evolutionofendemismonayoungtropicalmountain vincent s. f. t. merckx1,2, kasper p. hendriks1,3, kevin...

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Evolution of endemism on a young tropical mountainVincent S. F. T. Merckx1,2, Kasper P. Hendriks1,3, Kevin K. Beentjes1, Constantijn B. Mennes1, Leontine E. Becking1,4,5,Katja T. C. A. Peijnenburg1,6, Aqilah Afendy7, Nivaarani Arumugam7,8, Hugo de Boer1,9,10, Alim Biun11, Matsain M. Buang11,Ping-Ping Chen1, Arthur Y. C. Chung12, Rory Dow1, Frida A. A. Feijen1, Hans Feijen1, Cobi Feijen-van Soest1, Jozsef Geml1,2,Rene Geurts13, Barbara Gravendeel1,2,14, Peter Hovenkamp1, Paul Imbun11, Isa Ipor15, Steven B. Janssens16, Merlijn Jocque17,18,Heike Kappes1,19, Eyen Khoo12, Peter Koomen20, Frederic Lens1,2, Richard J. Majapun12, Luis N. Morgado1, Suman Neupane21,Nico Nieser1, Joan T. Pereira12, Homathevi Rahman7, Suzana Sabran12, Anati Sawang7, Rachel M. Schwallier1, Phyau-Soon Shim11{,Harry Smit1, Nicolien Sol1, Maipul Spait11, Michael Stech1, Frank Stokvis1, John B. Sugau12, Monica Suleiman7, Sukaibin Sumail11,Daniel C. Thomas1,22,23, Jan van Tol1, Fred Y. Y. Tuh11, Bakhtiar E. Yahya7, Jamili Nais11, Rimi Repin11, Maklarin Lakim11 &Menno Schilthuizen1,2,7

Tropical mountains are hot spots of biodiversity and endemism1–3,but the evolutionary origins of their unique biotas are poorlyunderstood4. In varying degrees, local and regional extinction,long-distance colonization, and local recruitment may all contrib-ute to the exceptional character of these communities5. Also, it isdebated whether mountain endemics mostly originate from locallowland taxa, or from lineages that reach the mountain by long-range dispersal from cool localities elsewhere6. Here we investigatethe evolutionary routes to endemism by sampling an entire trop-ical mountain biota on the 4,095-metre-high Mount Kinabalu inSabah, East Malaysia. We discover that most of its unique biodi-versity is younger than the mountain itself (6 million years), andcomprises a mix of immigrant pre-adapted lineages and descen-dants from local lowland ancestors, although substantial shiftsfrom lower to higher vegetation zones in this latter group wererare. These insights could improve forecasts of the likelihood ofextinction and ‘evolutionary rescue’7 in montane biodiversity hotspots under climate change scenarios.

In mountainous areas of the humid tropics, steep environmentalgradients coincide with high primary productivity and relative cli-matic stability to sustain large numbers of species, often with strik-ing degrees of endemism at higher elevations1,8. It has therefore beenrecognized that tropical mountains are biodiversity hot spots of greatconservation value1, especially because endemics on mountain topsare vulnerable to becoming trapped and then annihilated as a resultof global warming2,3.

The evolutionary origins of these unique biotas, however, are poorlyunderstood4. Like other insular habitats9, the endemic biota of anisolated mountain results from complex dynamics among coloniza-tion, in situ speciation, and local extinction. Each of these factors isdependent on the age and size of the habitat, and on the environmentalcontrast between the insular habitat and its matrix10. In the case ofa tropical mountain top, an added complication is the fact thatclimate fluctuations may have widened and restricted the geographicrange over which the montane conditions have extended in the past,

meaning that parts of the endemic biota may be relicts, and othercomponents may be novel in character4,5.

Disentangling these possibilities for a single tropical montanebiodiversity hot spot requires molecular phylogenetic study of a largenumber of fauna and flora elements. However, with only few excep-tions11,12, evolutionary studies in such hot spots have been limitedto single taxa13,14. This precludes broad understanding of the evolu-tionary and biogeographic origins of an endemic biota as a whole4.

We investigated the evolutionary routes to endemism by samplingan entire tropical mountain biota on the UNESCO World Heritage siteof Gunung Kinabalu in Sabah, East Malaysia. We demonstrate thatmost of its unique biodiversity is younger than the mountain itself andcomprises a mix of immigrant pre-adapted lineages as well as descen-dants from local lowland ancestors.

At 4,095 m, Kinabalu is the tallest mountain between the Himalayasand New Guinea. It is a solitary ‘sky island’, having emerged during thePliocene and early Pleistocene as a granite pluton within the surround-ing sandstone of the Crocker Range, the latter having formed muchearlier, between the Eocene and the early Miocene15. Because of thearea’s tectonic activity, as well as Pleistocene sea level changes, theexact historical progression of its elevation above sea level is notknown, but it is likely that a major rise, even beyond today’s elevation,of Kinabalu, as well as the central spine of the Crocker Range, tookplace between 6 million years ago and today (for more geologicalbackground see Methods). Since the early days of its exploration16,Kinabalu has been famous for its extremely high biological diversity,especially its richness in endemic species, with endemism proportionsreaching 25–30% for some taxa17.

To unravel the origins of the exceptionally rich Kinabalu biota, wemounted a Malaysian–Dutch expedition in which 47 taxonomistsworked at 37 localities, spanning the full range of elevations (Fig. 1).We used Sanger sequencing to sequence one or more fast-evolving locifor 1,852 individuals, belonging to 18 genera representing gastropods,annelids, insects, arachnids, vertebrates, pteridophytes, bryophytes,and angiosperms. We also obtained 3.7 million basidiomycete

{Deceased.

1Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, The Netherlands. 2Institute of Biology Leiden, Leiden University, Sylviusweg 72, 2333 BE Leiden, The Netherlands. 3Groningen Institute forEvolutionary Life Sciences, University of Groningen, Nijenborg 7, 9747 AG Groningen, The Netherlands. 4Wageningen University & Research centre, Marine Animal Ecology Group, PO Box 338, 6700 AHWageningen, The Netherlands. 5Department of Environmental Science, Policy, & Management, University of California Berkeley, 130 Mulford Hall #3114, Berkeley, California 94720, USA. 6Institute forBiodiversity and Ecosystem Dynamics, University of Amsterdam, Science Park 904, 1098 XH Amsterdam, The Netherlands. 7Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah,Jalan UMS, 88400 Kota Kinabalu, Sabah, Malaysia. 8Faculty of Earth Science, Universiti Malaysia Kelantan, Jeli Campus, Locked bag No.100, 17600 Jeli, Kelantan Darul Naim, Malaysia. 9Department ofOrganismal Biology, Uppsala University, Norbyvagen 18D, 75236 Uppsala, Sweden. 10Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, NO-0318 Oslo, Norway. 11Sabah Parks, Lot 45 &46, Level 1-5, Blok H, KK Times Square, 88806 Kota Kinabalu, Sabah, Malaysia. 12Forest Research Centre, Sabah Forestry Department, P.O. Box 1407, 90175 Sandakan, Sabah, Malaysia. 13WageningenUniversity, Department of Plant Sciences, Laboratory of Molecular Biology, 6700AP Wageningen, The Netherlands. 14University of Applied Sciences Leiden, Zernikedreef 11, 2333 CK Leiden,The Netherlands. 15Faculty of Resource Science and Technology, Universiti Malaysia Sarawak, 94300 Kota Samarahan, Sarawak, Malaysia. 16Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium.17Royal Belgian Institute of Natural Sciences, Aquatic and Terrestrial Ecology, Vautierstraat 29, 1000 Brussels, Belgium. 18Rutgers, The State University of New Jersey, Department of Biological Sciences,195 University Avenue, Boyden Hall, Newark, New Jersey 07102, USA. 19Zoological Institute, University of Cologne, Zulpicher Strasse 47b, D-50674 Cologne, Germany. 20Natuurmuseum Fryslan,Schoenmakersperk 2, 8911 EM Leeuwarden, The Netherlands. 21EEB Department, University of Connecticut, 75 N. Eagleville Road, Storrs, Connecticut 06269-3043, USA. 22School of Biological Sciences,University of Hong Kong, Pok Fu Lam Road, Hong Kong, China. 23Singapore Botanic Gardens, 1 Cluny Road, 259569 Singapore, Republic of Singapore.

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and glomeromycete ITS2 rDNA sequences from soil cores with ionsemiconductor sequencing. In addition, we retrieved data from eightpreviously published single-taxon studies on vascular plants.

We analysed all data within a phylogenetic framework to estimatethe times of origin of endemic species, and to determine whetherendemic species had descended from local or distant congeners(Methods, Fig. 2, Extended Data Table 1 and Extended Data Figs 1–9).Note that we define ‘endemic’ as restricted to the area in whichour expedition took place. Although the present study offers themost comprehensive evolutionary analysis of any mountain biota todate, the taxa covered are, by necessity, an uneven and fragmentarysampling of the full diversity. Nonetheless, we expect that ourresults are representative for the Kinabalu biota as a whole, as ourselected taxa encompass organisms with a wide variety of phylogen-etic backgrounds, ecologies, and life history traits.

Similar to Mesoamerican endemic cloud forest seed plants andvertebrates11, our molecular dating results show that the estimatedmean stem-node ages of 33 endemic species span a wide range, from0.12 million years to 14.6 million years (Fig. 3). However, 76% of these

fall within the past 6 million years, the time span during whichKinabalu is likely to have reached its present elevation. Only twoendemics, the frog Kalophrynus baluensis and the flowering plantIlex kinabaluensis, are markedly older than the mountain itself.These may be explained as artefactual if we failed to identify theclosest non-endemic sister lineage, thereby inflating their recon-structed age, or if these species are actually not endemics, but morewidespread. Alternatively, they may truly be old endemics that evolvedduring cooler periods at lower elevations in Borneo before Kinabalu’sformation.

Our phylogenetic and biogeographic analyses (Extended Data Table 1and Extended Data Figs 1–4, 7–9) suggest the existence of two categoriesof endemics18: ‘eccentric’ (12 taxa) and ‘centric’ (25 taxa). The eccentrictype of endemic has sister taxa that occur either in temperate climates(seven cases) or in other tropical mountains outside of Borneo (fivecases). To this group belong all bryophytes, pteridophytes, some of thefungal lineages and also the endemics in the flowering plant groupsHedyotis, Ilex, Impatiens, Ranunculus and Euphrasia, and the animalsCoeliccia and Tritetrabdella. Eccentric endemics predominantly occur at

Layang

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Figure 1 | Map of the study area. Inset left, location of the study area in theWorld and in Borneo. Inset middle, detail of the summit trail in Kinabalu Park.The eight expedition stations in Kinabalu Park and Crocker Range Park are

indicated with red markers, ten additional sampling sites with blue markers.Not indicated separately are 15 sites along the summit trail, and four sites veryclose to Mahua, Gunung Alab, and Inobong.

I. aureliana (China: Yunnan)

I. poilanei (Vietnam)

I. eberhardtii (Vietnam)

I. kinabaluensis (Kinabalu HQ, Gn. Alab,

Poring)

I. chevalieri (Vietnam)

I. latifolia (South India)

I. flaccida (South India)

I. cuspidata (South India)

I. hawkeri (New Guinea)

I. thamnoidea (Tanzania)

I. pseudohamata (Tanzania)

I. chinesis (Yunnan)

I. platypetala (Bali)

I. serpens (Tanzania)

I. usambarensis (Tanzania)

I. walleriana (Eastern Africa)

I. balsamina (India)

P. n. sp. (Sugud)

P. bryantioides (Poring)

P. bryantioides

(Kota Kinabalu & Kiansom)

P. latimanus (Sayap)

P. kinabaluensis (Kinabalu HQ & Gn. Alab)

P. similipes (Sugud)

P. similipes (Sugud)

P. similipes (Sayap)

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C. vitreopileatus (New Zealand)

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C. austroturmalis (Chile)C. argutus (Norway)C. cliduchus (Germany)C. glaucopus (Germany)

C. sp. (2,900 m)C. rotundisporus (Australia)C. delibutus (Germany)C. lustrabilis (Norway)C. sp. (2,100 m)

C. sp. (1,000 m)

C. sp. (2,900 m)C. sp. (1,900 m)C. calyptratus (USA)

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Figure 2 | Examples of phylogenetic reconstructions. a, Centric endemism(Ptomaphaginus). b, Eccentric endemism (Impatiens). c, Centric endemismsuperimposed on eccentric endemism (detail of the Cortinarius phylogeny).Clades are coloured with the same colour scheme as in Fig. 1, by elevational

zone at which they occur (see legend). High-elevation endemics in a and b areindicated in red. For support values and other details on these and other trees,see Extended Data Figs 1–4.

