Craig’s research centers on the use of molecular approaches to study ecology and evolution and addresses questions including; (1) the use of molecular markers to infer current and historical population processes at various spatial and temporal scales; (2) the effects of historical changes in habitat on current distributions and diversity of faunas, with particular reference to rainforest biotas; (3) improving the use of molecular information in conservation biology and the development of strategies that recognize evolutionary processes. The geographic focus of this research program spans the tropical forests of Australasia, especially the wet tropics of northeastern Australia and western North America

1. Impact of past and future climate change on diversity in tropical rainforests Evolutionary biogeography: historical biogeography & speciation processes Using knowledge of responses to past climate change to help predict the future Craig Moritz UC Berkeley

2. The problem - it is likely to get hotter than within the last 1 Myr [+ habitat loss + pathogens +…] Disappearing climates are concentrated in the montane tropics…. Rovito et al. 2009 PNAS … and we are losing sensitive species from cloud forests Williams et al. 2007 PNAS

4. Extraordinary endemism of rainforest reptiles and frogs (Moritz et al. 2005) + PNG montane 10 4 56 47 24 11 49 33 CY: 189 Kha NEQ: 791 kha MEQ: 173 Kha SEQ/NSW: 436 Kha Laura Gap

5. COOKTOWN CAIRNS TOWNSVILLE The Australian Wet Tropics Rainforest : 0.1% land area, but richest and most endemic region of Australia; World Heritage Area on basis of ancient, diverse biota 1 1 Yeates et al. 2002

6. Modelled distribution of rainforest under variable Pleistocene/Holocene climates predicts refugia Pollen record: Kershaw et al. 2005 Spatial modeling: (Graham et al. 2006; PNAS) BMC

7. Effects of past climate change on habitat distribution and diversity Past Multiple refugia Past Single refuge Current High diversity & local endemism High diversity Low diversity

8. Macroecology suggests late Quaternary rainforest contraction => local extinction 1. Irregularly shaped subregions have fewer endemic species (Williams & Pearson 1997) 2. More stable subregions have more endemic species (Graham et al. 2006)

9. Fine-scale phylogeography matches modelled refugia in a rainforest snail Hugall et al. 2002, PNAS

10. Paleomodel predictions and phylogeography for montane vs broad-niche Saproscincus skinks Warm-wet restricted Cold-dry restricted S. basiliscus : Broad-niched - high persistence, high diversity S. czechurai : Narrow-niched - low persistence & diversity Moussalli et al. 2009

11. A whole bunch of trees later…

12. Variable responses to past climate change Hugall et al. 2002 PNAS Phylogeographic supertree; Cameron et al, in review

13. Species hotspots and predicted Pleistocene refugia in Amazonia The refuge speciation model: Haffer 1969 Did this long history of climate-induced contraction of rainforests drive speciation as well as extinction?

14. Most endemic vertebrates are old (Miocene) species with sister taxa outside the wet tropics Musky rat kangaroo Golden bowerbird Carrot-tailed gecko Phylogenetic relics Sister taxa in PNG, SEQ Chowchilla Green-eyed tree frog Leaftail gecko Green & lemuroid ringtail possums Within WT speciation: Barred frogs (3) Ringtail possums (2) Tree kangaroos (2) Cophixalus frogs (11)

15. Extreme local endemism: 7/11 WT species of Cophixalus are restricted to single mountain-tops Hoskin 2004

16. C. zweifeli (Melville) 99 100 99 100 100 100 95 <59 100 99 100 99 92 100 <59 96 100 100 C. ornatus (sthn) C. ornatus (lowland) C. ornatus (nthn) C. infacetus (central WT) C. neglectus (Mt Bartle Frere) C. neglectus (Mt Bellenden Ker) C. mcdonaldi (Mt Elliot) C. crepitans C. aenigma (Mt Lewis) C. aenigma (Thornton Pk) C. exiguus (Big Tableland) C. bombiens (Windsor Tbld) C. saxatilis (Black Mtn) C. concinnus (Thornton Peak) C. monticola (Mt Lewis) C. hosmeri (Mt Lewis) A. pluvialus (central WT) A. robusta (sthn WT) A. fryi (nthn WT) A. gracilipes (Cape York) These species are very old (mid-Miocene to Pliocene?), but sister taxa in northern clades are adjacent (Hoskin, 2004). N WT narrow endemics Sth WT (& Melville) CY 0.05 substitutions/site 0.052 0.001 0.010 0.015 0.029 0.018 0.026 0.038 0.038 0.020 0.075 0.069 0.025 0.062 0.008 0.008 0.058 0.133 0.008 0.017 0.012 0.009 0.033 0.002 0.023 0.040 0.050 0.058 0.012 0.018 0.032 0.048 0.019 0.010 0.043 0.048 0.042 0.100 84% 99% 88% 78% 58% 26% 46% 80% 88% 85% 100% 67% 100% 100% 88% 78% 84% 100%

17. Deeply divergent phylogeographic lineages are morphologically conserved: eg. lizards across the BMC (MANOVA, p) Character G. queenslandiae C. laevis S. cornutus Snout-vent 0.253 0.645 0.104 Hind limb 0.553 0.937 0.134 Front limb 0.493 0.907 0.999 Gape width 0.412 0.995 0.119 Head length 0.466 0.815 0.657 Wilk's lambda 0.632 0.383 0.521 Mt Seq divergence: 7% 5% 8% Schneider & Moritz 1999 Proc. Roy. Soc Lond B

