BMC Evolutionary Biology

Comparative phylogeography of arthropod communities through the lens of an island chronosequence Famous for adaptive radiation … But not all radiations are created equal

Only by understanding the differences,

&2018 © IIthe Joint Congress conditions on Evolutionary Biology 2018. All rights reservedcreating - Any reproduction even in them,part is prohibited. will we ever understand the process of adaptive radiation

Rosemary Gillespie, Univ California,2018 © II Joint CongressBerkeley on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. nature.berkeley.edu/evolab2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Hormiga, Arnedo & Gillespie 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2003. Systematic Biology201852: 70-88 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

No. species/ No. species/ area 0 Hawaii Maui Nui Oahu Kauai Orsonwelles

O. torosus O. calx 1 O. calx 2 O. calx 3 O. iudicum O. bellum O. ventus 1 O. ventus 2 Classic “non O. ventus 3 O. malus 2 O. malus 3 adaptive” radiation O. malus 1 O. arcanus 1 O. arcanus 2 O. ambersonorum 1 O. ambersonorum 2 O. polites 1 Similar stories in 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. O. polites 2 O. polites 3 • Laupala crickets O. othello 3 O. othello 2 O. othello 4 (Shaw et al) O. othello 1 O. macbeth 5 O. macbeth 4 O. macbeth 2 O. macbeth 3 O. macbeth 1 • Silverswords O. falstaffius 1 O. falstaffius 2 O. falstaffius 3 (Baldwin et al) O. graphicus 1 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. O. graphicus 2 Gillespie 2016, Evolutionary2018 Applications © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . M. anguliventris Hawaii M. anguliventris Maui Adaptive radiation M. anguliventris Oahu M. anguliventris Kauai * with early ecological M. naevigera Hawaii M. naevigera Maui diversification M. naevigera Oahu * • Ecological entities then M. discreta Molokai diverge in allopatry M. discreta Kauai * M. cavata Hawaii M. cavata Necker * M. nigrofrenata Hawaii M. nigrofrenata Kauai * M. rufithorax Oahu

M. imbricataOahu

M. facunda Hawaii

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproductionM. arida evenMaui in part is prohibited. M. sp Maui M. kanakana Hawaii Similar story in Hyposmocoma M. kanakana Maui moths M. kanakana Oahu (Rubinoff et al) M. kanakana Kauai * M. spinosa Oahu M. hiatus Maui

M. perkinsi Oahu 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 M.© edita Oahu48th EuropeanGarb & Gillespie, 2009. Molecular Contact Ecology 18:1746–1764 Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Gene-tree (mt Gold – under leaves COI & nuclear EF1 seqs); output by Beast. White – on lichen

2018 © II Joint CongressGillespie on Evolutionary et Biology al. 2018. 2018. All rights reservedCurrent - Any reproduction Biology even in part is prohibited. 2018 © 48th European 28Contact(6), pp.941-947. Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . T. hawaiiensis Oa .. general progression down T. hawaiiensis Evolution of ecomorphs Ha island chain T. kauaiensis Ka T. pilosa Ka T. perreirai Oa T. polychromata Oa T. tantalus Oa T. quasimodo Oa T. quasimodo Ha T. quasimodo Ma T. anuenue Ha Repeated evolution Spiny leg T. kikokiko Ma Tetragnatha T. kukuiki Oa T. obscura Ha T. kukuhaa 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. Ha Phylogeny T. waikamoi Ma based on T. kamakou Ma mtDNA, T. restricta Ha allozymes, & T. restricta Ma minisatellites T. brevignatha Ma T. macracantha Ma 2018 © II Joint Congress on EvolutionaryGillespie Biology 2018. 2004All rights reservedScience - Any303: reproduction 356 -even359 in part is prohibited. T. brevignatha 20180.1 © 48thHa EuropeanPons & Gillespie Contact 2004 J Mol Evol Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Argyrodes trigonum T. hawaiiensis Ha A. huinakolu Kauai T. kauaiensis Ka A. kahili Kauai T. pilosa Ka A. uwepa Oahu T. perreirai Oa T. polychromata Oa A. sp (white) Oahu T. tantalus Oa A. makue Oahu T. waikamoi Ma A. sp (black) Molokai T. kamakou Ma A. sp (gold) Molokai T. restricta Ha A. sp Molokai T. restricta Ma (like melekalikimaka) 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. T. brevignatha Ma A. poele Molokai T. macracantha Ma A. melekalikimaka Maui T. brevignatha Ha A. laau Maui T. quasimodo Oa A. alepe Maui T. quasimodo Ha A. corniger Maui T. quasimodo Ma T. anuenue Ha A. waikula Hawaii T. kikokiko Ma A. hiwa Hawaii T. kukuiki Oa 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. Rhomphaea metaltissima Ariamnes attenuata T. obscura Ha Gillespie 2016, Evolutionary Applications Spintharus flavidus 2018 © 48th European T.Contact kukuhaa Ha Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . Insights from chronosequence Can the apparent “overshoot” Ariamnes 4,5 provide information on early stages 4 of the diversification process? 3,5 3 2,5 2 1,5 1 0,5

