1 biogeography of hawaii volcano tops from ocean floor remote colonization rare adaptive radiation...
TRANSCRIPT
![Page 1: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/1.jpg)
1
Biogeography of Hawaii
Volcano tops from Ocean Floor
Remote
Colonization Rare
Adaptive Radiation of Founder Species
Diagrams in this lecture are liberally adapted from:
Wagner, W. L. and V. A. Funk. 1995. Hawaiian biogeography: evolution on a hot spot archipelago. Smithsonian Institution Press, Washington, D.C.
![Page 2: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/2.jpg)
2
Origin of Island Archipelago
• Hot spot moves or plates move over hot spot• Volcanoes build to surface then erode or subside• Current islands arose after previous islands were
already submerged• Exceedingly limited island hopping
North America
Hawaiian Emperor Bend
Hawaiian Ridge
KureMidway
LayannNecker
NihoaKauai
OahuMaui
HawaiiLoihi
Aleutian Islands
Pacific Ocean
Meiji SeamountE
mperor Seam
ounts
![Page 3: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/3.jpg)
3
22°
20°
18°
160° 158° 156° 154°
Gardner Pinnacles
La Perouse PinnaclesNecker
Nihoa
7.5 Million Years Ago
![Page 4: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/4.jpg)
4
22°
20°
18°
160° 158° 156° 154°
Necker
Kauai
5 Million Years Ago
Nihoa
NihauKaula
![Page 5: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/5.jpg)
5
22°
20°
18°
160° 158° 156° 154°
Kauai
2.5 Million Years Ago
Nihoa
NihauKaula
Oahu
![Page 6: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/6.jpg)
6
22°
20°
18°
160° 158° 156° 154°
Kauai
1 Million Years Ago
Nihoa
NihauKaula
OahuMaui
![Page 7: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/7.jpg)
7
22°
20°
18°
160° 158° 156° 154°
Kauai
0.5 Million Years Ago
Nihoa
NihauKaula
Oahu
Maui
Mahukona
![Page 8: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/8.jpg)
8
22°
20°
18°
160° 158° 156° 154°
Kauai Present
NihauKaula
Oahu
Maui
Loihi Seamount
Molokai
LanaiKahoolawe Hawaii
![Page 9: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/9.jpg)
9
Island Development• Island Appears• Colonized by Plants• Biota Establishes• Biota Stabilizes• Island Moves Away
from Hot Spot• Waif Founders Lack
Competition• Speciation: Adaptive
Radiation to fill empty niches
• Endemic Species• Sequence of Islands
should reflect patterns of speciation
• Endemics found on just one island, one volcano
• Humans (about 10 BC)
• Introduced Exotic Species
• Endemics Poor Competitors…Extinct
![Page 10: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/10.jpg)
10
Endemic Terrestrial Species
Taxon/Character Kauai to Hawaii Nihoa to Kure
Insects 2,300 50
Land Snails 1,000 8
Land Birds 70 4
Ferns and Allies 120 0
Flowering Plants 850 12
Area (km2) 16,576 8
Endemics/10 km2 38 1
![Page 11: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/11.jpg)
11
Cladistic Analysis• 1960s Development of Cladistic Analysis
– Determines Possible Pathways of Evolution
• Characters and Polarized Character States– Anatomy, Morphology, Behavior, Chemical, Gene
Sequence– Plesiomorphies-Ancestral character states– Apomorphies-Derived character states
• Construction of Cladogram-computers!– Reveal homoplasies (parallel, convergent), reversals– Most-parsimonious (fewest steps) cladogram– Not necessarily unique!
![Page 12: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/12.jpg)
12
Cladogram
B D F G
grade clade
B F G D
Taxon 1 2 3 4 5 6 7
B 1 0 0 0 0 1 0
D 1 1 0 0 0 0 1
F 1 1 1 1 0 1 0
G 1 1 1 1 1 0 0
12
34
566
7
Equally parsimonious
Rotation Here
Phenotypic Character (1-7) States (0/1)
Plesiomorphic-->Apomorphic Homoplasy
![Page 13: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/13.jpg)
13
Island-Hopping Crickets!
