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.

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

3

22°

20°

18°

160° 158° 156° 154°

Gardner Pinnacles

La Perouse PinnaclesNecker

Nihoa

7.5 Million Years Ago

4

22°

20°

18°

160° 158° 156° 154°

Necker

Kauai

5 Million Years Ago

Nihoa

NihauKaula

5

22°

20°

18°

160° 158° 156° 154°

Kauai

2.5 Million Years Ago

Nihoa

NihauKaula

Oahu

6

22°

20°

18°

160° 158° 156° 154°

Kauai

1 Million Years Ago

Nihoa

NihauKaula

OahuMaui

7

22°

20°

18°

160° 158° 156° 154°

Kauai

0.5 Million Years Ago

Nihoa

NihauKaula

Oahu

Maui

Mahukona

8

22°

20°

18°

160° 158° 156° 154°

Kauai Present

NihauKaula

Oahu

Maui

Loihi Seamount

Molokai

LanaiKahoolawe Hawaii

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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

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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

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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!

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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

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34

566

7

Equally parsimonious

Rotation Here

Phenotypic Character (1-7) States (0/1)

Plesiomorphic-->Apomorphic Homoplasy

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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

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maukapunaspadixkohalaweliwaikemoikipahulukukuihanastridulansmakaialternatusalapaopuakaheaawilioahuensiselongatusepimeceshypomacronkahiliparakahilialatusflavidushearobustuspiheavictoriae

Prognathogryllus

Kauai

Oahu

Maui

HawaiiIsland FormationMigrationTime

Island Hopping Crickets

Equivocal

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prunacerasinakonahualalaifugaxparanigrakohalaensiskoleanigraproseaeukoleavespertinapacificatantalushapapakokeensiskeahua

Laupala

Kauai

Oahu

Maui

Hawaii

Island FormationMigrationTime

More Island Hopping Crickets

Equivocal

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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

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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

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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

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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

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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

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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

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Loxops caeruleirostrisLoxops coccineusHemignathus virens virensHemignathus virens wilsoniHemignathus virens steinegeriPalmeria doleiHimatione sanguineaVestiaria coccineaHemignathus parvusOreomystis bairdiTelespiza cantansParoreomyza montanaCarpodacus mexicanus OG

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Honeycreeper Cladogram

15

11

1117

8

23

1119

29

33

26

37

19

11

11

7

67

7

8

6

10

mtDNA RFLP

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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 rkoning@snet.net

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Silverswords

N

Kauai-11 endemic

Oahu-2 endemic

Maui Nui-7 endemic

Hawaii-3 endemic

Wilkesia gymnoxiphiumDubautia paleata Argyroxiphium

sandwicense

Plus 5 indigenous species

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QuickTime™ and aTIFF (Uncompressed) decompressor

are needed to see this picture.

http://www.botany.hawaii.edu/faculty/carr/images/dpal_chl.jpg

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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

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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

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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!

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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

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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

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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

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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

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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

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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