Flora of Hawaii•• since so isolated, it has a depauperate flora and consists entirely of waif since so isolated, it has a depauperate flora and consists entirely of waifelements derived from LDDelements derived from LDD

Argyroxiphium sandwicense - Haleakala silversword

native

Passiflora mollissima - Banana poka

or weeds

Flora of Hawaii•• individual islands tend to fit the expected number of species based on area individual islands tend to fit the expected number of species based on area

- except most recent Hawaii- except most recent Hawaii

Flora of Hawaii•• endemism high in flowering plants endemism high in flowering plants

1,817 including introduced1,817 including introduced

649 including introduced649 including introduced

87 native families87 native families

•• 0% endemic 0% endemic

146 including introduced146 including introduced

956 native species956 native species

•• 89% endemic 89% endemic

216 native genera216 native genera

•• 15% endemic 15% endemic

BrighamiaLobeliaceae

Flora of Hawaii•• endemism highest in the world endemism highest in the world

BrighamiaLobeliaceae

•• speciation far exceeds immigration speciation far exceeds immigration

•• Hawaiian flora so distinctive, often assigned its own floristic region Hawaiian flora so distinctive, often assigned its own floristic region

•• Why so distinctive? Why so distinctive?

Flora of Hawaii•• distance from source area and thus few immigrants distance from source area and thus few immigrants

•• archipelago effect and inter-island archipelago effect and inter-islandspeciationspeciation

Flora of Hawaii•• extraordinary rich and diverse habitats available extraordinary rich and diverse habitats available

•• 38 community types 38 community typesrecognizedrecognized

Flora of Hawaii•• disharmonic flora disharmonic flora

•• only 3 native orchid only 3 native orchidspeciesspecies

•• but many introduced but many introducedorchids that becomeorchids that becomenaturalizednaturalized

Dendrobium & honey bees

Flora of Hawaii•• disharmonic flora disharmonic flora

Pritchardia beccariana

•• only 1 native palm genus only 1 native palm genus

•• native gymnosperms and primitive native gymnosperms and primitiveangiosperms absentangiosperms absent

Flora of Hawaii•• disharmonic flora - 15 largest families of native species disharmonic flora - 15 largest families of native species

•• largelarge source area families source area familiesunder representedunder represented

•• small Campanulaceae -small Campanulaceae -lobeliads and bellflowers -lobeliads and bellflowers -over representedover represented

Flora of Hawaii•• immigrants: 270 (or less!) gave rise to 956 native angiosperms immigrants: 270 (or less!) gave rise to 956 native angiosperms

Cyrtandra Pelea

Cyanea

•• 1/10 1/10thth of colonist generates of colonist generates 1/2 of species ! 1/2 of species !

Flora of Hawaii•• immigrants: similar story with other biota immigrants: similar story with other biota

•• one American finch gave rise to 33 extant and 14 extinct species of honeycreepers one American finch gave rise to 33 extant and 14 extinct species of honeycreepers

Flora of Hawaii•• immigrants: similar story with other biota immigrants: similar story with other biota

•• 800 land molluscs from 20 colonists 800 land molluscs from 20 colonists

•• 5,800 insects from 250 colonists 5,800 insects from 250 colonists

Flora of Hawaii•• immigrants: but different story for ferns! immigrants: but different story for ferns!

•• 168 ferns from 135 colonists - implying constant immigration (lack of genetic 168 ferns from 135 colonists - implying constant immigration (lack of geneticisolation) and/or few speciation eventsisolation) and/or few speciation events

Sadleria - 6 species Polypodium - 1 species

vs.vs.

Flora of Hawaii•• LDD mechanisms LDD mechanisms

internal internalBIBI

mud mudBMBM

viscidviscidBVBV

barbs/hooksbarbs/hooksBBBB

air airAA

driftdriftDFDF

drift - raredrift - rareDRDR

39%

13%

10%

12%

1%

14%

9%

Leptocophylla BIBI

Argyroxiphium BBBB

Acacia - koa DRDR Viola BMBM

Flora of Hawaii•• floristic affinities floristic affinities

LeptocophyllaEpacridaceae

•• despite proximity of despite proximity ofHawaii to N America,Hawaii to N America,almost half of colonists arealmost half of colonists areof Austral-asian originof Austral-asian origin