Island Caconemobius Leptogryllus Thaumato-gryllus
Prognatho-gryllus
Trigonidium
Prolaupala Laupala
Nihoa 1 0 1 0 0 0 0Kauai 2 3 1 16 27 0 7Oahu 1 7 0 13 34 1 11
Molokai 2 5 0 0 13 1 1Maui 2 7 1 4 30 1 8Lanai 1 1 0 0 4 0 1
Hawaii 6 5 1 3 27 0 7
![Page 14: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/14.jpg)
14
maukapunaspadixkohalaweliwaikemoikipahulukukuihanastridulansmakaialternatusalapaopuakaheaawilioahuensiselongatusepimeceshypomacronkahiliparakahilialatusflavidushearobustuspiheavictoriae
Prognathogryllus
Kauai
Oahu
Maui
HawaiiIsland FormationMigrationTime
Island Hopping Crickets
Equivocal
![Page 15: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/15.jpg)
15
prunacerasinakonahualalaifugaxparanigrakohalaensiskoleanigraproseaeukoleavespertinapacificatantalushapapakokeensiskeahua
Laupala
Kauai
Oahu
Maui
Hawaii
Island FormationMigrationTime
More Island Hopping Crickets
Equivocal
QuickTime™ and aTIFF (Uncompressed) decompressor
are needed to see this picture.
http
://w
ww
.wam
.um
d.ed
u/~f
ergu
sd/L
aup3
.jpg
![Page 16: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/16.jpg)
16
Amazing Fruit Flies!• 511 of 2,300 endemic insects are drosophiloids• DNA sequences show 10 MYBP arrived Necker• Necker has subsided and lost habitat…now n=0• All 511 endemic species are a single clade!• Analyzed by morphology, karyotype, molecular
analysis
Drosophila sproati
http://www.msu.edu/~taylo110/DSproati.JPG
![Page 17: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/17.jpg)
17
D. melanogasterD. robustaD. melanicaD. funebrisD. pinicolaD. imigransD. repletaE. crassifemurS. exiguaD. mimicaD. sproatiZaprionusChymomyzaHirtodrosophilaScaptodrosophila
Outgroups
Hawaiian
Continental
Numbers ofmtDNA base transitions
Drosophilid Comparisons15
16
1676
9
1010
141113
101111
3816
10
6
9
8
8
77
8
7
7
Purine AGPyrimidine CT
![Page 18: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/18.jpg)
18
Primitive Phylogram of Drosophiloidsbostrycha disjuncta
affinidisjuncta
grimshawl
recticilia
musaphilia
hawaiiensis
gradata
spectabilis
setosimentum
clavisetae
adiastolapeniculipediscilifera
planitibia silvestris
heteroneurahemipeza
cyrtoloma
neoperkinsineopicta
hanaulaeoahuensis
nigribasismelanocephala
obscuripes
picticornis
mimica
arcuatuslongiseta
adunca
cognatayoonitanythrix
differens
chromosomal changes
![Page 19: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/19.jpg)
19
Drosophilid Area Cladogram
ContinentalOahuMolokaiWest MauiEast Maui 1East Maui 2Hawaii KonaHawaii Hilo
Just as observed among crickets, these fruit flies radiate in ways that correlate with the biogeography of the Hawaiian Islands
![Page 20: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/20.jpg)
20
Spiders Spinning Southeast!
T. kauaiensis
T. tantalus
T. waikamoi
T. polychromata
T. macracantha
T. brevignatha
T. brevignatha
T. stelarobusta
T. pallescens
T. restricta
T. restricta
T. quasimodo
T. quasimodo
T. quasimodo
T. quasimodo
T. kamakou
T. kamakou
T. perreirai
T. pilosa
N
1
1
1
1
11
2
2
22
3
3
3
3
33
4
4
4
4
44
6
5
364
6
4
3
Tetragnatha
http
://w
ww
.ent
omol
ogy.
corn
ell.e
du/F
acul
ty_S
taff
/Bla
ckle
dge/
tetr
agna
tha.
