the styliform process (in K. lineata and K. sublineata the process reaches to the fourth thoracic segment while in K. musteli it reaches only to the posterior margin of the second thoracic segment), and in the lesser number of eggs and relatively longer egg sac.

This species takes its trivial name from the generic name of the type host.

Family Nothadichelidae fam, nov.

Family diagnosis.-Caligidea, Dichelesthioidea. Female:Head fused with first thoracic segment. Thoracic segments two and three more or less free; remaining segments fused with

31

genital segment. Genital segment elongate and about as wide as carapace. Abdomen 1-segmented, short, attached either ventrally or dorsally to genital segment. Caudal rami small, longer than wide, consisting of two segments. Egg sacs long, straight or coiled; eggs moderately flattened. First antenna 4- to 3-segmented, setiferous, non-prehensile. Second antenna 2- to 3-segmented, uncinate, strongly developed.Mouth tube conical, rounded. Mandibles styliform, unarmed. Maxilla biramous with bifid exopod and smaller endopod, when present, consisting of nodular base armed with a spine. One pair of 4-segmented maxillipeds present; fourth segment terminating in a bifid claw. Four pairs of biramous thoracic legs present; fifth pair lacking; first three pairs multi­jointed with two or three segments per ramus; fourth pair reduced or rudimentary with one or two segments per ramus.Bases of all legs (though much reduced in fourth pair) with lateral lobes, either smooth or spinate.

Type genus: Nothadichelidium gen. nov.

Discussion. The erection of Nothadichelidae fam. nov. is necessary to accommodate two new genera which do not fit existing families. Addition of a new family rather than the emendation of an existing one was necessitated by the lack of general agreement with any one family.

The new genera do not fit the diagnosis of Anthosomatidae Yamaguti, 1963 for the following reasons: 1. body withoutplates and carapace processes as in Anthosomatidae; 2* lack

32

of a second pair of maxillipeds as is present in Anthosomatidae3. third and fourth legs not fused into a plate; 4. abdomen not covered by a dorsal plate. Exclusion from the family Catlaphilidae Ttipathi, 1960 is obvious because in the new genera: 1. the head is fused with the first segment and the fifth segment is not free; 2. the second antenna is 3- segmented and prehensile; 3. the fifth thoracic leg is absent4. the caudal furcae are not rudimentary. The new genera do not belong in Dichelesthidae Bassett-Smith, 1899 because they:1. possess no lobes on the thoracic segments; 2. have only one pair of maxillipeds; 3. have biramous third thoracic legs. The new genera are not compatible with the diagnosis for Eudactylinidae Yamaguti, 1963 due to the lack of two pairs of maxillipeds and the presence of a bifurcate maxilla. Affiliation with the family Pseudocycnidae Yamaguti, 1963 is ruled out as the new genera do not possess two pairs of maxillipeds but have four pairs of biramous thoracic appendages.

Basically, the characters that distinguish members of this family from all others in the superfamily are the lack of two pairs of maxillipeds and the presence of four pairs of biramous thoracic legs. Although every other family description included mention of at least one free thoracic segment, the author did not observe a truly free segment in the specimens of the new family. Manipulation and careful observation re­vealed no articulation or visible suture lines despite the flexibility of the area between segments.

33

The name of the new family, Nothadichelidae (L), alludes to its maverick nature in the superfamily*

Genus Nothadichelidium gen. nov.

Generic diagnosis.-Caligidea, Dichelesthioidea, Notha­dichelidae. Female; Body elongate. Carapace wider than long; posterior margin not overlapping first thoracic segment. Thorax with lateral processes. First segment fused with head; second and third segments more or less free; remaining segments fused with genital segment. Genital segment con­stituting over one-half total body length yet not appreciably wider than carapace; posterior margin truncate with slight lobes. Abdomen small, 1-segmented, attached to ventral surface of genital segment. Caudal rami small, narrow, 2-segmented. Egg sacs long, coiled; eggs flattened. First antenna 5-segmented, non-prehensile. Second antenna 3-segmented, uncinate; main organ of prehension. Mouth cone rounded, directed posteriorly; mandibles styliform, slightly curving. Maxilla biramous with bifid exopod and spinate, nodular endopod. Maxilliped (one pair only) 4-segmented, terminating in a bifid claw; weakly prehensile. Four pairs of biramous thoracic legs present with the following segmentation; first leg, two segments per ramus; second leg, 3-segmented exopod,2-segmented endopod; third leg, two segments per ramus; fourth leg, reduced, 2-segmented exopod, 1-segmented endopod. All legs armed with spines; bases with spinate lateral lobes.

34

Type species: Nothadichelidium polyprionum n. sp.