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high elevations (mean lower elevational boundary, 2,212 m; s.d., 837 m),they are strict Kinabalu endemics (they do not occur on nearby, lowermountains), and are further characterized by high dispersal capacities(one, two and seven clades with eccentric endemism have small, mediumand large dispersal, respectively).

The centric type of endemic has sister taxa that occur locally, in theBorneo lowlands. Six out of the eight endemic animal species, some ofthe fungal lineages, the conifers, and 17 of the 20 flowering plantsbelong to this type. Centric endemics occur on average at lower eleva-tions (mean lower elevational boundary, 1,724 m; s.d., 728 m) andtend to have lower dispersal capacities than the eccentric endemics(eight, seven and ten clades with centric endemism have small, med-ium and large dispersal, respectively). Some centric endemics arenot strict Kinabalu endemics, as they are also found on other moun-tains in Kinabalu’s vicinity. Our ancestral state reconstructions(Supplementary Table 6) confirm a pattern of altitudinal bottom-upcladogenesis in the centric endemics, with 8 out of 11 species rangingpartially or entirely outside of the 95% credibility interval for thereconstructed elevation of the most recent common ancestor, andthe remaining 3 out of 11 falling entirely within this interval. Whenwe performed the same analysis with Kitayama’s (ref. 19) seven vegeta-tion zones, rather than elevation, we found that only three of theseeight elevation shifts may represent a shift towards a higher vegetationzone. This suggests that, even in high-elevation centric endemics, sub-stantial niche shifts are rare. Niche conservatism is probably evenunderestimated, since the Massenerhebung effect causes an upwardshift of vegetation zones on Kinabalu compared with other Borneanmountains14,20.

Our multi-taxon study shows that Kinabalu’s biodiversity hot spotis of recent origin. This means the mountain is an evolutionarycradle, accumulating neo-endemics, as has been suggested for otheryoung, high-elevation biodiversity hot spots at low latitudes21, suchas the Tibetan plateau22, the Andean highlands8, and Afrotropicalvolcanoes23. However, probably as a consequence of the rapid emer-gence of the mountain15 and its unique alpine summit conditions19,many of these neo-endemics have not evolved by drastic nicheshifts from local ancestors, but rather by immigration of pre-adaptedpropagules from elsewhere. This explains the multiple independentcolonization events in some taxa (for example, Glomus, Rhododendronand Coeliccia). In addition, local lowland taxa have also generatedmontane species, but although some of these have reached vegetationzones above 2,000 m, most do not show substantial niche shifts away

from their ancestral niche. The fact that the endemic biota of Kinabaluappears to be composed largely of pre-adapted (eccentric) species andlocally derived (centric), ecologically conserved endemics is in linewith niche conservatism24.

We suggest that our novel approach of molecular dating of multipleclades be applied to larger communities in this and other tropicalmontane biodiversity hot spots25. In combination, such informationshould allow a detailed dissection of the relative roles of ecologicalspeciation, colonization and habitat filtering in the formation ofendemic biotas in this and other tropical mountains. Moreover, suchunderstanding could improve predictions of the likelihood of extinc-tion and evolutionary rescue of endemic species experiencing chan-ging climate conditions7.

Online Content Methods, along with any additional Extended Data display itemsandSourceData, are available in the online version of the paper; references uniqueto these sections appear only in the online paper.

Received 9 December 2014; accepted 14 July 2015.

Published online 12 August 2015.

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17. Wong, K. M. & Phillips, A. Kinabalu, Summit of Borneo (Sabah Society & SabahParks, Kota Kinabalu, 1996).

18. van Steenis, C. G. G. J. Plant geography of the mountain flora of Mt. Kinabalu. Proc.R. Soc. Lond. B 161, 7–38 (1964).

19. Kitayama, K. An altitudinal transect study of the vegetation on Mount Kinabalu,Borneo. Vegetatio 102, 149–171 (1992).

20. Grubb,P. J. InterpretationofMassenerhebungeffecton tropicalmountains.Nature229, 44–45 (1971).

21. Favre, A. et al. The role of the uplift of the Qinghai–Tibetan Plateau for the evolutionof Tibetan biotas. Biol. Rev. Camb. Philos. Soc. 90, 236–253 (2015).

22. Schwery, O. et al. As old as the mountains: the radiations of the Ericaceae. NewPhytol. 207, 355–367 (2015).

23. Price, T. D. et al. Niche filling slows the diversification of Himalayan songbirds.Nature 509, 222–225 (2014).

24. Crisp, M. D. et al. Phylogenetic biome conservatism on a global scale. Nature 458,754–756 (2009).

25. Culmsee, H. & Leuschner, C. Consistent patterns of elevational change in treetaxonomic and phylogenetic diversity across Malesian mountain forests.J. Biogeogr. 40, 1997–2010 (2013).

Supplementary Information is available in the online version of the paper.

Acknowledgements We acknowledge the mountain guides of Sabah Parks for theirhelpwith material collection, ongoing assistance tocontinuing research in thearea, andrecent efforts following the 5 June 2015 earthquake. R. Butot, M. de Jong, and

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Figure 3 | Elevations and ages for endemic species. Elevations (mid-points,minima and maxima) and dates of origination derived from moleculardating (averages and 95% credible intervals) for endemic species; eccentricspecies (see main text) are indicated with a black circle. The vertical dashed lineindicates the oldest possible date for Kinabalu to have reached its currentelevation. For details, see Extended Data Figs 5 and 6.

