Origin of Island Archipelago Empe Biogeography of Hawaii Aleutian Islands ror Se amou Volcano tops from Ocean Floor Remote Colonization Rare Adaptive Radiation of Founder Species North America Meiji Seamoun t Pacific Ocean nts Hawaiian Emperor Bend Kure Midway Layann Necker Nihoa Kauai Oahu Maui Hawaii Loihi Haw aiian Ridg e • Hot spot moves and plate moves over hot spot • Volcanoes build to surface then erode or subside • Current islands arose after previous islands were already submerged • Exceedingly limited island hopping 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. 1 Detroit 80 Ma Suiko 61 Ma Nintoku 56 Ma Koko 49 Ma Diakakuji 47 Ma Midway 27 Ma Necker 10 Ma Time/motion data adapted from: Tarduno, et al. 2009. The bent Hawaiian-Emperor hotspot track: inheriting the mantle wind. Science 324: 50-53 Motion of hotspot magma plume under plate? 2 Necker Plate motion over fixed hotspot? 3 4 1 Kauai Oahu Nihau Molokai Necker Maui Lanai Kahoolawe Hawaii 5 6 7.5 Million Years Ago 22° 5 Million Years Ago 22° Gardner Pinnacles Necker 20° Nihoa La Perouse Pinnacles 20° Necker Kauai Nihoa 18° Kaula 18° 160° 158° 156° 154° 160° 7 158° Nihau 156° 154° 8 2 2.5 Million Years Ago 1 Million Years Ago 22° 22° Nihoa Kauai Nihoa Kauai 20° 20° Oahu Nihau Kaula 18° Oahu Nihau Kaula Maui 18° 160° 158° 156° 154° 160° 158° 156° 154° 9 10 0.5 Million Years Ago 22° 22° Kauai Nihau Kaula Nihau Kaula Oahu Maui 20° Present Kauai Nihoa Oahu Molokai Maui Lanai Kahoolawe Hawaii 20° Mahukona Loihi Seamount 18° 18° 160° 158° 156° 154° 160° 11 158° 156° 154° 12 3 Island Development • • • • • • Island appears Marine/shore colonized Island colonized by plants Terrestrial biota establishes Biota stabilizes Island moves away from hot spot • Waif founders lack competition • Speciation: adaptive radiation to fill empty niches Endemic Terrestrial Species • Endemic species evolve • Sequence of islands should reflect patterns of speciation • Endemics found on just one island, one volcano • Humans arrive (about 10 BC) • Introduced exotic species are released • Endemics that may be poor competitors…extinct 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 16,576 8 38 1 Area (km2) Endemics/10 km2 13 Cladistic Analysis 14 How do you DO cladistics? 1. Look at a group of organisms that you think are related • 1960s Development of Cladistic Analysis – Determines Possible Pathways of Evolution 2. Find a not-too-distantly related (primitive?) out-group • Characters and Polarized Character States 3. Select characters that will help to distinguish the organisms – Anatomy, Morphology, Behavior, Chemical, Gene Sequence – Plesiomorphies-Ancestral character states – Apomorphies-Derived character states 4. Polarize the character states by: • Stratigraphic sequence (fossil sequence) • Developmental sequence (ontogeny recaps phylogeny) • Outgroup comparison • Construction of Cladogram-computers! 5. Build a data matrix – Reveal homoplasies (parallel, convergent), reversals – Most-parsimonious (fewest steps) cladogram – Not necessarily unique! 