Flora of Hawaii•• floristic affinities floristic affinities

•• few from Arctic-Beringia few from Arctic-Beringia

Viola

Plantago - plantain

Flora of Hawaii•• floristic affinities floristic affinities

•• woody violets considered closely woody violets considered closelyrelated to South American violetsrelated to South American violets

Viola

•• all native Hawaiian violets are polyploid all native Hawaiian violets are polyploidand derived from polyploid races of oneand derived from polyploid races of oneBeringian tundra species - the AlaskanBeringian tundra species - the Alaskanvioletviolet

Viola langsdorfii

Flora of Hawaii•• floristic affinities floristic affinities

•• 18-22% from western 18-22% from westernNorth AmericaNorth America

Argyroxiphium

Flora of Hawaii•• adaptive radiation of Hawaiian silverswords adaptive radiation of Hawaiian silverswords

•• silversword alliance of 3 genera and 28 silversword alliance of 3 genera and 28speciesspecies

•• trees, shrubs, subshrubs, rosette plants, trees, shrubs, subshrubs, rosette plants,cushion plants, vinescushion plants, vines

Flora of Hawaii

•• B. Baldwin verified that the Hawaiian B. Baldwin verified that the Hawaiiansilversword radiation derived from silversword radiation derived from CaliforniaCalifornia

•• the closest species is now the closest species is nowcalled called Carlquistia muiriiCarlquistia muirii

•• this origin occurred aboutthis origin occurred about5 mya5 mya

•• rapid radiation at base rapid radiation at base

•• adaptive radiation of Hawaiian silverswords adaptive radiation of Hawaiian silverswords

Flora of Hawaii•• inter-island biogeography [ inter-island biogeography [““island hoppingisland hopping””]]

Argyroxiphium

•• most speciation events occur on individual islands most speciation events occur on individual islands

•• few successful inter-island colonizationsfew successful inter-island colonizations

•• movement is from older to younger islands movement is from older to younger islands

Flora of Hawaii•• inter-island biogeography [ inter-island biogeography [““island hoppingisland hopping””]]

Spiders Spinning Southeast!T. kauaiensis

T. tantalus

T. waikamoi

T. polychromata

T. macracantha

T. brevignatha

T. brevignatha

T. stelarobustaT. 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

Geranium (pandemic) OGarboreum E. Maui sub-alpinemultiflorum E. Maui sub-alpine cuneatum Hawaii sub-alpine tridens E. Maui alpinehanaense E. Maui boghumile W. Maui bog kauaiense Kauai bog

Geranium arboreum

Geranium hanaenseGeranium cuneatum

Geranium colonized“late” E. Maui and

radiated and migratedspeciating most recentlyon Kauai with ecological

correlations!

Flora of Hawaii•• inter-island biogeography - not all the same story! inter-island biogeography - not all the same story!

Flora of Hawaii•• extinction: on-going process on islands (as well as immigration) extinction: on-going process on islands (as well as immigration)

•• but due to humans . . . but due to humans . . .

•• 107 of 956 angiosperm species 107 of 956 angiosperm species —— 11% extinct 11% extinct

•• 423 of 956 species 423 of 956 species —— 38% extinct or threatened 38% extinct or threatened

Flora of Hawaii•• extinction: rogue gallery of exotics extinction: rogue gallery of exotics

Passiflora mollissima - banana poka

Miconia introduced intoHawaii as ornamental

Miconia calvescens“green cancer”

Flora of Hawaii•• extinction: rogue gallery of exotics extinction: rogue gallery of exotics

Why the Hawaiian lobeliads? Why the Hawaiian lobeliads?

•• largest grouplargest group: : 6 genera,6 genera,115 species)115 species)•• 1/9th of native flora1/9th of native flora•• considered derived from considered derived from3-5 separate colonizations3-5 separate colonizations•• phenomenal phenomenal variation invariation inhabitat, life form, flowers,habitat, life form, flowers,and fruitsand fruits

Hawaiian LobeliadsWhy the Hawaiian lobeliads? Why the Hawaiian lobeliads?

•• appear to have appear to have co-co-evolved evolved with the endemicwith the endemicHawaiian Hawaiian honeycreepershoneycreepers

Hawaiian Lobeliads

Why the Hawaiian lobeliads? Why the Hawaiian lobeliads?

