The California Hotspots Project: I.
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1 The California Hotspots Project: I. Identifying regions of rapid diversification of mammals Ed Davis, M. Koo, C. Conroy, J. Patton & C. Moritz Museum of Vertebrate Zoology, UC Berkeley *Funded by Resources Law Group & CA State Parks *Thanks to J. Grinnell & colleagues, T. Smith, V. Sork, D. Ackerly, T. Barnosky, D. Wake, A. Vandergast, A. Bohonak. Goal: To maintain evolutionary processes and the viability of species and functional landscapes necessary to achieve this. Moritz 2002 (After Frankel (1974))
2 A policy-driven challenge to evolutionary biologists Active process of land acquisition for conservation in CA: CSP, TNC & others: $0.5B last 5 yrs for purchase Prop 84: $6B Get it right, make it count.. Include Evol Processes as a major factor in planning to assist resilience Rayburn - take a bow! Where are the areas of active diversification: Evolutionary hotspots?
3 Environmental surrogates for evolutionary processes: South Africa (Cowling et al.) Diversification across lowland-upland gradients, edaphic gradients, macroclimatic gradients
4 Neo-endemism of CA plants (polyploid derivatives) Stebbins & Major 1965 Hypothesis: Elevated neoendemism in regions with recent (Plio-Pleistocene) development of full Mediterranean climate AND not subject to recent glaciation (also Raven & Axelrod 1978)
5 Endemism-richness for species vs lineages (Rissler et al. 2006)
6 Key evolutionary processes Generative processes Adaptive diversification Abiotic gradients (climate, soils) Biotic interactions -> character displacement, (eg. novel communities, suture zones) Non-adaptive diversification Genetic drift Opportunity Niche space Genetic isolation Stability
7 What biological data do we have/need to map Evol hotspots? Geographic distributions & phylogeny (species, phylogeographic lineages) Hotspots for short-branch taxa [neo-endemism] Hotspots of phylogeographic endemism Suture zones Spatial information on ecotypic differentiation (phenotypes) Subspecies endemism as a surrogate Today: mammals only
8 Subspecies as surrogates for geographic patterns of phenotypic (adaptive?) divergence Skepticism from some systematists May not reflect historical lineages (good) May be artefacts of inadequate sampling/analysis of clines Variable criteria for recognition (bad) BUT, evolutionists (eg. Grinnell, Mayr) have long used subspecies to describe geographic patterns of phenotypic variation and view subspecies as intermediates in the speciation process Diverse CA taxa: Dipodomys (30) Tamias (16) Thomomys (16) Microtus (12) Chaetodipus (12)
9 General approach Select (near) endemic taxa (>75% range in CA) 26 species 130 subspecies Estimate 1km resolution (MVZ points -> range-map-clipped MaxENT models) Richness maps mtdna distance to sister species (GenBANK, unpubl) Endemism-richness =!"1/area) per cell Species neo-endemism richness =!"(1/area) X 1/distance) per cell
10 Endemic species - richness, endemismrichness & neo-endemism !"1/area)!"1/area)(1/distance) Note: island endemics excluded 1. SF Bay area: R. raviventris 2. Interior Coast Ranges: D. ingens, D. nitratoides, A. nelsoni 3. Tehachapi: P. alticola 4. Central Sierra: S. lyelli, T. alpinus 3,4 = recent lineages cf. 1,2 = old; Note: north/central coast
11 Endemic species: foci of recent vs old taxa (not 1/area weighted) Richness Neo-Richness -! (1/distance) Paleo-Richness! distance Central coast, Tehachapi, Central Sierra are foci Essentially no difference to richness; ie random
12 Richness & endemism-richness of endemic subspecies North coast, SF Bay, Tehachapi, San Bernadino, CV, Owens Valley, Central coast
13 Neo- and subspecies endemism (top 10%) & comparison with CA/Fed Protected Areas Species & subspecies neoendemism areas distinct except Tehachapis, St. Lucia, San Bernadino Ranges Environmental correlates: coastal gradients, biogeographic overlap areas (gradients) Good potential for integrated reserve design in most areas (except Nth coast?). Note - other taxa & approaches needed (eg. Sork, Vandergast/Bohonak etc.)
14 MVZ Grinnell Project: change over 100 yrs Across Yosemite transect, many small mammal show upwards range shifts of m since 1920 (with c. 3C increase in monthly minimum temperature)
15 Management & policy implications Specific - don t get excited yet - more taxa/approaches to come! General: Rates of diversification are not randomly distributed - cold & hotspots Common theme is gradients/ecotones, stability? Location of hotspots likely to shift with climate Protection of EHs requires large-scale and coordinated planning to protect connectivity across gradients, and large, heterogeneous isolates
16
17 Caveats Spatial errors in distribution estimates Minimized by combining models & point estimates Error in inferring relative divergence time from mtdna distance ancestral coalescence, introgression etc.) - likely overestimating divergence times; adds noise Vagaries of subspecies classifications But most from Grinnell (or his students) so relatively consistent approach Current patterns of neo-endemism may reflect environmental, but not geographic space of origin (eg. late Q range shifts)
18 Components of Hotspots Project Evolutionary theory: Selection, isolation, hybridization etc. Conceptual model: => environmental predictors; e.g topographic, climatic and geological complexity, ecological stability Spatial layers for environmental predictors Build & test Spatial patterns of diversity (genetic, phenotypic, phylogenetic, species): Mammals, amphibian, reptiles, birds, plants Spatial prediction of rates of evolution => hotspots Apply Conservation values; gap analysis to identify priority areas for acquisition
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EVOLUTION Evolution - changes in allele frequency in populations over generations. Sources of genetic variation: genetic recombination by sexual reproduction (produces new combinations of genes) mutation
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