Nothadichelidium po lyprionum gen, et sp. nov,(Figures 45 to 54)

Hosts and number examined: Polyprionum oxygenios (Bloch and Schn.), Groper; family Serranidae, (Type host) (8). Leptocephalus verreauxi (Kaup), Conger Eel; family Leptocephalidae. (6),

Location: Gills,Localities: 1. Akaroa (43°48fS, 172°58,E), Canterbury Province,

South Island, N. Z.; Akaroa Harbor (4-10 fathoms; mud, sand, horse mussels),

2. Cape Campbell (41°43'S, 174°17IE), Marlborough Province, South Island, N. Z.; 6.9 miles,158° from Cape Campbell (60 fathoms; soft mud),

3. Timaru (44°24«S, 171°15«E), Canterbury Province, South Island, N. Z.; 8.7 miles, 068° from Timaru (9 fathoms; sand).

Number of copepods recovered: P. oxygenios 13, L. verreauxi 16.Number of copepods examined: 10

Description. Female: Body elongate without dorsal plates, 7.22 long by 1.05 wide at its widest point. Head fused with first segment, wider than long (0.83 wide by 0.62 long); covered dorsally with rectangular carapace which does not overlap the first thoracic segment. Head followed by narrowed thorax consisting of five segments, four (segments

35

2 through 5) defined by lateral lobes. Segment one fused with head, segments two and three more or less free (no definite articulation points observed), and segments four and five fused with genital segment. Spinate lobes belonging to the basal structure of legs one and two seen in dorsal view behind head and between segments two and three. Genital segment large, accounting for approximately three-fifths of total body length; ovaries appearing as darkened areas on either side of intestine. Posterior end of genital segment truncated with slight lobes on either side of abdomen. Small (0.39 long by 0.30 wide),1-segmented abdomen attached to ventral surface of genital segment. Caudal rami small, narrow, 0.27 long by 0.07 wide,2-segmented, with- short spine on terminal segment. Egg sacs (0.24 wide) long, coiled; eggs flattened.

First antenna 5-segmented, weak, non-prehensile; segments increasing in length and decreasing in width from proximal to distal end. Naked setae on anterior margins of all segments plus a tuft of seven or more at terminus of segment five. First antenna easily seen in dorsal view of whole mount.

Second antenna 3-segmented, uncinate, strong; apparently the main organ of prehension.

Mouth cone rounded and bent posteriorly, enclosing styliform mandibles. Mandibles unarmed, curving medially to form smooth hooks.

36

Maxilla biramous; exopod delicate, bifid, with base lateral to and at level of base of mouth cone. Endopod small, consisting of a spine on a nodular base anterior to base of maxillipeds.

Maxilliped 4-segmented, directed posteriorly, with segment four a bifid claw. (Above appendages arise at level of terminus of mouth cone and from their weak structure seem more suited for assisting the mandibles in tearing flesh rather than for prehension.)

First leg biramous with two segments per ramus; armed with spines and fringing bristles (Figure 51). Bases with large lateral lobes armed with small spines.

Second leg biramous, with an exopod of three segments and endopod of two segments. Armament consisting of spines and a small denticulated area on segment three of the endopod (Figure 52). Bases with similar spiniferous lobes as in leg one, only smaller.

Third leg biramous, with two segments per ramus; armed as in previous appendages (Figure 53). Basal lobe smaller than in leg two.

Fourth leg biramous, greatly reduced with exopod of two segments and endopod of one segment (Figure 54). Basal lobe present but very small.

Male: Unknown.

37

Discussion, Species takes its trivial name from generic name of type host.

Genus Neonothadiche1idium gen, nov.

Generic diagnosis.-Caligidea, Dichelesthioidea, Nothadichelidae. Female: Body elongate. Carapace wider than long; posterior margin overlapping all or most of first thoracic segment. Thorax with lateral processes. First segment fused with head; second and third segments more or less free (no definite articulation points observed); remaining segments fused with genital segment. Genital segment length constituting over one-half total body length yet not appreciably wider than carapace; posterior margin rounded. Abdomen small, t-segmented, attached to dorsal surfaceof genital segment. Caudal rami small, narrow, 2-segmented.

(

Egg sacs long, straight; eggs flattened. First antenna 4- segmented, non-prehensile. Second antenna 2-segmented, uncinate, with peg-like process in segment two which fits into a small hole in segment one; main organ of prehension. Mouth cone rounded, directed posteriorly; mandibles styliform, slightly curving. Maxilla biramous with bifid exopod and unarmed nodular endopod. Maxilliped (one pair only) 4-segmented, terminating in bifid claw; weakly prehensile. Four pairs of biramous thoracic legs present with following segmentations first leg, two segments per ramus; second leg, 2-segmented exopod, 3-segmented endopod; third leg, 2-segmented exopod,3-segmented endopod; fourth leg, modified, fleshy, with one

38

segment per rarmis. All legs armed with spines or tubercles.Bases with smooth lateral lobes (lobe of fourth leg less dis­tinguishable from rest of leg structure).

Type species: Neonothadichelidium leptocephali gen, etsp. nov.

