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 Hawaiian biogeography: evolution on a hot spot archipelago. Smithsonian Institution Press, Washington, D.C.

2. Origin of Island Archipelago
North America
Meiji Seamount
Aleutian Islands
Pacific Ocean
Emperor Seamounts
Hawaiian Emperor Bend
Kure
Midway
Layann
Necker
Nihoa
Kauai
Hawaiian Ridge
Oahu
Maui
Hawaii
Loihi
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

3. 7.5 Million Years Ago
22°
Gardner Pinnacles
La Perouse Pinnacles
20°
Necker
Nihoa
18°
160°
158°
156°
154°

4. 5 Million Years Ago
22°
Necker
Nihoa
20°
Kauai
Nihau
Kaula
18°
160°
158°
156°
154°

5. 2.5 Million Years Ago
22°
Nihoa
Kauai
20°
Oahu
Nihau
Kaula
18°
160°
158°
156°
154°

6. 1 Million Years Ago
22°
Nihoa
Kauai
Oahu
20°
Nihau
Maui
Kaula
18°
160°
158°
156°
154°

7. 0.5 Million Years Ago
Nihoa
22°
Kauai
Oahu
Nihau
Kaula
Maui
20°
Mahukona
18°
160°
158°
156°
154°

8. Present
Kauai
22°
Oahu
Nihau
Molokai
Kaula
Maui
Lanai
Kahoolawe
Hawaii
20°
Loihi Seamount
18°
160°
158°
156°
154°

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

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
Flowering Plants
850 12
Area (km2)
16,576
Endemics/10 km2 38 1

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!

12. Cladogram grade clade Equally parsimonious B D F G B F G D6 6 7 5 4 3 2 1
Rotation Here
Phenotypic Character (1-7) States (0/1)
Taxon 1 2 3 4 5 6 7 B D F G
Plesiomorphic-->Apomorphic
Homoplasy

13. Island-Hopping Crickets!
Caconemobius
Leptogryllus
Thaumato-gryllus
Prognatho-gryllus
Trigonidium
Prolaupala
Laupala
Nihoa 1
Kauai 2 3 16 27 7
Oahu 13 34 11
Molokai 5
Maui 4 30 8
Lanai
Hawaii 6

14. Island Hopping Crickets
Prognathogryllus
mauka
puna
spadix
kohala
weli
waikemoi
kipahulu
kukui
hana
stridulans
makai
alternatus
alapa
opua
kahea
awili
oahuensis
elongatus
epimeces
hypomacron
kahili
parakahili
alatus
flavidus
hea
robustus
pihea
victoriae
Hawaii
Equivocal
Island Formation
Migration
Time
Maui
Oahu
Kauai

15. More Island Hopping Crickets
Laupala
pruna
cerasina
kona
hualalai
fugax
paranigra
kohalaensis
kolea
nigra
prosea
eukolea
vespertina
pacifica
tantalus
hapapa
kokeensis
keahua
Hawaii
Island Formation
Migration
Time
Equivocal
Maui
Oahu
Kauai

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

17. Drosophilid Comparisons15
D. melanogaster
D. robusta
D. melanica
D. funebris
D. pinicola
D. imigrans
D. repleta
E. crassifemur
S. exigua
D. mimica
D. sproati
Zaprionus
Chymomyza
Hirtodrosophila
Scaptodrosophila 16
Continental 16 8 9 9 7 7 6
Numbers of
mtDNA base
transitions 8 10 10 7 14 6 7
Purine AG 11 7 13
Pyrimidine CT 8 10
Hawaiian 10 11 11 38
Outgroups 16

18. Primitive Phylogram of Drosophiloids
chromosomal changes
recticilia
bostrycha
disjuncta
musaphilia
gradata
affinidisjuncta
hawaiiensis
grimshawl
planitibia
silvestris
spectabilis
differens
heteroneura
adiastola
peniculipedis
cilifera
setosimentum
hemipeza
cyrtoloma
picticornis
clavisetae
hanaulae
oahuensis
neoperkinsi
arcuatus
neopicta
longiseta
nigribasis
melanocephala
adunca
mimica
obscuripes
yooni
tanythrix
cognata