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B. Driessen carried out DNA extractions and PCRs. P. Wilkie provided a tissue samplefor Payena. For the fungi, T. Semenova extracted DNA from the soil samples andE. Duijm generated the reads on the Ion Torrent sequencer. B. Blankevoort andE. J. Bosch prepared Fig. 1. J. Holloway (Natural History Museum, London, UK), I. Das(Universiti Malaysia Sarawak), and T. S. Liew (Naturalis, UMS) provided background onKinabalu endemism, and R. Hall (University of London) on its geological history.R. Etienne (University of Groningen), R. Hall (University of London) and F. Bossuyt (FreeUniversity Brussels) read a draft of the paper. Collection and export permits weresupplied by the Sabah Biodiversity Centre. Besides institutional support viaNetherlands FES-funding, this project received funding via grants 821.01.002 and825.12.007 of the Netherlands Organization for Scientific Research, the AlbertaMennegaFoundation, theEcology Fundof theRoyalNetherlandsAcademyofSciences,the Uyttenboogaart-Eliasen Foundation, and Pro Acarologia Basiliensis. G. van Uffelen,R. van Vugt, and P. Kessler (Botanic Gardens, Leiden) and T. de Winter, G. Thijsse,B.Hoeksema, H.Dautzenberg, J.Mols, andE. van Nieukerken (Naturalis) gaveadviceontransportation of specimens. Artist I. Kopelman and Naturalis general director E. vanHuis (Naturalis) accompanied the expedition participants in the field. J. Gunsalam andP. Jilan (Sabah Parks Board of Trustees) officiated the opening and closing ceremony,respectively. P.Basintal, E. van HuisandE.Smets supervised theexpeditionasdirectorsfor the organizing institutions. Media coverage and communications were facilitated byJ. van Alphen, A. Mohd. R. P. Dahlan, K. de Greef, A. Kromhout, J. Schilthuizen,E. E. Soudi, L. Stek, A. Taminiau, and M. Z. Abdul Wahab. Sabah Parks staff membersL. Apin, B. Busin, B. Butit, J. Gampoyo, G. Gangko, B. Gangku, S. Gangku, P. Gimpiton,M. Gunggutou, T. Gunggutou, G. Gunsalam, J. Guntabid, I. Isnain, K. Ag. Jalil, Y. Johalin,R. M. Karim, Y. Kumin, D. Laban, A. Lahi, J. Lapidin, M. Latim, N. Maidin, N. Majuakim,R. Miadin, Y. Miki, M. Moguring, H. Mujih, F. Simin, K. C. Sion, D. Sumbin, D. Tingoh,E. Yakin, were invaluable for all manner of logistics and other support before, during,and after the expedition, in the field and at the stations and administrative offices. For

the Forest Research Centre researchers, J. B. Dawat, M. Gumbilil, A. J. Jaimin, Jei.Jumian, Jem. Jumian, J. L. Yukang, P. J.Miun,M. A. T. Mustapha, and B. Saludin assistedin the field. T. Siew of Borneo Indah Sdn. Bhd. was the local supplier for consumablesand field equipment. We thank the ESEB Outreach Fund and the Treub Foundation forfunding outreach activities in Kinabalu Park and the Crocker Range Park. N. Raeshelped with the niche modelling.

Author Contributions M. Sch., V.S.F.T.M., M.Sp., M.M.B., C.B.M., L.E.B., K.T.C.A.P., J.N.,R.R., and M.L. conceived and guided the study. M.Sch. and V.S.F.T.M. wrote the paper.K.H., F.S. and K.B. administrated all samples and sequences. K.H. and L.E.B. wereresponsible for DNAsampling in the field. V.S.F.T.M., A.A.,N.A., A.B., P.-P.C., A.Y.C.C., R.D.,H.F., C.F.-v.S., J.G., R.G., P.H., P.I., I.I., M.J., H.K., E.K., P.K., F.L., R.J.M., L.N.M., N.N., J.T.P.,H.R., S.S., A.S., R.M.S., P.-S.S., H.S., N.S., J.B.S., M.Sul., S.Sum., J.v.T., F.Y.Y.T., B.E.Y., C.B.M.,R.R. and M.Sch. collected material in the field and identified it following currenttaxonomy. K.H., V.S.F.T.M., H.d.B., F.A.A.F., J.G., B.G., P.H., S.B.J., S.N., M.St., D.C.T. andM.Sch. conducted the analyses.

Author Information GenBank accession numbers for newly determined sequencesare: KF369344, KF492848, KF839944–KF839949, KM378672–KM378719,KM489525–KM489544, KM982697–KM982715, KP152323–KP152394,KP978313–KP979097, KR002164–KR002181, KR030167–KR030238 andKR232819–KR232836. A large portion of the data is also publicly available on BOLD(www.boldsystems.org), as dataset DS–KINA (http://dx.doi.org/10.5883/DS-KINA).Voucher specimens have been archived in public natural history collections. Reprintsand permissions information is available at www.nature.com/reprints. The authorsdeclare no competing financial interests. Readers are welcome to comment on theonline version of the paper. Correspondence and requests for materials should beaddressed to M.Sch. ([email protected]).