Discussion. Neonothadichelidium gen. nov, differs from Nothadichelidium gen. nov., the type genus of the new family Nothadichelidae, in the following characters: 1. carapacenot only covers the head but also covers most of the first thoracic segment; 2. first antenna is 4-segmented vice 5- segmented; 3. thoracic leg segmentation is different in the second, third, and fourth legs; second and third have 2- segmented exopods with 3-segmented endopods; fourth with both rami 1-segmented; fourth legs are somewhat rudimentary;4. egg sacs are straight rather than coiled; 5. abdomen is attached to the dorsal surface of the genital segment instead of the ventral surface as in No thad iche1id ium.

At this taxonomic level the most important characters that can be used to separate the two new genera would seem to be the following as they are found in Neonothadichelidium: the extension of the carapace over a good portion of the first thoracic segment, the straight egg sacs, and the dorsally attached abdomen.

39

Neono thad ichelid ium leptocephali gen, et sp, nov.(Figures 55 to 63)

Hosts and number examined: Polypr ionum oxygenios (Bloch and Schn.), Groper; family Serranidae. (8),Leptocephalus verreauxi (Kaup), Conger Eel; family Leptocephalidae. (Type host) (6).

Lo cat ion: Gills.Localities: 1. Akaroa (43°48'S, 172058'E), Canterbury

Province, South Island, N. Z.; Akaroa Harbor (4-10 fathoms; mud, sand, horse mussels).

2. Cape Campbell (41°43,S, 174°17*E), Marlborough Province, South Island, N. Z.; 6.9 miles,158° from Cape Campbell (60 fathoms; soft mud).

3. Timaru (44°24*S, 171°15,E), Canterbury Province, South Island, N. Z.; 8.7 miles,068° from Timaru (9 fathoms; sand).

Number of copepods recovered: P. oxygenios 11, L. verreauxi 10. Number of copepods examined: 10.

Description. Female; Body elongate without dorsal plates, 3.75 long by 0.82 wide at its widest point. Head fused with first segment, wider than long (0.70 wide by 0.55 long); covered dorsally with oval carapace which overlaps the greater part of the first thoracic segment. Thorax produced laterally at two points, delineating segments two and three which are more or less free and equal in width. Remaining segments fused to genital segment. Smooth lobes, belonging

40

to the basal structure of legs one, two, and three, seen in dorsal view behind the head, lateral to segment two, and behind segment three, respectively. Genital segment large, accounting for two-thirds of total body length when measured with fused fourth and fifth segments; ovaries filling entire genital segment on either side of intestine. Posterior margin of genital segment rounded. Abdomen small (0.27 wide by 0.26 long), 1-segmented, attached to dorsal surface of genital segment. Caudal rami small, narrow, 0.20 long by 0.06 wide, 2-segmented, with short spine on terminal segment. Egg sacs (0.24 wide) long (two times total body length), straight; eggs flattened.

First antenna 4-segmented, weak, non-prehensile; segments tapering distally with segment two being longest.Two naked setae on anterior margin of segment one and tuft of setae on terminus of segment four. First antenna easily seen in dorsal aspect.

Second antenna 2-segmented, uncinate, strong, with peg-like process in segment two which fits into a hole (or socket) in segment one; this main organ of prehension usually visible in dorsal view.

Mouth cone rounded, directed posteriorly, and enclosing inwardly hooking styliform mandibles; mandibles unarmed.

Maxilla biramous; exopod delicate, bifid, originating laterally to base of mouth cone. Rounded, unarmed nodule

41

situated anterior to base of maxilliped, representing only remnant of endopod.

Maxilliped 4-segmented, arising at end of mouth cone, directed posteriorly; segment four in the shape of a bifid claw. (Above appendages apparently have little to do with prehension due to their weak development and therefore probably assist the mandibles in rending host tissue.)

First leg biramous, with two segments per ramus; armed with stout spines (Figure 60). Bases with large, lateral, unarmed lobes«

Second leg biramous, with exopod of two segments and endopod of three segments. Armament consisting of shorter, even more strongly developed spines than in leg one; basal lobes also present, slightly larger than leg one (Figure 61).

Third leg biramous, with exopod of two segments and endopod of three segments. Armed with stout spines, three of which have been reduced to tubercles (Figure 62). Basal lobe present as in previous legs but somewhat smaller.

Fourth leg biramous, with one segment per ramus.Entire leg structure, including basal process, modified into a large basal area and two fleshy lappets, each of which terminates in weak spines (Figure 63).

Male s Unknown»

42

Discussion. This species takes its trivial name from the generic name of the type host.

Order Lerneopodidea Yamaguti, 1963 Family Chondracanthidae Dana, 1853

Subfamily Chondracanthinae Dana, 1853Genus Chondracanthus De la Roche, 1811

Ghondracanthus genypteri Thomson, 1890 (Figures 64 to 79)

Host and number examined: Genypterus blacodes (Bloch and Schn.),Ling; family Ophidiidae. (4).