•• appear to have appear to have co-co-evolved evolved with the endemicwith the endemicHawaiian Hawaiian honeycreepershoneycreepers•• honeycreepers representhoneycreepers representa separatea separate adaptiveadaptiveradiationradiation

Hawaiian Lobeliads

Lobelia gloria-montisLobelia gloria-montis

Lobelia telekii - Lobelia telekii - Mt. KenyaMt. Kenya

What are the Hawaiian lobeliads? What are the Hawaiian lobeliads?

Hawaiian Lobeliads

BrighamiiBrighamii

What are the Hawaiian lobeliads? What are the Hawaiian lobeliads?

Hawaiian Lobeliads

DelissiaDelissia

What are the Hawaiian lobeliads? What are the Hawaiian lobeliads?

Hawaiian Lobeliads

TrematolobeliaTrematolobelia

What are the Hawaiian lobeliads? What are the Hawaiian lobeliads?

Hawaiian Lobeliads

ClermontiaClermontia

What are the Hawaiian lobeliads? What are the Hawaiian lobeliads?

Hawaiian Lobeliads

CyaneaCyanea

What are the Hawaiian lobeliads? What are the Hawaiian lobeliads?

Hawaiian LobeliadsThe Hawaiian lobeliads represent a The Hawaiian lobeliads represent a single lineage/colonizationsingle lineage/colonization!!

Hawaiian Lobeliads

The Hawaiian lobeliads - more spectacular adaptive radiation The Hawaiian lobeliads - more spectacular adaptive radiation

Hawaiian LobeliadsRemarkable divergent andRemarkable divergent andconvergent character evolution - butconvergent character evolution - butfruit color is perfectly consistent fruit color is perfectly consistent withwithrelationshipsrelationships

Hawaiian Lobeliads

Two clock calibrations - using AsteridTwo clock calibrations - using Asteridfossils or using Hawaiian Island ages -fossils or using Hawaiian Island ages -place the differentiation of place the differentiation of HawaiianHawaiianlobeliads at 13-14 myalobeliads at 13-14 mya

Hawaiian Lobeliads

Original colonist arrived inOriginal colonist arrived inLaPerouse/Gardner Pinnacles LaPerouse/Gardner Pinnacles - large- largevolcanic islands 10-15 myavolcanic islands 10-15 mya

Hawaiian LobeliadsTwo clock calibrations - using AsteridTwo clock calibrations - using Asteridfossils or using Hawaiian Island ages -fossils or using Hawaiian Island ages -place the differentiation of place the differentiation of HawaiianHawaiianlobeliads at 13-14 myalobeliads at 13-14 mya

Hawaiian LobeliadsTwo clock calibrations - using AsteridTwo clock calibrations - using Asteridfossils or using Hawaiian Island ages -fossils or using Hawaiian Island ages -place the differentiation of place the differentiation of HawaiianHawaiianlobeliads at 13-14 myalobeliads at 13-14 mya

Original colonist arrived inOriginal colonist arrived inLaPerouseLaPerouse//Gardner Pinnacles Gardner Pinnacles - large- largevolcanic islands 10-15 myavolcanic islands 10-15 mya

Hawaiian LobeliadsOne of the oldest radiations of extant Hawaiian biota - honeycreeper pollinators wouldOne of the oldest radiations of extant Hawaiian biota - honeycreeper pollinators wouldnot arrive for another 7-9 mynot arrive for another 7-9 my

Hawaiian LobeliadsBut . . . a 2008 paper shows that the 5 recently extinctBut . . . a 2008 paper shows that the 5 recently extinctHawaiian Honeyeaters Hawaiian Honeyeaters –– thought to be recently derived thought to be recently derivedfrom from AustralAsian Honeyeaters AustralAsian Honeyeaters –– are in fact 14-17 my are in fact 14-17 myold and derived from an American clade - waxwings,old and derived from an American clade - waxwings,silky flycatchers, and palm chatssilky flycatchers, and palm chats

Hawaiian lobeliadsHawaiian lobeliads13-14 mya13-14 mya

Hawaiian HoneycreepersHawaiian Honeycreepers7 mya7 mya

““Assume nothing in Hawaiian naturalAssume nothing in Hawaiian naturalhistoryhistory”” Frank Howarth, 1992