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http://www.ncbi.nlm.nih.gov/nuccore/KF369344




http://www.ncbi.nlm.nih.gov/nuccore/KM378672






http://www.ncbi.nlm.nih.gov/nuccore/KP152323




http://www.ncbi.nlm.nih.gov/nuccore/KR002164

http://www.ncbi.nlm.nih.gov/nuccore/KR002181

http://www.ncbi.nlm.nih.gov/nuccore/?term=KR030167




www.boldsystems.org

http://dx.doi.org/10.5883/DS-KINA

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METHODSNo statistical methods were used to predetermine sample size. The experimentswere not randomized and the investigators were not blinded to allocation duringexperiments and outcome assessment.Expedition. Material was sampled chiefly between September 10th and 25th,2012, by the participants of the Crocker Range/Kinabalu Scientific Expedition,which was conceived, organized, funded and conducted jointly by Sabah Parks(Malaysia) and Naturalis Biodiversity Center (The Netherlands). The expeditionwas set up with the explicit aim to investigate evolutionary patterns of endemism(see main text), and participants were selected to maximize taxon coverage andecological diversity. The 47 expedition members were asked to identify one ormore congeneric species complexes (hereafter termed ‘genera’) within their taxonof expertise that included high elevation endemics and preferably also more wide-spread lowland species (although there is a possibility that this approach wouldleave out any monotypic endemics, in the groups studied by us, such species arenot known). These, plus outgroup species, were then sampled (using taxon-specific methods; see Supplementary Table 2 for further details) at eight expeditionstations located at a range of elevations (Fig. 1), and also in additional locations inboth parks and their vicinity. Data on time, latitude and longitude, elevation, andhabitat were collected with each sample. Material was subsampled for DNA ana-lysis, mostly in the field, by preserving tissues or entire organisms, usually eitherin pure ethanol or in silica gel (see Supplementary Table 2 for taxon-specificsubsampling and storage). In the case of fungi, mycelial DNA was obtained bybulk extraction from composite soil cores (see Supplementary Table 2 and below).In addition to the data collected during the expedition, the expedition memberswere asked to provide a complete assessment of elevational and geographicaldistribution for each of the species within their taxon of expertise, based onpublished sources and previous collections.Molecular analysis. Upon arrival in the Naturalis DNA barcoding facility, sub-samples were processed in a semi-automated way. DNA extractions on animalmaterial were performed on either tissue samples or individual arthropod legs, orusing whole arthropods in a non-destructive extraction method. DNA wasextracted using the NucleoMag 96 Tissue kit (Macherey-Nagel Gmbh & Co.,Duren, Germany) on the KingFisher Flex magnetic particle processor (ThermoScientific). DNA extractions on plant tissue were performed on dried leaf samples,using the NucleoMag 96 Plant kit (Macherey-Nagel Gmbh & Co., Duren,Germany). The KingFisher Flex protocols were slightly adjusted from the manu-facturer’s protocol (http://www.thermoscientific.com) to include longer mixing ofbeads with lysed sample and a drying step before elution. A volume of 150ml wasused for elution in both animal and plant samples. Soil samples containing fungalmycelia were processed by pooling 40 cores (for sampling details, seeSupplementary Table 2), resulting in a composite soil sample for each site.Genomic DNA was extracted from 1 g of this sample using the NucleoSpin Soilkit (Macherey-Nagel Gmbh & Co., Duren, Germany), according to the manufac-turer’s protocol. For each sample DNA extraction was carried out twice andreplicates were combined.

For each specimen, the accepted barcode marker for its taxon was sequenced, inmany cases augmented with one or more other phylogenetically informative loci.Supplementary Tables 1 and 2 provide lists of the loci and primers used for eachgenus included in the study. Where relevant for outgroup comparison or furthergeographic resolution, we then obtained additional sequences from Borneo orelsewhere from GenBank or from our own, unrelated studies. GenBank accessioncodes for these are given in Supplementary Table 2. After sequencing, (remains of)voucher specimens were mostly deposited in the collections of Sabah Parks and/orNaturalis Biodiversity Center. Voucher collection codes are listed in Supple-mentary Table 2.

PCR reactions generally contained 18.75ml ultrapure H2O, 2.5ml 103 PCRbuffer, 1.0ml 10 mM of each primer, 0.5ml 2.5 mM dNTPs and 0.25ml 5Uml21

Qiagen Taq. Most PCRs were performed using an initial denaturation step of 180 sat 94 uC, followed by 40 cycles of 15 s at 94 uC, 30 s at 50 uC and 40 s at 72 uC,and ended with a final extension of 300 s at 72 uC and pause at 12 uC.Bi-directional Sanger sequencing was performed at either Macrogen (http://www.macrogen.com) or BaseClear (http://www.baseclear.com). Sequences wereedited manually (primers trimmed off and obvious reading errors corrected) usingSequencher 4.10.1 (Gene Codes Corporation). For the fungal soil samples, theamplicon library was sequenced using an Ion 318TM Chip in an Ion TorrentPersonal Genome Machine (PGM; Life Technologies, Guilford, CT, USA). PCR,sequencing, raw data filtering and all bioinformatic analyses were performed asdescribed in detail previously26. From these fungal soil sequences, we selected non-redundant sequences of the ectomycorrhizal genus Cortinarius and the arbuscularmycorrhizal genus Glomus because they were distributed along the entire altitu-dinal range of our sampling.