Location: Gills.Localities: ' 1. Dunedin (45°53,S, 170°31,E), Otago Province,

South Island, N. Z.; 4.3 miles, 045° fromKaritane (45°34'S, 170°51,E).

2. Timaru (44°24,S, 17l°15,E), CanterburyProvince, South Island, N. Z.; 8.7 miles,068° from Timaru (9 fathomsJ sand).

Number of copepods recovered: 73.Number of copepods examined: 15.

Description. Female: Body elongate, 5.79 (4.89 - 7.20) long by 2.13 (2.06 - 2.25) wide at widest point. Head oblongin shape, fused with thorax though a slight constrictiondelineates head from thorax. First two thoracic segments fused^ with laterally and posteriorly directed processes at point of fusion; first and second thoracic legs attached ventrally.

43

Rest of body, consisting of remaining thoracic segments fused with genital segment, constricted in center, forming two portions of approximately equal area as in the body of a violin; posterior margin of both anterior and posterior portions with lappets. Abdomen small (0.48 long by 0.47 wide),1-segmented, situated between terminal set of lapetts. No caudal rami. Egg sacs three-fourths length of body; numerous small eggs (0.21 diameter).

First antenna, 1-segmented, fleshy, armed with approximately seven naked setae at terminus; easily dis- cernable from dorsal aspect of whole mount.

Second antenna 1-segmented, uncinate, strongly developed; main organ of prehension. (These appendages are brittle and tend to break off and remain in the host's tissue if care is not exercised when removing them. )

Mandible apparently 2-segmented; terminal segment falciform with serrations lining the entire medial or grinding edge and ventral stir face.

Maxilla, located between mandible and first maxilliped, consisting of a nodular base with two stout spines directed posteriorly.

First maxilliped indistinctly 2-segmented; basal segment strong; terminal segment tapering to dull point, with most of medial edge armed with denticles.

44

Second maxilliped indistinctly 3-segmented, strongly developed with claw-like tip; prehensile.

First and second thoracic legs (Figure 65) biramous, reduced to 1-segmented fleshy lobes attached to soft bases. Remnant of third leg reduced to soft palp attached to anterior margin of fused thorax-genital segment,

Male: Small, 0.45 long by 0.29 wide, attached to female at fusion of abdomen and genital segment. Head fused with first thoracic segment, larger than rest of body. Second thoracic segment more or less free. Trunk, consisting of remaining thoracic segments, genital segment, and abdomen, indistinctly 4-segmented, directed ventrally. Caudal rami1-segmented, well developed, with basal setae.

First antenna soft, as in female, with approximately six naked setae at terminus; located directly behind second antenna as seen in lateral view.

Second antenna 1-segmented, strong, uncinate. Very prominent from all aspects; main organ of prehension.

Mandible 2-segmented, with falciform terminal segment bearing serrations both on cutting edge and ventral surfaces as in the female counterpart.

Maxilla with nodular base armed with two spiniformsetae.

45

First maxilliped 2-segmented; basal segment strong with spine on anterior margin; terminal segment one-half length of basal segment with two spines on margin.

Second maxilliped 2-segmented; basal segment swollen; terminal segment broad-based, tapering to claw-like point.

First and second thoracic legs reduced to 1-segmented lamelliform processes with two terminal setae.

Antennae and mouth parts of the male are conspicuously similar to those of the female, only smaller.

Discussion. This redescription of Chondracanthus genypterl is presented because Thomson's (1890a) description of this species is short and lacks the detail now necessary to separate the species of this rather large genus. The mouth parts, the structure of the female's abdomen, and the attached male omitted by Thomson are described for the first time.

Family Lerneopodiae Olsson, 1869 Subfamily Clavellinae Dana, 1853

Genus Clave1lodes Wilson, 1915

Clavenodes intermedius (Quidor, 1906) Wilson, 1915(Figures 80 to 85)

Syn: Anchorella intermedia Quidor, 1906

46

Hosts and number examined: Trematomus bernacchil Boulenger; family Nototheniidae. (Type host) (19).T. borchgrevinki Boulenger; family Nototheniidae. (9).2- hansoni Boulenger; family Nototheniidae. (l) Rhigophila dearborni DeWitt; family Zoarcidae. (2).

Location: Gills.Localities: 1. McMurdo Sound area (77°50,S, 166°45,E),

Antarctica.2* Windmill Island group near Wilkes Station

(66°15 *S, 110°32 *E), Antarctica.

Latitude StLongitude Depth in fathoms Bottomtypea. 66° 15 8 S, 110°35 0 E 7 Sand, rockb. 66°15 9 s, 110°34 4 E 10 Rockc. 66° 15 9 s, 110°34 3 E 12 Rockd. 66°15 9 s, 110°35 1 E 9 Rocke. 66° 15 9 s, 110°31 2 E 30-50 Rockf. 66° 16 0 s, 110°32 3 E 20 Rocks. 66° 16 0 s, 110°31 0 E 30-50 Rockh. 66°2l!0 s, 110°27 0 E 200 Mud

Previously reported host and locality: Nothostenia sp. fromAntarctica (from Jean Charcot expedition) (Quidor, 1906).