No. species/ area species/ No. 0 Hawaii Maui Nui Oahu Kauai Youngest -- Island age -- Oldest

Tetragnatha – spiny leg clade

All in 3

habitat 2

1,5

0,5

2018 © II JointHawaii Congress on EvolutionaryMaui Biology Nui 2018. All rightsOahu reserved - Any reproductionKauai even in part is prohibited. 2018 © 48th EuropeanYoungest -- Island ageContact-- Oldest Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . What is the process? Sticklebacks - Divergence in habitat use among closely related species is a common characteristic of adaptive radiations. (Matthews et al 2010. Ecology 91: 1025–1034)

Can the apparent “overshoot” provide information on the early stages of the diversification process?

Ariamnes Tetragnatha – spiny leg clade

4,5 3

No. species/ area species/ No. 0 0

Hawaii Maui Nui Oahu Kauai 2018 © II JointHawaii Congress on EvolutionaryMaui Biology Nui 2018. All rightsOahu reserved - Any reproductionKauai even in part is prohibited. Youngest2018 -- Island ©age 48th -- Oldest EuropeanYoungest -- Island ageContact-- Oldest Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Taxa used in current study (green ecomorph) T. waikamoi range T. brevignatha range T. macracantha range

Related ecomorphs T. kamakou T. restricta

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. Cotoras et al. 20182018 BMC Evolutionary © Biology48th18(1):100 European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . Hawaiian Tetragnatha - Genetic Structure of Maui & Hawaii Spp of Green Ecomorph - based on exon capture

Lanai T. macracantha East Maui W Maui Why so many of the green ecomorph on this T. brevignatha one volcano? T. waikamoi East Maui East Maui

T. brevignatha

SplitsTree 4.13.1 (Huson and Bryant, 2006)

Darko Cotoras 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. Cotoras et al. 20182018 BMC Evolutionary © Biology48th18(1):100 European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Taxa used in current study (green ecomorph) T. waikamoi range T. brevignatha range T. macracantha range

Related ecomorphs T. kamakou T. restricta

1. Allopatry ➔ genetic isolation 2. Secondary sympatry ➔interaction 3. Subsequent ecological divergence allows “stable co-occurrence”2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . Interplay between isolation & host shifts in generating diversity within single species, Nesosydne umbricata • Host generalist, on unrelated hosts Clermontia and Pipturus • What is relative importance of geography and host in structuring populations in the early stages of differentiation. Photo: Karl Magnacca

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - AnyKari reproduction Goodman even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Kari Goodman Stefan Prost Nesosydne planthoppers2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Parallel radiation Big Is Clermontia & Cyanea on Pipturus

Radiation on Clermontia

Photo: Karl Magnacca 1. Allopatry ➔ genetic isolation ➔ 2. Secondary sympatry interaction2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 3. Subsequent2018 ecological © 48th divergence European ➔co-occurrence Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] .

2018 © II Joint Congress different very on be can Processes but process;the Evolutionary intoinsight Provide Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . Fitness 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] . – all:

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. entire communityentire fromthe insightsneed accumulates,biodiversity Tohow understand Community Aassembly Across the Chronosequence Metabarcoding

floor

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th EuropeanHenrik ContactKrehenwinkel Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

Identity & abundance Invasion impact & of all members vulnerability

0.7-1.8 my

Krehenwinkel et al. 2018. PloS one 13: e0189188. Krehenwinkel et al. 2018. Methods in Ecology and Evolution, 8(1): 126-134 2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. Krehenwinkel et al. 2017 Scientific Reports 7, 17668. 0-0.4 my Krehenwinkel et2018 al. 2017 Methods in ©Ecology 48th and Evolution. DOI:10.1111/2041European-210X.12647. Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .

2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited] . Daniel Gruner, John Harte, Neo Martinez, Rasmus Nielsen, Patrick O’Grady, Diana Percy, Donald Price, Dan Rabosky, George Roderick, Kerry Shaw Michael Brewer, Darko Cotoras, Curtis Ewing, Kari Goodman, Karl Magnacca, Andy Rominger, Elske Tielens UC Berkeley, U Maryland, Cornell, UH Hilo

nature.berkeley.edu/hawaiidimensions2018 © II Joint Congress on Evolutionary Biology 2018. All rights reserved - Any reproduction even in part is prohibited. 2018 © 48th European Contact Lens Society Of Ophthalmologists. All rights reserved - Any reproduction even in part is prohibited] .