6. Group by number of synapomorphies (shared derived) 7. Sketch possible cladograms 8. Seek simplest (most parsimonious) cladogram 15 16 4 Asia ATG Area cladogram showing progression process Asia OG OG Oahu ATC Kauai ATC Hawaii ATC Maui ATC B A Character: Apomorphic State 1 Eye size: Large Black Wide 4 Body width: Wide 5 Body color: Dark 6 Wing width: Wide 7 Wing length: Long 8 Leg length: Long 9 Leg thickness: Thick 10 DNA transversion: G>C 2 Eye color: 3 Neck width: C Taxon: A B C D E 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 1 1 1 0 0 0 0 1 1 0 0 1 0 1 0 1 1 1 0 1 1 1 0 0 0 1 1 1 0 1 1 1 1 0 0 0 0 1 0 0 0 1 E 2 1 1 3 2 5 1 2 3 5 Nihoa Kauai Oahu Molokai Maui Lanai Hawaii Oahu Maui Hawaii A D C autapomorphies -- 2 black eye -- 7 long wing -- 3 wide neck -- 8 long leg -- 5 dark body -- 1 large eye -- 9 thick leg -- 4 wide body apomorphies E B A D C -- 2 -- 7 -- 3 -- 8 -- 5 -- 1 OG -- 9 -- 4 -- 10 -- 6 -- 6 wide wing plesiomorphies -- 10 DNA transversion G>C 18 17 Island Hopping Crickets Island-Hopping Cricket Species! Island Kauai B Kauai ATC D OG E Hawaii Caconemobius Leptogryllus Thaumato -gryllus Prognathogryllus Trigonidium Prolaupala Laupala 1 2 1 2 2 1 6 0 3 7 5 7 1 5 1 1 0 0 1 0 1 0 16 13 0 4 0 3 0 27 34 13 30 4 27 0 0 1 1 1 0 0 0 7 11 1 8 1 7 Island Formation Migration Time Oahu Kauai 19 Maui Prognathogryllus Equivocal 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 20 5 http://www.wam.umd.edu/~fergusd/Laup3.jpg More Island Hopping Crickets pruna cerasina kona hualalai fugax paranigra kohalaensis kolea nigra prosea eukolea vespertina pacifica tantalus hapapa kokeensis keahua Hawaii Island Formation Migration Time Amazing Fruit Flies! Laupala Equivocal Maui Oahu Kauai • • • • • 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 21 Primitive Phylogram of Drosophiloids Drosophilid Comparisons 15 16 Continental 8 9 Numbers of mtDNA base transitions Purine ADG Pyrimidine CDT Outgroups 7 8 6 16 7 6 9 10 10 7 7 7 10 Hawaiian 11 11 38 16 8 22 14 11 13 10 bostrycha 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 Proper cladograms have branch length correlated to the number of steps disjuncta chromosomal changes gradata affinidisjuncta planitibia differens hemipeza cyrtoloma picticornis neoperkinsi neopicta nigribasis obscuripes melanocephala 23 musaphilia hawaiiensis grimshawl silvestris heteroneura hanaulae oahuensis recticilia spectabilis adiastola setosimentum peniculipedis clavisetae cilifera arcuatus longiseta mimica adunca yooni cognata tanythrix 24 6 Drosophilid Area Cladogram 1 4 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 3 5 T. pilosa T. kauaiensis ck 3 1 6 T. perreirai Ba T. tantalus dis per 4 T. kamakou 2 saT.l kamakou T. quasimodo T. waikamoi 4 3 4 2 2 T. quasimodo 6 3 T. quasimodo 3 4 T. polychromata 6 T. macracantha T. quasimodo 3 1 1 T. restricta 2 3 4 T. restricta 4 N T. brevignatha 1 1 http://www.entomology.cornell.edu/Faculty_Staff/Blackledge/tetragnatha.JPG Spiders Spinning Southeast! Tetragnatha T. brevignatha 3 4 25 T. stelarobusta T. pallescens 26 http://www.state.hi.us/dlnr/mfbp/images/akohekohe.jpg Honeycreepers of Hawaii Vestiaria coccinea http://www.stanford.edu/~petelat1/ani5.jpg Hemignathus virens Telespiza cantans http://www.uatuahine.hawaii.edu/hana/NaManu/iiwi/iiwi.jpeg Vestiaria coccinea Extreme isolation limits vertebrate fauna No native amphibians 6 7 7 7 6 27 19 11 11 1 mammal (bat) Oreomystis bairdi 15 mtDNA RFLP 8 1 bird established and radiated into 33 extant and 14 extinct species! 9 10 No native reptiles Hemignathus parvus http://www.stanford.edu/~petelat1/hcreepr1.jpg http://www.oahunaturetours.com/ images/laysanfinch.jpg Palmeria dolei Honeycreeper Cladogram 11 11 17 8 23 11 19 29 33 26 37 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 Is this a phylogram, a cladogram, or an area cladogram? 