Number of copepods recovered: T. bernacchii 20, T. bor chgrevinki16, T. hansoni 1, R. dearborni 2.

Number of copepods examined: 15.

Description. Female: Body consisting of three regions: head, neck, and trunk. Head, averaging 1.2 long, well

47

delineated from neck by a dorsal cephalic shield; all mouth parts except first maxillipeds contained under shield. Neck, about 4.5 long, smooth, slightly increasing in diameter from anterior end to union with trunk where the displaced first maxillipeds lie; neck generally reflexed against dorsal side of trunk. Trunk wider than long (4.1 wide by 3.9 long), plump though depressed, slightly tapering anteriorly, truncated posteriorly; ventral surface with genital processes bearing four genital lappets (two dark seminal receptacles can be seen in genital process); round reproductive structures on either side of process. No abdomen or caudal rami. Egg sacs cylindrical, long (6.2 long by 1.9 wide); eggs multiseriate.

First antenna slender, 3-segmented, originating behind base of second antenna; terminal segment with four soft spines.

Second antenna large, biramous, indistinctly 3-segmentedt originating posterior and lateral to origin of maxilla; exopod2-segmented, endopod 1-segmented; rami terminating in denticles.

Mouth cone oval with fine hairs around entire margin of orifice. Mandibles styliform with slight terminal knobs.

Maxilla at base of mandibles, tripartite; palp with single terminal seta.

Second maxilliped subchelate, 2-segmented; terminal claw with accessory claw and row of short teeth on inner

48

margin near tip; basal segment with small peg-like structure on inner margin*

First maxilliped located at base of neck, indistinctly segmented* (These appendages are so short that the bulla appears to be attached directly to the neck.)

No thoracic legs present on adult.

Male: Body almost square, 0.9 long by 0.8 wide. Headwith dorsal carapace, folded to lie on ventral side of trunk; head and trunk fused with no demarcation between them.Anterior end of body truncated and with mouth parts.Appendages and genital process pointing forward.

First antenna indistinctly 3-segmented, very small; seen in lateral view as slight projection at anterior margin.

Second antenna biramous as in female; dorsal ramus blunt with slight denticulation; ventral ramus 2-segmented with larger denticles on terminal segment.

Maxilla as in female but smaller.

Both pairs of maxillipeds strongly developed, uncinate; first pair somewhat larger and more heavily armed with denticles on terminal segment.

Male attached to female on or near to genital process.

49

Discussion. The specimens of my population fit the generic diagnosis of Clave1lodes as found in Wilson (1915) and Yamaguti (1963). The present study reveals that the copepods in this collection are probably conspecific with Clavellodes intermedius (Quidor, 1906) Wilson, 1915, though the original description is sketchy (Wilson did not redescribe the species when he removed it from Anchorella (Cuvier, 1830) and placed it in Clavellodes). Most of Quidor1s description is devoted to comparisons between the male and female of the species rather than to a clear description of either sex to properly describe the animal. The comparison of my population with that represented by Quidor (1906) is based on measurements, shape of the maxilla, shape and armature of the mouth cone, the genital processes and seminal receptacles, and the similarity in hosts and localities. Further, C. intermedius is the only species of that genus found in Antarctic waters or on the host genera.

The validity of Rhigophila dearborni as a host ofClavellodes intermedius must be questioned. It is believedpossible that a mixup in labeling and host records might haveoccurred. This decision is based on host-parasite data utilizingboth monogenetic trematodes and parasitic copepod records.The ectoparasitic fauna of the two fish specimens in questionare, from past experience, typically trematomid, not zoarcidin nature. This discrepancy is being checked in every waypossible, however, to date nothing tangible has been found to eliminate R. dearborni as a host of C. intermedius.

50

Family Naobranchiidae Yamaguti, 1939 Genus Naobranchia Hesse, 1863

Naobranchia helicolena sp. nov.(Figures 86 to 91)

Host and number examined: Helicolenus percoides (Richardson),Sea Perch; family Scorpaenidae. (Type host) (17),

Location: Gills.Locality: Cape Campbell (41°43IS, 174°17,E), Marlborough

Province, South Island, N. 2.; 6.9 miles, 158° from Cape Campbell (60 fathoms; soft mud).

Number of copepods recovered: 19.Number of copepods examined: 10.

Description. Female: Body divided into three parts, head, neck, and trunk. Head, 3.9 long, well separated from neck and covered dorsally and laterally with a carapace; prominent spoon-shaped palps attached to ventral surface of head. Neck 1.75 long, smooth, increasing in diameter posteriorly to join with trunk. Trunk as long as head and neck together, tapered anteriorly, expanding posteriorly to enclose egg sacs in a transparent capsule; egg sacs situated posteriorly withrespect to trunk and not laterally as in some species. Eggslarge, multiseriate, and rather loosely contained. Abdomen 1-segmented, barely visible, fused for most of its length with the genital area of the trunk; no caudal rami visible.

