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Deconstructing Diversity Starting Out, Getting There, Staying Alive

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Deconstructing Diversity Starting Out, Getting There, Staying Alive Deconstructing diversity starting out, getting there, staying alive Kartik Shanker Centre for Ecological Sciences Indian Institute of Science Bangalore What causes diversity ? Why do some areas within the tropics have greater diversity? Biodiversity hotspots Wiens 2011 Cracraft 1985 Map: Conservation International Diversity in clades Clade - group composed of ancestor and all its descendants Why is the diversity of some clades greater? > 200 million years old & 2 species Alfaro et al. 2009 Image:http://reptilis.net Diversity in time and space Evolutionary radiations Pratt 2005 Seehaunsen 2006 Jonsson et al. 2012 Givnish 2010 Losos 2009 Connecting diversity in space and in radiations Diversity in space/hotspot = Summation of patterns among clades + Pratt 2005 biogeographic processes (dispersal) Seehaunsen 2006 Givnish 2010 Cracraft 1985 Losos 2009 Todays talk: diversification in the Western Ghats ➢ An evolutionary biogeography perspective of diversity ➢ Starting out: an evolutionary perspective ➢ The challenge of delimitation ➢ Understanding evolutionary origins ➢ Getting there and staying alive: a macroecological view ➢ Staying alive: factors influencing persistence ➢ Getting there: the role of dispersal ➢ Combining environment and range ➢ The road from distribution to diversity: a brief synthesis Determinants of species range Climate Environmental Topography variables (barriers) Species geographic range Species- Inter-specific specific associations traits Determinants of species richness Environmental variables Species Richness Range Area related factors General conceptual framework Environmental variables Species Species- Inter-specific geographic specific associations range traits Species richness General conceptual framework Persistence (Staying Alive) Persistence Species Dispersal (Staying Alive) geographic (Getting there) range Species richness Synthetic framework Origins Dispersal (Starting Out) (Getting there) Species Biotic factors Abiotic factors geographic (Staying Alive) (Staying Alive) range Species richness Two detective stories The Linnaean shortfall The Wallacean shortfall Bush frogs - Raorchestes Wrinkled frogs - Nyctibatrachus Large scale sampling framework • Capture topographic, bioclimatic and geological heterogeneity • Massifs - elevational range from 0 to 2650m • Delimited 14 mountain complexes >1400m • Sampled most of the massifs (over a 5 year period) Palghat Gap • Across available elevational gradients Elivalmalai 2600 Anaimalai Nilgiri Camels Camels Hump Agasthya- Kottamala Kudremukh and Bababudan 1800 malai Thadiyendamol Nishani Pushpagiri Kurichiarmala Brahmagiri Nelliampathy Elevation>1400 m 1000 PalghatGap Elevation (m) South 1000 km North Site selection and sampling methods Across broad elevational bands (vegetation based) Nilgiri Anaimalai 2000 m High Nilgiri 1400 m Mid 700 m Anaimalai Low No. of sites = 250 No. of points = 20,000 Elevational range =100 - 2650m Lineage delimitation : A step wise multi- criteria approach Step 1: Haplotype phylogeny: ML Tree • Combination of genetics, geography and morphology. Step 2: Bayesian Poisson Tree • ML Tree serves as input for bPTP which estimates process the number of putative lineages. • Each lineage is then treated as a hypothesis and Step 3: Pairwise genetic distance evaluated using pairwise genetic distance, geographical overlap and morphology. • Sister lineages with ‘high’ divergence treated as Step 4: Geographic range overlap independent lineages/species • Remaining pairs examined for morphological Step 5: Morphological distinctness distinctness or geographical separation Raorchestes Nyctibatrachus CESS072 CESS156 P1 P3 M G CESS111 CESS170 CESS277 CESS158 CESS015 CESS288 L 4 CESS256 CESS066 L 6 CESS517 CESS527 CESS513 CESS512 CESS190 CESS070 L 7 CESS276 CESS108 CESS167 * CESS226 CESS262 CESS172 CESS223 Ahaetulla CESS404 L 5 Trimeresurus CESS329 CESS086 CESS183 CESS182 CESS141 * CESS164 CESS275 CESS053 CESS311 CESS154 CESS089 CESS090 CESS515 CESS145 CESS144 L 6 CESS155 CESS088 II CESS087 L 5 CESS525 CESS143 CESS273 CESS114 CESS058 CESS054 CESS059 IV CESS103 CESS166 CESS057 L 3 CESS104 L 3 CESS181 CESS422 CESS421 CESS178 CESS505 L 4 CESS257 Ahaetulla nasuta RAP0517 L 2 Ahaetulla sp. CESS506 CESS074 Ahaetulla sp. CESS211 Ahaetulla sp. CESS244 L 1 CESS101 Ahaetulla sp. CESS528 Ahaetulla sp. CESS521 CESS100 Ahaetulla sp. CESS159 L 7 CESS032 * A. pulverulenta RS-C L 8 A. nasuta FMNH255023 III L 9 A CESS056 A. dispar CESS188 * A. dispar CESS187 CESS285 L10 A. dispar CESS189 A. dispar CESS261 T. gramineus B261 Ahaetulla sp. CESS075 L11 L 1 A. perroteti CESS286 CESS147 L12 A. perroteti CESS287 CESS369 A. fronticincta CAS245684 A. fronticincta CAS222635 L13 CESS378 A. fronticincta CAS204966 Ahaetulla sp. CESS347 CESS033 Ahaetulla sp. CESS340 L14 Ahaetulla sp. CESS260 L 2 CESS272 A. prasina APR2 L15 V CESS040 * A. prasina