28 7 How about Plants? Hibiscadelphus How about Plants? Hibiscadelphus Hibiscadelphus distans Hibiscadelphus distans fruit pr og re ss distans Kauai io wilderianus Maui n crucibracteatus Lanai giffardianus Hawaii hualalaiensis Hawaii bombycinus Hawaii adaptive radiation http://www.flickr.com/photos/ 53189052@N08/8236595322/ Could Velcro-like seeds be a protoadaptation for hitchhiking via bird feathers? Seeds are favored as food by introduced rats! The phylogram is also an area cladogram showing the positive correlation between evolution and oceanic island biogeography. This plant is critically endangered (pollinator: honeycreepers susceptible to malaria from exotic birds). 29 Silverswords http://biology.usgs.gov/s+t/lrgimage/t172w01.jpg http://hbs.bishopmuseum.org/good-bad/images/iliau.jpg Photo Source: Unknown…If yours, please send email to koningre@gmail.com Kauai-11 endemic Oahu-2 endemic Wilkesia gymnoxiphium Dubautia paleata Maui Nui-7 endemic Argyroxiphium sandwicense Hawaii-3 endemic N http://eol.org/data_objects/17602931 30 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 Plus 5 indigenous species http://www.botany.hawaii.edu/faculty/carr/images/dpal_chl.jpg 31 32 8 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 Sanicula gregaria of California/ Oregon 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 The long distance path probably argues for the sticky surfaces of “tar” weeds as the protoadaptation for hitch-hiking…on what? http://www.insectimages.org/images/ 384x256/5459645.jpg http://www.calflora.net/bloomingplants/images/ redrocktarweed4.jpg Silversword Cladogram (simplified) http://www.stanford.edu/ ~rawlings/kengif/jpeg/ tarweed.jpeg Silversword Cladogram (simplified) 33 For further reading: 34 http://www.sherwincarlquist.com/tarweed-silversword.html Guess how this plant got to Hawaii from Alaska/Oregon? Sanicula mariversa of Hawaii (endemic) Viola kauaensis endemic to Kauai Viola of Hawaii (open fruit with seeds) http://www.flickr.com/photos/49147273@N04/5106688086/ http://www.botany.hawaii.edu/faculty/carr/ images/san_mar_cu_555.jpg 35 http://www.botany.hawaii.edu/faculty/carr/images/vio_kau_fl.jpg http://www.botany.hawaii.edu/faculty/carr/images/ vio_cha_tra_regan_ritchie.jpg 36 9 http://www.botany.hawaii.edu/ faculty/carr/images/ger_arb.jpg Not all cladograms tell the same story! Unlike anywhere else on earth…Hawaiian violets are woody! An adaptation to year-round warm temperature? Another example of adaptive radiation across the archipelago! http://www.botany.hawaii.edu/ faculty/carr/images/ ger_cun_cu.jpg 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 http:// www.botany.hawaii.edu/ faculty/carr/images/ kauaiense Kauai bog ger_han_cu.jpg Geranium colonized “late” E. Maui and radiated and migrated, speciating most recently on Kauai with ecological correlations! http://www.botany.hawaii.edu/faculty/carr/images/vio_lan_mid.jpg 37 humile E. Maui capillare W. Maui Tetramolopium remyi W. Maui, Lanai colonized “late” E. Maui and radiated and rockii Molokai sylvae Molokai, Maui migrated to the other islands tenerrimum Oahu arenarium E. Maui consanguineum Hawaii conyzoides Maui Nui, Hawaii lepidolum E. Maui, Lanai, Oahu Geranium cuneatum 38 Geranium hanaense For Further Reading on Hawaiian Plant Origins Baldwin B. G. and W. L. Wagner. 2010. Hawaiian angiosperm radiations of North American origin. Ann Bot. 105(6): 849-879. www.ncbi.nlm.nih.gov/pmc/articles/PMC2876002/ http://www.botany.hawaii.edu/faculty/carr/images/tet_hum.jpg Not all cladograms tell the same story! Tetramolopium humile 39 40 10