19. Drosophilid Area Cladogram
Continental
Oahu
Molokai
West Maui
East Maui 1
East Maui 2
Hawaii Kona
Hawaii Hilo
Just as observed among crickets, these fruit flies radiate in ways that correlate with the biogeography of the Hawaiian Islands

20. Spiders Spinning Southeast!
Tetragnatha 5
T. pilosa 1
T. kauaiensis 6
T. perreirai 4 4 2
T. tantalus
T. kamakou 4 3
T. kamakou 4 3 3 2 2 1
T. quasimodo
T. waikamoi
T. quasimodo 3
T. quasimodo 1 6 3 1 4
T. polychromata
T. quasimodo 3
T. restricta 4 6 1
T. macracantha
T. restricta 2 3 N
T. brevignatha 4 1
T. brevignatha 3
T. stelarobusta 4
T. pallescens

21. Honeycreepers of Hawaii
Honeycreepers of Hawaii
Vestiaria coccinea
Palmeria dolei
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!
Hemignathus virens
Telespiza cantans
Hemignathus parvus
Vestiaria coccinea
Oreomystis bairdi

22. Honeycreeper Cladogram9
Loxops caeruleirostris
Loxops coccineus
Hemignathus virens virens
Hemignathus virens wilsoni
Hemignathus virens steinegeri
Palmeria dolei
Himatione sanguinea
Vestiaria coccinea
Hemignathus parvus
Oreomystis bairdi
Telespiza cantans
Paroreomyza montana
Carpodacus mexicanus OG 10 15
mtDNA RFLP 11 19 6 7 11 17 7 7 8 11 23 11 11 8 19 29 6 33 26 37

23. How about Plants? Hibiscadelphus
Hibiscadelphus distans
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.
Photo Source: Unknown…If yours, please send to

24. Wilkesia gymnoxiphium
Silverswords
Kauai-11 endemic
Oahu-2 endemic
Wilkesia gymnoxiphium
Dubautia paleata
Maui Nui-7 endemic
Argyroxiphium
sandwicense
Hawaii-3 endemic N
Plus 5 indigenous species

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

26. Silversword Cladogram (simplified)
Tarweed (California USA) OG
Dubautia paleata Kauai
Dubautia waialealae Kauai
Dubautia laxa indigenous
Wilkesia gymnoxiphium Kauai
Wilkesia hobdyi Kauai
Argyroxiphium caliginis Maui
Argyroxiphium grayanum Maui
Argyroxiphium virescens Maui
Argyroxiphium sanwicense M&H
Argyroxiphium kauense Hawaii

27. Not all cladograms tell the same story!
Geranium arboreum
Geranium (pandemic) OG
arboreum E. Maui subalpine
multiflorum E. Maui subalpine
cuneatum Hawaii sub-alpine
tridens E. Maui alpine
hanaense E. Maui bog
humile W. Maui bog
kauaiense Kauai bog
Geranium colonized “late” E. Maui and radiated and migrated speciating most recently on Kauai with ecological correlations!
Geranium cuneatum
Geranium hanaense

28. Not all cladograms tell the same story!
Tetramolopium colonized “late” E. Maui and radiated and migrated to the other islands
humile E. Maui
capillare W. Maui
remyi W. Maui, Lanai
rockii Molokai
sylvae Molokai, Maui
tenerrimum Oahu
arenarium E. Maui
consanguineum Hawaii
conyzoides Maui Nui, Hawaii
lepidolum E. Maui, Lanai, Oahu
Not all cladograms tell the same story!
Tetramolopium humile

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

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

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

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

34. Greater diversity and abundance in Estuarine waters
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