51

First antenna short, 5-segmented, fourth segment with sharp spine, fifth segment tapering to a point; no other armament.

Second antenna biramous, with stout basal segment; ventral ramus 2-segmented, terminal segment ending in three small claws; dorsal ramus 1-segmented, rounded terminally with small knob on end. Second antenna protruding a relatively short distance from the side of the head, not encircling the anterior margin as in some other lerneopodids.

Mouth cone directed anteriorly, rounded, enclosing straight, unarmed mandibles.

Maxilla bipartite with sharply pointed tips; originating at base of mouth cone, with the longer branch extending the full length of the cone.

Second maxilliped, 3-segmented, uncinate; first segment broad, unarmed; second segment of less diameter, tapered anteriorly to form socket to receive base of third segment; third segment claw-shaped, three-quarters as long as second segment, always found tightly flexed against the medial margin of segment two.

First maxillipeds in form of two large loops of rmiscularized tissue originating from base of neck at fusion of neck and trunk. First maxillipeds wrapped around the hostfs gill filaments.

52

No thoracic legs present.

Male: Small 0.3 long by 0.4 wide, attached to female^ genital area between egg sacs. Cephalothorax and anterior trunk fused in same line without any trace of segmentation, both covered with a dorsal carapace giving a humpbacked appearance to the copepod. Posterior trunk turned ventrally at right angle to anterior portion, appearing as a large process beneath the first maxillipeds.

Antennae and mouth parts as in female only smaller, with the exception of the first maxillipeds whose terminal claws interlock with conical protuberances.

No thoracic legs present.

Discussion. The copepods of my population clearly belong to the family Naobranchiidae and thus to the genus Naobranchia (as the family as monotypic) by virtue of the capsule enclosing together the genital portion of the trunk, the egg sacs, and the abdomen. No other species of Naobranchia possesses all of the following characters considered taxonomically important: 1. head and neck together equal inlength to the trunk; 2. first maxillipeds large, almost as long as the length of the neck region; 3. egg sacs situated at posterior margin of trunk and not extending laterally or anteriorly; 4. no elongated caudal rami.

53

N. occidentalis (Wilson, 1915) is most similar to N. helicolena however, it is eliminated from conspecificity on the basis of the structure of the second antenna, the extent of the first maxillipeds, and in the degree of capsule enclosure.

The new species takes its trivial name from the generic name of the type host.

SUMMARY

The techniques for collecting and preparing parasitic copepods for study are described. , A systematic treatment of seven copepods from New Zealand waters and two copepods from the Antarctic area is presented.

Five species are described for the first times Kr^yeria musteli sp. nov., Nothadichelidium polyprionum gen., et sp. nov. , Neono thad i che1id ium leptocephali gen. et sp. nov. , Naobranchia helicolena sp. nov. from New Zealand and Neo s cute11id ium yeatmani gen. et sp. nov. , from Antarctica. Three copepods are redescribeds Nesippus incisus Heegaard,1962, Chondracanthus genypteri Thomson, 1890, from New Zealand

\

and Clavellodes intermedius (Quidor, 1906) Wilson, 1915 from Antarctica. Eudactylina sp. from New Zealand is described but a decision whether it is a new species is withheld pending further investigation of the literature.

New host records are reported for Nesippus incisus Heegaard, 1962 and Clavellodes intermedius (Quidor, 1906) Wilson, 1915. New locality records are also established for the two species above.

\

A tisbid copepod (NeoscuteHidium yeatmani) is reported from a fish for the first time.

54

LITERATURE CITED

Bassett-Smith, P. W. 1898a. Further new parasitic copepodsfound on fish in the Indotropic region. Ann. Mag. Nat. Hist. Ser. 7, 8(2):77-98.

............ . 1898b. Some new or rare parasiticcopepods found on fish in the Indotropic region. Ibid. Ser. 7, ll(2):357-372. _____ . 1899. A systematic description of

parasitic Copepoda found on fishes, with an enumeration of the known species. Proc. Zool. Soc. London, 2s 438-507.

Beneden, P. J. van. 1853. Notice sur un genre nouveau de la tribu des Caligiens (genre Kr^yeria van Bened.).Bull. Acad. Roy. Belg. 20(1)I 23-30.

Brady, G. S. 1910. Die marinen Copepoden: 1. Ueber die Copepoden der Stamme Harpacticoida, Cyclopoida, Notodelphoida und Caligoida. Trans1. from English by Vanhoffen. Deutsch. Sudpolar-Expedition 1901-1903 (Drygalski) v. 11 Zool. 3(5)s 497-593.