JPG
![Page 21: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/21.jpg)
21
Honeycreepersof
HawaiiVestiaria coccinea
Hemignathus virens
Extreme isolation limits vertebrate
fauna
No native amphibians
No native reptiles
1mammal (bat)
1 bird established and
radiated into 33 extant
and 14 extinct species!
Telespiza cantans
Vestiaria coccinea
Hemignathus parvus
Palmeria dolei
Oreomystis bairdi
http://www.state.hi.us/dlnr/mfbp/images/akohekohe.jpght
tp://
ww
w.o
ahun
atur
etou
rs.c
om/
imag
es/la
ysan
finc
h.jp
g
http://www.uatuahine.hawaii.edu/hana/NaManu/iiwi/iiwi.jpeg
http
://w
ww
.sta
nfor
d.ed
u/~p
etel
at1/
hcre
epr1
.jpg
http
://w
ww
.sta
nfor
d.ed
u/~p
etel
at1/
ani5
.jpg
![Page 22: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/22.jpg)
22
Loxops caeruleirostrisLoxops coccineusHemignathus virens virensHemignathus virens wilsoniHemignathus virens steinegeriPalmeria doleiHimatione sanguineaVestiaria coccineaHemignathus parvusOreomystis bairdiTelespiza cantansParoreomyza montanaCarpodacus mexicanus OG
9
Honeycreeper Cladogram
15
11
1117
8
23
1119
29
33
26
37
19
11
11
7
67
7
8
6
10
mtDNA RFLP
![Page 23: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/23.jpg)
23
How about Plants? Hibiscadelphus
distans Kauai
wilderianus Maui
crucibracteatus Lanai
giffardianus Hawaii
hualalaiensis Hawaii
bombycinus Hawaii
The phylogram is also an area cladogram showing the positive correlation between evolution and oceanic island biogeography.
Hibiscadelphus distans
Photo Source: Unknown…If yours, please send email to [email protected]
![Page 24: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/24.jpg)
24
Silverswords
N
Kauai-11 endemic
Oahu-2 endemic
Maui Nui-7 endemic
Hawaii-3 endemic
Wilkesia gymnoxiphiumDubautia paleata Argyroxiphium
sandwicense
Plus 5 indigenous species
http
://bi
olog
y.us
gs.g
ov/s
+t/l
rgim
age/
t172
w01
.jpg
http
://hb
s.bi
shop
mus
eum
.org
/goo
d-ba
d/im
ages
/ilia
u.jp
g
QuickTime™ and aTIFF (Uncompressed) decompressor
are needed to see this picture.
http://www.botany.hawaii.edu/faculty/carr/images/dpal_chl.jpg
![Page 25: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/25.jpg)
25
Silversword Origins• Closest relatives are California tarweeds
• 3,900 km open ocean crossing
• Fruits too thin-walled to survive bird gut
• Fruits do not float in sea water
• Fruits too heavy to float on wind
• Hairy and sticky sepals suggest bird attachment
• Preened off on Kauai, then radiated and migrated
![Page 26: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/26.jpg)
26
Tarweed (California USA) OGDubautia paleata KauaiDubautia waialealae KauaiDubautia laxa indigenousWilkesia gymnoxiphium KauaiWilkesia hobdyi KauaiArgyroxiphium caliginis MauiArgyroxiphium grayanum MauiArgyroxiphium virescens MauiArgyroxiphium sanwicense M&HArgyroxiphium kauense Hawaii
Silversword Cladogram (simplified)
QuickTime™ and aTIFF (Uncompressed) decompressor
are needed to see this picture.
http://www.calflora.net/bloomingplants/images/redrocktarweed4.jpg
![Page 27: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/27.jpg)
27
Geranium (pandemic) OGarboreum E. Maui subalpinemultiflorum E. Maui subalpine cuneatum Hawaii sub-alpine tridens E. Maui alpinehanaense E. Maui boghumile W. Maui bog kauaiense Kauai bog
Not all cladograms tell the same story!