Phylogenetic analysis. We performed initial exploratory alignment and createdneighbour-joining trees. This allowed an assessment of individuals that (on thebasis of cursory morphological examination) had been included as potential(undescribed) members of the species complexes of choice. In some cases, thesequence analysis showed that these individuals were in fact unrelated and suchsequences were then removed before further analysis. Then, using Geneious (ver-sion 6.0.5; created by Biomatters; available from http://www.geneious.com/) as aworkbench, sequence data were aligned with MUSCLE25. We ran ModelTest27 todetermine, using Akaike Information Criterion (AIC), the best model of nucleo-tide substitution per alignment, for each genetic marker separately. We thenperformed a Bayesian analysis using MrBayes 3.2.2 (ref. 28) with settings formultiple markers unlinked, running two replicates of a minimum of 107 genera-tions each. Convergence was checked in Tracer (v1.5; ref. 29) and, after discardinga 25% burn-in, the resulting majority-rule consensus trees were preserved. For thefungal data sets, multiple sequence alignments were built using MUSCLE and, dueto the large size of these data sets, phylogenetic analyses of maximum likelihood(ML) were carried out in Garli 2.0 (ref. 30) using the GTR1I1G model withestimated values of the alpha parameter of the gamma distribution of variablesites (G) and proportion of invariable sites (I). Clade support was assessed by non-parametric bootstrap analyses with 100 pseudo-replicates.Molecular dating. The times of divergence of Mount Kinabalu endemics fromtheir most closely related extant relatives were estimated with a molecular clockapproach. We included not only our own data, but also incorporated publishedstudies on genera other than the ones we sampled. For each genus, taxon samplingand molecular data used are specified in Supplementary Table 4. All data setswere assembled in Geneious (ver. 5.5.6; created by Biomatters; available fromhttp://www.geneious.com/). Data sets were pruned to contain a single taxon perspecies by selecting a random sequence from each species. Sequence alignmentswere obtained with the MAFFT (ver. 6.814b) alignment tool implemented inGeneious. Divergence time estimates were obtained using a relaxed molecularclock approach with BEAST (ver. 2.1.3; ref. 31). For each DNA region used, anunlinked substitution model was implemented as selected with jModeltest (ver.2.1.5; ref. 27) under the AIC. For the tree prior, a Yule birth rate prior was selectedwith a flat prior on the Yule birth rate. A single clock and tree model was used foreach analysis. All analyses were run with an uncorrelated log-normal clock model,to which an exponential prior was assigned for both the mean (ucldMEAN; meanset at 10.0) and the standard deviation (ucldStdev; mean set at 0.33). Time cal-ibration was done using: (1) fossil constraints; (2) secondary calibration points; or(3) substitution rates, depending on the information available for each particulargenus. In cases where suitable fossil or secondary calibration information was notavailable, we used a priori information about the substitution rate of one of thegene partitions—as opposed to the ‘fix the mean substitution rate’ option across allpartitions—to optimize the analyses. The full details on calibration methods usedfor each taxon are given in Supplementary Table 5. For each run, posterior dis-tributions of parameters were approximated using a Markov chain Monte Carloanalysis of 5 3 107 generations, sampling every 5,000 generations, discarding aburn-in of 5 3 106 generations (10%). Convergence of the chains was checked byevaluating the ESS values of each parameter with Tracer v1.6.Biogeographical, elevational and vegetation zone analyses. For each genus, anindependent estimation of the dispersal neighbourhood size was obtained fromthe spatial ecology literature on the genus concerned or related genera, and/orexpert knowledge. We used this information to classify dispersal neighbourhood(two standard deviations for the distribution of distances between birth andreproduction for individuals of a certain species) in the genus in question as ‘small’,,1 km (mostly flightless animals), ‘medium’, 1–100 km (flying or ballooninganimals, and plants dispersed by flying animals), or ‘large’, .100 km (plantsand fungi with very small wind-dispersed seeds or spores). An overview ofdispersal assessments is in Supplementary Table 3.

Ancestral ranges were estimated for each taxon group under maximum like-lihood with BioGeoBears32 using the three-parameter DEC1J model. The ultra-metric tree from the Bayesian relaxed molecular clock analysis was used as inputtree. This tree was pruned to include the clade with the endemic species and at leastone deeper node (for most groups two deeper nodes were included). Each specieswas scored to be absent or present in Mount Kinabalu (K), Borneo (B), and the restof the world (W). For each group the analysis was run with equal probability forexchange between the three areas over the entire diversification time of the tree.The maximum number of areas was assigned as the maximum number of areasobserved in the extant taxa (either 2 or 3). The optimizations, with the ancestralrange with the highest likelihood for each node, are shown in Extended DataFigs 7–9). We classify a ‘W’ to ‘K’ or ‘KW’ shift in the lineage leading to a speciesendemic to Kinabalu as ‘eccentric endemism’. All other ancestral ranges leadingto Kinabalu endemics are scored as ‘centric’.

LETTER RESEARCH


http://www.thermoscientific.com

http://www.macrogen.com

http://www.macrogen.com

http://www.baseclear.com

http://www.geneious.com

http://www.geneious.com

For each genus sampled during the expedition that showed centric endemism,we added the elevation at which it had been collected as a continuous character. InBayesTraits 2.0 (ref. 33) we reconstructed ancestral states for elevation at the nodeleading to the endemic species and its sister clade using the continuous randomwalk model which implements a generalized least squares model for continuouslyvarying traits. The elevation of the most-recent common ancestor of the node ofinterest was inferred using a MCMC approach on a set of 2,700 trees randomlysampled after the phylogenetic inference with MrBayes (excluding burn-in). TheMCMC chain of BayesTraits was run for 106 generations, discarding the resultssampled during the first 105 generations as burn-in. We then inspected whetherthe current range of elevation of the endemic species stretches beyond the 95%credible interval for the most-recent common ancestor. If it stretches above thisinterval, we considered this indicative of a ‘bottom-up’ scenario, whereas thereverse would suggest a ‘top-down’ scenario. The results are summarized inSupplementary Table 6.

In parallel with the elevational analysis, we conducted an analysis in which wecoded the vegetation belt(s) as taxon traits. We used the zones defined for Mt.Kinabalu in ref. 19, namely: (i) lowland vegetation (below 600 m); (ii) hill vegeta-tion (600–1,200 m); (iii) lower montane vegetation (1,200–2,000 m); (iv) uppermontane vegetation (2,000–2,800 m); (v) subalpine vegetation (2,800–3,400 m);(vi) upper subalpine vegetation (3,400–3,700 m); (vii) alpine vegetation (above3,700 m). The probabilities at each vegetation belt for the most-recent commonancestor of the node leading to the endemic clade and its sister group were inferredwith the continuous-time Markov model for discrete data implemented inBayesTraits 2.0. The same input trees were used as for the elevation reconstruction,and the MCMC was run for 106 generations, discarding the results sampled duringthe first 105 generations as burn-in. The resulting probabilities are shown inSupplementary Table 7.Geological summary. The following is based on refs 15,34 and R. Hall, personalcommunication, 2014.