DeWitt, H. H. and J. C. Tyler. 1960. Fishes of the Stanford Antarctic Biological Research Program, 1958-1959. Stanford Ichthyoi. Bull. 162-199.

55

56

Dillon, W. A. and W. J. Hargis, Jr. 1965. Monogenetictrematodes from the southern Pacific Ocean. Part I. Monopisthocotyleids from New Zealand fishes, p. 229- 249... In Biology of the Antarctic Seas II, Antarctic Research Series 5. American Geophysical Union.

Gooding, R. U, 1957. On some Copepoda from Plymouth, mainly associated with invertebrates, including three new species. J. Mar. Biol. Assoc. U. K., 3j>(2): 195-221.

_____________ . 1960. North and South American copepods ofthe genus Hemicvclops (Cyclopoida: Clausidiidae).Proc. U. S. Nat. Mus. 112(3434): 159-&95.

Graham, D. H. 1956. A treasury of the New Zealand fishes.A. H. & A. W. Reed, Wellington. 424 p.

Hargis, W. J. Jr. 1953. Chloretone as a trematode relaxer,and its use in mass-collecting techniques. J. Parasito1. 39(2): 224-225.

Heegaard, P. 1940. Some new parasitic copepods (Ghondra-canthidae and Lernaeopodidae) from western Australia. Vidensk. Medd. fra Dansk. Naturh. Foren. 104: 87-101.

___________. 1943. Parasitic copepods mainly from tropicaland antarctic seas. Ark. Zool. 34A(l8)s 37pp.

. 1962. Parasitic Copepoda from Australian waters.Rec. Aust. Miis. 25(9): 149^234.

Hewitt, G. C. 1963. Some New Zealand parasitic Copepoda of thefamily Caligidae, Tr. Roy. Soc. New Zealand 4(3): 61-115.

57

Hewitt, G. C. 1964a. The occurrence of Lepeophtheirus insignis Wilson (Copepoda parasitica) in New Zealand waters and its relationship to L. molae Heegaard. Tr. Roy. Soc.New Zealand 4(8): 153-155.

. 1964b. A new species of Lophoura (Shyriidae, Copepoda) from New Zealand waters. Tr. Roy. Soc. New Zealand 5(3): 55-58.

___________. 1964c. A new species of Caligus (Copepoda) ona species of Tripterygion from New Zealand. Tr. Roy.Soc. New Zealand 5(10): 123-130.

. 1964d. A redescription of Gloiopotes huttoni (Thomson, 1889), with a key to the species of the genus. Tr. Roy. Soc. New Zealand 5(8): 85-96.

Hutton, F. W. 1904. Index Faunae Novae Zealandiae.Philosophical Institute of Canterbury, New Zealand,Dulau Sc Co., London. 274 pp.

Lewis, A. G. 1964a. Caligoid copepods (Crustacea) of theHawaiian Islands: parasitic on fishes of the family Acanthuridae. Proc. U. S. Natl. Mus. 115(3482):137-244.

___________. 1964b. The caligid copepod genus Dentigryps(Crustacea: Caligoida). Proc. U. S. Natl. Mus. 115(3487): 347-380.

Parrott, A. W. 1957. Sea angler's fishes of New Zealand.Hodder and Stoughton, London. 176 pp.

. 1958. Big game fishes and sharks of New Zealand.Hodder and Stoughton, London. 127 pp.

58

Phillips, W. J. 1922. On the occurrence of a parasiticcopepod on the New Zealand ling. New Zealand J. Sci. Tech. 4(6): 315.

_____________ . 1927. Bibliography of New Zealand fishes.New Zealand Marine Dept. Fish. Bull. No. 1.

Quidor, A. 1906. Sur les Copepodes recueillis par la mission Charcot dans les mers Antarctiques. Bull. Mus, Nat.

Hist., Paris, 12: 27-33.,. 1913. Copepodes parasites. 2. Exped. Antarct.

Franc. (1908-1910) (Charcot). Sci. Nat. 197-215.Shiino, S. M. 1952. Copepods parasitic on Japanese fishes,

1: On the species of Caligus and Lepeophtheirus. Rep. Fac. Fish., Pref. Univ. Mie, .1(2): 79-112.

Thomson, G. M. 1890a. Parasitic Copepoda of New Zealand with descriptions of new species. Tr. New Zealand Inst,22: 353-376.

• 1890b. A new parasitic copepod. Tr. New ZealandInst. 23: 227-229.

Vervoort, W. 1964. Free-living Copepoda from Ifaluk Atollin the Carolina Islands. U. S. Natl. Mus. Bull. 236, 431 pp.

Waite, E. R. 1923. The fishes of South Australia. R. E. E« Roger, Government Printer, Adelaide. 243 pp.

Wilson, C, B. 1915. North American parasitic copepods belonging to the family Lernaeopodidae, with a revision of the entire family. Proc. U, S. Natl. Mus. 47(2063): 565-729.