18. A more nuanced view of refugial divergence? Refuge model: Haffer 1969 Vanishing refugia: Vanzolini & Williams 1981 AWT forests: now vs modeled refugia (VanderWal et al. 2008) Pre-European Late Q refugia

19. Phenotypic divergence in microrefugia: the (nearly) vanishing refuge model 1. Cophixalus ornatus (Hoskin et al. sub.) LGM paleomodel Multilocus phylogeography Eco-morphology 2. Lampropholis robertsi (Bell et al., in prep.) TU CU AU

20. The Wet Tropics Suture Zone : a natural laboratory for comparative studies of speciation Gnypetoscincus queenslandiae Litoria genimaculata , L. rheocola , Saltuarius cornutus Carlia rubrigularis, Lampropholis coggeri, Gnarosophia bellendenkerensis, Austrochaperina fryi/robusta Litoria nannotis, G. bellendenkerensis Carphodactylus laevis Bettongia tropica, L. genimaculata Litoria nannotis Carphodactylus laevis Hypsilurus boydii Eulamprus tigrinus Saproscincus czechuri Nyctimystes dayi, Glaphyromorphus fuscicaudis Cophixalus ornatus Cophixalus exiguus/aenigma Temnoplectron reyi/politulum Saproscincus lewisi/basiliscus Moritz et al. 2009 PRSL <8Kya expansion Hybridization Admixture for mtDNA Parapatry for mtDNA

22. Mitochondrial DNA ND4 : 500 bp Is there evidence for RI at secondary contacts?: Phylogeography of Carlia (Dolman & Moritz, 2006) N. Carlia rubrigularis S. Carlia rubrigularis Carlia rhomboidalis Rhom. 11.38 % 7.15 % North Rub. South Rub. Secondary Contact

24. Broad-scale phylogeography of L. genimaculata M. Cunningham PhD thesis

25. Cairns  Rapid allopatric speciation via reinforcement in the green-eyed tree frog (Hoskin et al. 2005, Nature) Incomplete isolation N <-> S Rapid evolution of prezygotic isolation between iS and S via reinforcement in iS/N contact zone N N N N N N N N N isolated N isolated N N N N N N L. genimacualta PNG 0.105 0.052 0.067 S S S S S isolated S isolated S isolated S S S S S S S S S S Tinaroo Dam Barron River LAMB RANGE Davies Ck Clohesey’s River Northern Haplotype Southern Haplotype N Rainforest Sclerophyll iN iS N S                                                         Mixed    Jum Rum Ck. transect Shoteel Ck. transect 6km

26. Narrow endemism of species & lineages is concentrated in coastal refugia Species & lineages Lineages only Suture zone Conservation values: Moritz et al. 2001, Moritz 2002 Herp species Herp lineages

27. Response to late Quaternary climate change has increased sensitivity to future climate change D. Hilbert, CSIRO Predicted declines in species richness (Williams et al. 2003) warming Today 2 o C warming

28. Predicted loss of montane endemics, e.g. ( C. neglectus )

29. Predicted loss of range in a widespread species ( C. ornatus ) Species persists, but major loss of phylogeographic and phenotypic diversity and of hybrid zones

30. But even well-fit models can fail to predict persistence under past climate change Eg. distinct lineages of S. basiliscus; Moussalli et al. 2009

31. Perhaps unexpected persistence reflects variation in physiology across lineages within species. (Langham et al. in prep.) B95 range CTmax CTmin Region

34. Improving biodiversity prediction in tropical hotspots (Carnaval et al. 2009 Science) Map predicted stable areas (putative refugia) distribution of genetic diversity Descriptive phylogeography timing of demographic processes Population genetic tests congruence across taxa Assemblage-scale hypothesis testing Spell out spatially-explicit hypotheses Re: Sample in stable and unstable areas Model validation Hypothesis formulation Diversity distribution modeling

35. Paleomodels predict endemism in Atlantic Forest (Carnaval & Moritz 2008 J. Biogeog.) Pre-European Atlantic Forest Predicted late Quaternary stability

36. Carnaval et al. 09 Science MtDNA data agree with modeled responses to Late Quaternary climate change Phylogeography of Brazilian Atlantic rainforest frogs

37. H 1 : long-term persistence ( θ  ) 1 ( θ  ) 2 H 2 : recent colonization ( θ  ) 2 ( θ  ) 1 θ 2 θ 1 θ 2 θ 1 Population 1 Population 2 Population 1 Population 2  2 (20 kybp)  1 (120 kybp – 1.2 Mybp) Carnaval et. al. 2009. Science. Population expansion into unstable areas in the Atlantic rainforest Testing for “assemblage-wide” responses to climate change with ABC Where Z 2 = # of species evolved under H 2

38. Conservation implications of a hotspot within a hotspot Stable central corridor functioned as a large refuge for biodiversity Yet, it is poorly known and under much higher threat relative to the unstable south 1500 today

39. “ As buds give rise by growth to fresh buds, and these, if vigorous, branch out and overtop on all sides many a feebler branch, so by generations I believe it has been with the great tree of life, which fills with its dead and broken branches the crust of the earth, and covers the surface with its ever branching and beautiful ramifications” (Darwin 1857)

40. Soooo.. many to thank! Mr February What’s with the blue shirts?