APR1 * A. prasina LSUHC8586 L16 T. trigonocephalus A. mycterizans LSUHC8913 L20 A. mycterizans ADM001 T. strigatus CESS142 A. prasina FMNH269042 L17 I A. prasina KU302954 L18 T. borneensis A. prasina KU326673 L19 A. fasciolata LSUHC8837 T. puniceus L21 A. fasciolata LSUHC9831 B Other Trimeresurus Ahaetulla sp. CESS318 L22 Ahaetulla sp. CESS259 OUTGROUP P1 P3 G M GD OUTGROUP Outgroup 0.07 Wallace meets Linnaeus and Darwin Allopatric Sympatric Parapatric Identifying sister lineages with confidence Micrixalus Nyctibatrachus Raorchestes Known lineages Putative new lineages New lineages across levels of divergence in all taxa 2 h 2 2 g i H 2 n o i t a r a p e S l a c i g o l o h Genetic divergence p r o M Shallow 1 1 Moderate w o High 1 L Allopatric Geographical distribution Sympatric 1 Parapatric Lineages in multi-dimensional space Geography Allopatry Sympatry Parapatry ? Morphology overlap distinct overlap distinct overlap distinct L 7 1 0 13 exclusive2 lineages ? ? 3 Genetic M distance 16 exclusive lineages H General lineage concept Not so general lineage concept Morphological, genetic and geographical separation can occur at different rates and in different sequences, even within the same clade Vijayakumar et al (2016) Proc B Shanker et al (2017) J Genetics Starting out: an evolutionary perspective Peninsular India 35 Calotes Trimeresurus Ahaetulla Raorchestes X North clade 25 X M Peninsular India 67% 33%U CLOSED CANOPY( OPEN CANOPY 40%) (60%) Palghat Gap Refuge C 3% 35 SC 27% M 33% South clade 37%U CLOSED CANOPY OPEN CANOPY (77.5%) (22.5%) Trimeresurus Ahaetulla Raorchestes Getting there and staying alive: a macroecological view Gradients in Species Richness MECHANISMS PATTERNS Temperature Latitudinal Precipitation Area Altitudinal Productivity Bathymetric Historical factors Rapoport’s rule Mid Domain Effect Environmental factors • Solar energy • Solar radiation and altitude/latitude • Temperature (Range, mean, maximum, minimum) • Wind and Rainfall • Precipitation • Water stress • AET – Actual evapotranspiration • PET – Potential evepotranspiration • Wind patterns (global and local scales) • Soil • Productivity (Biomass, Abundance) The mid domain effect • Geometric constraints (d) • Range size and position random 70 60 • Range limits constrained by 50 boundaries 40 30 • Alternate models with empirical Speciesrichness 20 10 range size distributions or empirical 0 0 2 4 6 8 10 12 range mid-points Elevational zone Colwell and Hurtt, 1994, Colwell and Lees, 2000 Combining range and environment ➢ Environmental patterns ➢ Range based explanations •Mid domain effect ➢ Range cohesion and range scatter models ➢ Modeling the distribution of organisms based on the assumption that ranges are contiguous (as one would expect from dispersal) and that they are not ➢ Using both frameworks with occurrence probabilities based on environmental parameters Rahbek et al 2007 Range cohesion models Range scatter models Implication: everything is everywhere (Bejerinck's law) A cohesion – temperature model Model that combines range overlap and temperature provides best visual fit to empirical data for many taxa Frog and Lizard richness maps for the WG Bird richness map for the WG 60 - 97 Tree richness map for the WG Range Cohesion in an empirical grid of WG Species RichnessSpecies Species Richness Species Feasible ranges and Linear Temperature Gradient Summary • Using a spatially explicit biogeographic approach for discovery and delimitation of lineages forms the basis of further analysis • A replicable step-wise multi-criteria approach for delimitation • Combining range, environment and evolution is critical for understanding diversity • Ecological dispersal plays a greater role in some groups (birds, some plants) • Diversification plays a greater role in some groups (frogs, some plants) • Significant role of both in some groups (lizards, snakes, some plants) • In-situ diversification of many clades in Peninsular India, with Western Ghats serving as a hotspot; different models of diversification Researchers and collaborators PROJECT researchers: PROJECT collaborators: SP Vijayakumar (PhD, frogs) KP Dinesh (Postdoc, frogs) Varad Giri (Lizards) Varun Torsekar (PhD, frogs) Aniruddha Dutta Roy (Skinks) Ashok Kumar Mallik (PhD, snakes) Ishan Agarwal (Geckos) Sneha Vijaykumar (PhD, birds) P. Gowri Shankar (Snakes) Navendu Page (PhD, plants) Deepak Veerappan(Lizards) Saunak Pal (Lizards and Snakes) Xavier (Canopy climbing) Mrugank Prabhu (Frogs) SR Chandramouli (Frogs) Mixed species research: Achyuthan Srikanthan (Snakes) Vijay Ramesh (Frogs) Hari Sridhar, Anne Theo, Gaurang Gowande (Lizards) Priti Bangal, Priyanka H. Anisha Jayadaven (Models) Riya Menezes, Aditi Jayarajan, Priyanka Swamy (Molecular genetics) Mayavan (Field Assistant) PROJECT SUPPORT • CEPF • INDIAN INSTITUTE OF SCIENCE (IISc) • ATREE • Ministry of Environment and Forests • Kerala Forest Department • Karnataka Forest Department • Tamil Nadu Forest Department • Maharashtra Forest Department • FIELD AND EXPEDITION Members Thank you.
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