Orogeny in the region began at about 23 million years ago (Mya) as a result ofthe northward movement of the Australian plate leading to the collision ofAustralia and Southeast Asia in what today is eastern Indonesia. Orogeny in theKinabalu area began at about 20 Mya after collision of north Borneo with theSouth China continental crust. This led to the emergence of most of Sabah. Soon

after this emergence much of Sabah became submerged, although a north–south-oriented mountain belt formed of tightly folded mudstones and sandstones (thefuture Crocker Range) remained. At the same time, the climate changed fromseasonal to ever-wet. Mapping of former shelf edges offshore shows that later theCrocker Range widened with time, implying that it became higher in its centralparts. Between 8 and 7 Mya a granitic body intruded the crust and cooled rapidlyfrom ,7 to 5 Mya. Melting could have resulted from a build-up of heat fromradioactive decay in the deep crust, or from decompression melting followingextension. After 6 Mya (the precise timing and duration is not known) thearea (including the surrounding Crocker Range) rose further, elevating to thepoint that it was glaciated recently, possibly after a lithospheric root detachedand sank into the deeper mantle, leading new mantle material to flow into thevacated space below the Crocker Range. Extensive glacial weathering of the sum-mit suggests the mountain had reached its present elevation by at least 1 Mya.The Pinosuk gravels on Kinabalu’s south-eastern slope are the accumulated ero-sional debris of young glaciation. Today, the mountain is still rising by 0.5 mmper year.

26. Geml, J. et al. Large-scale fungal diversity assessment in the Andean Yungasforests reveals strong community turnover among forest typesalong analtitudinalgradient. Mol. Ecol. 23, 2452–2472 (2014).

27. Posada, D. & Crandall, K. A. Modeltest: testing the model of DNA substitution.Bioinformatics 14, 817–818 (1998).

28. Ronquist, F. & Huelsenbeck, J. P. MrBayes 3: Bayesian phylogenetic inferenceunder mixed models. Bioinformatics 19, 1572–1574 (2003).

29. Rambaut, A. & Drummond, A. J., Tracer v1.5. http://beast.bio.ed.ac.uk/Tracer(2009).

30. Zwickl, D. J. Genetic Algorithm Approaches for the Phylogenetic Analysis of LargeBiologicalSequenceDatasets under the MaximumLikelihoodCriterion (The Universityof Texas, Austin, 2006).

31. Bouckaert, R. et al. BEAST 2: a software platform for Bayesian evolutionaryanalysis. PLOS Comput. Biol. 10, e1003537 (2014).

32. Matzke, N. J. Probabilistic historical biogeography: new models for founder-eventspeciation, imperfect detection, and fossils allow improved accuracy and model-testing. Front. Biogeogr. 5, 242–248 (2013).

33. Meade, A. & Pagel M. BayesTraits, version 2.0. http://www.evolution.rdg.ac.uk(2015).

34. Hall, R. et al. The Geology of Mount Kinabalu (Sabah Parks, 2008).

RESEARCH LETTER


http://beast.bio.ed.ac.uk/Tracer

http://www.evolution.rdg.ac.uk

0.06

RBINS.15995 K. sp.2

RBINS.15993 K. sp.2

RBINS.16009 K. baluensis

AB611921 Kalophrynus pleuros�gma



RBINS.16000 K. sp.2

AB611939 Metaphrynella sundana

RBINS.16006 K. sp.2

AB611913 K. interlineatus

AB611931 Metaphrynella pollicaris

RBINS.15994 K. sp.2

RBINS.15999 K. sp.2

RBINS.15998 K. sp.2

RBINS.16001 K. sp.2

RBINS.16007 K. sp.2


RBINS.15992 K. sp.1



AB611973 Phrynella pulchra

0.78

1

0.65

1

0.99

1

1

Anura - Kalophrynus

Aranea - Myrmarachne

0.4

NMF2012-01399.PK267 M. malayana

NMF2012-01399.PK024 M. mariaelenae












NMF2012-01399.PK373 M. cornutaAY297412 M. assimilis (Philippines)

NMF2012-01399.PK080 M. sp.5







NMF2012-01399.PK210 M. hanoii




EU815599 Ligonipes sp. (Australia)

NMF2012-01399.PK260 M. nov.sp.2





NMF2012-01399.PK256 M. nov.sp.2





NMF2012-01399.PK052 M. sp.4

















NMF2012-01399.PK264 M. nov.sp.2









JX145678 M. evidens (Gabon)


NMF2012-01399.PK213 M. mariaelenaeNMF2012-01399.PK215 M. mariaelenae

NMF2012-01399.PK257 M. nov.sp.2


NMF2012-01399.PK081 M. sp.5



NMF2012-01399.PK050 M. nov.sp.1


NMF2012-01399.PK183 M. sp.3










NMF2012-01399.PK206 M. sp.3


NMF2012-01399.PK036 M. christae




NMF2012-01399.PK073 M. sp.5



JX145679 M. foenisex (Gabon)



NMF2012-01399.PK051 M. sp.4












NMF2012-01399.PK357 M.cornuta


NMF2012-01399.PK077 M. sp.5


JX145676 M. cf. gedongensis (Malaysia)


0.54

0.84

1

0.72

0.5

1

0.87

0.99

0.96

0.75

1

1

1

1

1

0.79

1

0.98

0.85

1

0.68

1

1

0.73

0.86

1

0.03

EU329039 EU329082 A. nakanoi (Japan)

EU329067 EU329110 Diplazium nipponicum (Japan)

EU329055 EU329098 A. yokoscense (Japan)

EU329059 EU329102 A. x tokashikii (Japan)

EU329051 EU329094-A.tozanense (Japan)

EU329037 EU329080 A. kuratae (Japan)

EU329049 EU329092 A. strigillosum (Japan)

EU329026 EU329070 A. atkinsonii (Japan)

EU329058 EU329101 A. x hisatsuanum (Japan)

EU329042 EU329085 A. niponicum (Japan)

e4002185655 A. amoenum

EU329030 EU329074 A. deltoidofrons (Japan)

e4002185620 A. clemensiae

EU329025 EU329069 A. arisanense (Japan)EU329031 EU329075 A. eremicola (Japan)