Geranium arboreum
Geranium hanaenseGeranium cuneatum
Geranium colonized “late” E. Maui and
radiated and migrated speciating most recently on Kauai with ecological
correlations!
http
://w
ww
.bot
any.
haw
aii.e
du/
facu
lty/c
arr/
imag
es/g
er_a
rb.jp
g
http://www.botany.hawaii.edu/faculty/carr/images/ger_han_cu.jpg
http://www.botany.hawaii.edu/faculty/carr/images/ger_cun_cu.jpg
![Page 28: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/28.jpg)
28
humile E. Mauicapillare W. Mauiremyi W. Maui, Lanairockii Molokai sylvae Molokai, Mauitenerrimum Oahuarenarium E. Maui consanguineum Hawaiiconyzoides Maui Nui, Hawaiilepidolum E. Maui, Lanai, Oahu
Not all cladograms tell the same story!
Tetramolopium humile
Tetramolopium colonized “late” E. Maui and
radiated and migrated to the other islands
http://www.botany.hawaii.edu/faculty/carr/images/tet_hum.jpg
![Page 29: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/29.jpg)
29
![Page 30: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/30.jpg)
30
Biotic Factors
• Symbioses (living together)– Mutualism: zooxanthellae/corals/forams– Commensalism: bromeliads on trees– Parasitism: epiphytic woevine on shrubs
• Competitors: Epiphytes, Allelopathy, Exotics
• Herbivory
• Pollination, Seed Dispersal– Partners required for reproduction
![Page 31: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/31.jpg)
31
Abiotic Factors
• Light-wavelength (color), amplitude (intensity)– Water absorbs low energy wavelengths…leaving blue– Turbidity (milkiness) vs Turbulence (mixing)– Chlorophyll (purple, blue, yellow, red)– Carotenoids (green)– Phycobilins (green to red) depending on type/depth
• Gas Availability (N2=78%, O2=21%, CO2<1%)– Oxygen poisons photosynthesis (photorespiration)– CO2 highly soluble in water, O2 virtually insoluble– Most algae are C3, many terrestrials C4 or CAM
![Page 32: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/32.jpg)
32
Abiotic Factors
• Temperature-plants are essentially poikilotherms– Water in tidepools rises rapidly
• Eurythermal = wide tolerance, Stenothermal = narrow tolerance
– Dunes heat unless leaves shade them or burrow deeply– Ice damage, frost, heat shock– Large body size: thermal insulation vs heat load– Surface:Volume ratio (Neither Fucus nor Ascophyllum here!)
• Desiccation: evaporation, osmotic stress– Saccate thallus (Colpomenia or Valonia)– Mucilaginous sheath, Waxy Cuticle, Hairy Epidermis
![Page 33: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/33.jpg)
33
Abiotic Factors
• Salinity: osmotic stress, Na+ or Cl- toxicity– Salinity in small tidepools rises with evaporation– Salinity in small tidepools falls with rainfall
• Euryhaline = wide tolerance, Stenohaline = narrow tolerance
– Halophytes• Exudation: Avicennia via hydathodes (salt glands)• Exclusion: Rhizophora ion pumps keep salt out• Shedding: Spartina, annual dicots, leaves abscise taking salt out• Acclimitization: mannitol, sucrose, betaine, etc.
• Soil Minerals– Oligotrophic Waters are Depauperate in Minerals (N, P, Fe)– Eutrophic Estuarine waters: Algal Blooms
![Page 34: 1 Biogeography of Hawaii Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species Diagrams in this lecture are liberally](https://reader036.vdocument.in/reader036/viewer/2022070305/5515151d550346a80c8b5d43/html5/thumbnails/34.jpg)
34
Abiotic Factors
• Surge Stress, Drag Stress, Shear Stress– Turbulence in Rocky Intertidal– Scouring in Sandy Intertidal
• Euryhaline = wide tolerance, Stenohaline = narrow tolerance
– Benefits• Predator Restriction• Parasite Removal• Equalize Salinity and Temperature• Adds CO2 for photosynthesis, but also adds O2
– Rhodoliths in deep waters…detached coralline red algae– Phytoplankton in Photic Zone of open ocean only 5000 spp!
Greater diversity and abundance in Estuarine waters