59

Wilson, C. B. 1922. North American parasitic copepods belong­ing to the family Dichelesthiidae. Ibid. 60(2400): 1-100.

____________ • 1932. The copepods of the Woods Hole region,Massachusetts. U. S. Natl. Mus. Bull. 158. 635 pp.

Yamaguti, S. 1963. Parasitic Copepoda and Branchiura offishes. Interscience Publishers, New York. 1104 pp.

Yamaguti, S. and T. Yamasu. 1959. Parasitic copepods fromfishes of Japan, with descriptions of 26 new species and remarks on two known species. Biol. J. Okayama Univ. 5(3-4): 89-165.

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PLATES I-VII

(All appendages of female unless otherwise noted)

EXPLANATION OF PLATE I

Neoscute1lidium yeatmani gen* et sp. nov.Figures

I. Dorsal view of female2. First antenna.3* Second antenna.4* Mandible.5. Second maxi1la•6. Maxilliped.7a First thoracic leg.8. Second thoracic leg.9a Third thoracic leg.10. Fourth thoracic leg.11. Fifth thoracic leg.

8

EXPLANATION OF PLATE II

Nesippus incisus Heegaard, 1962Figures

12. Dorsal view of female.13. Dorsal view of male.14. First antenna.15. Second antenna.16. Mouth cone and maxilla17. First maxilliped.18. Second maxilliped.19. First thoracic leg.

•oCM Second thoracic leg.

21. Third thoracic leg.22. Fourth thoracic leg.

EXPLANATION OF PLATE III

Eudactvllna sp.Figures

23. Dorsal view of female24. Egg sac.25. First antenna.26. Second antenna.27. Maxilla.28. First maxilliped.29. Second maxilliped.30. Mandible.31. First thoracic leg.32. Second thoracic leg.33. Third thoracic leg.34. Fourth thoracic leg.

/

26

2324

27 30

25

28

3 ?

29

32

3 3

34

EXPLANATION OF PLATE IV

Kg^veria musteli sp. nov.Figures

35. Dorsal view of female36. First antenna.37. Second antenna.38. Mandible.39. First maxilliped.40. Second maxilliped.41. First thoracic leg.42. Second thoracic leg.43. 'Third thoracic leg.44. Fourth thoracic leg.

EXPLANATION OF PLATE V

Nothadichelidium Dolvorionum Ken. et: sp. nov.Figures Figures

45. Dorsal view of female 50. Maxilliped.46. First antenna. 51. First thoracic leg.47. Second antenna. 52. Second thoracic leg48. Endopod of maxilla. 53. Third thoracic leg.49. Exopod of maxilla. 54. Fourth thoracic leg

Neonothadichelidium leotoceohali Ken. et sp. nov.Figures Figures

55. Dorsal view of female. 60. First thoracic leg.56. First antenna. 61. Second thoracic leg57. Second antenna. 62. Third thoracic leg.58. Maxilla, a 63. Fourth thoracic leg59. Maxilliped.

46 4? 484549

50 52

53 54

5657

555859

62

60

63

EXPLANATION OF PLATE VI

Chondracanthus genvpteri Thomson, 1890Figures

64* Dorsal view of female.65* Ventral view of female.66* First antenna of female.67* Second antenna of female.68. Mandible of female.69. Maxilla of female.70. First maxiliiped of female.71* Second maxilliped of female72. Lateral view of male*73. First antenna of male.74. Second antenna of male.75. Maxilla of male.76. Mandible of male.77. First maxilliped of male.78. Second maxilliped of male.79. Second thoracic leg of male

EXPLANATION OF PLATE VII

Clave1lodes intermedius (Quidor, 1906) Wilson, 1915 Figures

80. Ventral view of female^81. Lateral view of male.82. In situ mouth parts of female.83. Maxilla of female.84. First maxilliped of female.85. Second maxilliped of female.

Naobranchia helicolena sp. nov.Figures

86. Dorsal view of female.87. Lateral view of male.88. First antenna of female.89. Second maxilliped of female.90. Maxilla of female.91. Second antenna of female.

0*

V ITA

David Eric Zwerner

Born in-Indianapolis, Indiana, June 16, 1937.Graduated fronr Washington-Lee High Schoo 1 Ar 1 ington,Virginia June, 1955. B.S. (Zoology) The George Washington University, Washington, D* C., June 1959. Entered U. S.Coast Guard, July 1959, and served as Navigator and Operations Officer on the CGC Conifer (WAGL-301) and CGC Jonquil (WAGL-330) at Morehead City, North Carolina and as Search and Rescue Controller and Duty Officer for the Fifth Coast Guard District in Portsmouth, Virginia. Released to ready reserve in December 1962; present rank Lieutenant, USCGR. Entered School of Marine Science (Virginia Institute of Marine Science) of The College of William and Mary in February 1963; graduate assistantship in Parasitology October 1963 - October 1965; Research Assistant in Parasitology October 1965 to present.