EU329032 EU329076 A. filix-femina-cul�vated


EU329046 EU329089 A. setuligerum (Japan)

EU329044 EU329087 A. otophorum (Japan)

EU329041 EU329084 A. nikkoense (Japan)

EU329040 EU329083 A.neglectum (Japan)

EU329057 EU329100 A. x akiense (Japan)

EU329063 EU329106 Cornopteris decurren� alata (Japan)

EU329035 EU329078 A. iseanum var. angus�sectum (Japan)

EU329050 EU329093 A. tashiroi (Japan)



EU329028 EU329072 A. clivicola (Japan)


EU329038 EU329081 A. melanolepis (Japan)

EU329043 EU329086 A. oblitescens (Japan)

EU329034 EU329077 A. iseanum (Japan)


e4002185539 A.amoenum

EU329068 EU329111 Diplazium hachijoense (Japan)

EU329061 EU329104 Cornopteris christenseniana (Japan)

EU329060 EU329103 Cornopteris christenseniana (Japan)

EU329045 EU329088 A. rupestre (Japan)EU329056 EU329099 A. yokoscense var. dilatatum (Japan)

e4002185646 Diplazium simplicivenium

EU329062 EU329105 Cornopteris crenulatoserrulata (Japan)

EU329047 EU329090 A. sheareri (Japan)EU329029 EU329073 A. cuspidatum (China)

EU329036 EU329079 A. kirisimaense (Japan)

EU329054 EU329097 A. wardii (Japan)

EU329027 EU329071 A. brevifrons (Japan)

EU329053 EU329096 A. viridescen�pe (Japan)

EU329064andEU329107-Deparia.petersenii-Japan

EU329052 EU329095 A. vidalii (Japan)



EU329065andEU329108-Deparia.conilii-Japan

EU329048 EU329091 A. subrigescens (Japan)

EU329066andEU329109-Deparia.lobatocrenata-Japan

0.95

1 1

1

0.91

0.99

1

1

1

0.96

0.99

0.75

0.7

1

0.75

0.91

0.81

0.68

1

1

0.69

0.57

0.99

1

1

0.96

1

1

0.64 0.85

0.94

1

1

0.6

1

0.99

0.92

1

1

1

Athyriaceae - Athyium

Balsaminaceae - Impatiens

0.0070

FJ826636 I. chevalieri (Vietnam)

DQ147814 I. aureliana (China, Yunnan)

DQ147849 I. hians Africa

DQ147854 I. la�folia (Indian)

FL073 I. kinabaluensis

DQ147869 I. poilanei (Vietnam)

tbd-I.serpens-

tbd-I.thamnoidea-

tbd-I.pseudohamata-

CM0015 I. kinabaluensis

DQ147850 I. hawkeri (Papua New Guinea)



DQ147845 I. flaccida (India)


DQ147839 I. eberhard�i (Vietnam)

DQ147832 I. cuspidata (India)


DQ147892-I.walleriana-African_origin

DQ147825 I. chinensis (China, Yunnan)

DQ147868 I. platypetala (Indonesian, Bali)

tbd-I.usambarensis

DQ147816 I.balsamina (Indian)

1

1

1

1

1

1

1

1

1

1

1

1

1

0.68

Begoniaceae - Begonia

0.0030

FL040 B. burbidgei

SNP25535_2 B.vaccinioides

FL066 B.cauliflora

PH115 B. spec2

FL074 B. cf. beryllae

FL071 B. beryllae

LLF141 B. amphioxus

DCT08-102 B. kingiana (Malaysia)

DCT07-26 B. burbidgei

DCT09-03 B.pendula (Malaysia)

FL062 B. spec1

FL065 B.cauliflora

SNP25535_1 B.vaccinioides

FL075 B. imbricata

DCT07-24 B. masoniana (China)

1

1

0.98

1

0.89

0.57

1

1

3500-4000m3000-3500m2500-3000m2000-2500m1500-2000m1000-1500m500-1000m0-500m

Extended Data Figure 1 | Phylogenetic reconstructions for Anura, Araneae, Athyriaceae, Balsaminaceae and Begoniaceae. Details of materials and methodsused are given in Supplementary Table 2.

LETTER RESEARCH


Coleoptera - Ptomaphaginus

0.08

RMNH.INS.549270 Ptomaphaginus bryan�oides


RMNH.INS.555623 Ptomaphaginus ater

RMNH.INS.555603 Ptomaphaginus kinabaluensis


RMNH.INS.555633 Ptomaphaginus.nr. fagei

RMNH.INS.555610 Ptomaphaginus similipes

RMNH.INS.555611 Ptomaphaginus c.f. la�manus



RMNH.INS.555619 Ptomaphaginus n.sp. bryan� complex


RMNH.INS.549243 Prionochaeta harmandi (Japan)








RMNH.INS.555607 Ptomaphaginus latescens

RMNH.INS.555631 Ptomaphaginus kinabaluensisRMNH.INS.555601 Ptomaphaginus kinabaluensis















RMNH.INS.555632 Ptomaphaginus nr. fagei







RMNH.INS.555624-Ptomaphaginus ater



1

1

0.79

0.86

1

0.85

1

1

1

1

1

1

1

1

0.98

0.92

0.95

1

1

0.88

1

1

Cortinariaceae - Cortinarius

5 changes

“up”“mid”“low”

FJ039601.1 Cor�narius olivaceopictusAF539707.1 Cor�narius luteostriatulusAF539708.1 Cor�narius obscurooliveus

GU233323.1 Cor�narius alienatusOTU 12584

OTU 12602AY669588.1 Cor�narius collinitus

OTU 7973OTU 12746

OTU 17155OTU 16366OTU 25865

OTU 25299OTU 2634OTU 21065OTU 19102

OTU 18261GU233334.1 Cor�narius vitreopileatus

OTU 1813DQ652235.1 Cor�narius lustrabilis

OTU 1149OTU 4508OTU 7114

EU525980.1 Cor�narius calyptratusAY174787.1 Cor�narius glaucopus

AY669558.1 Cor�narius langeiAY669535.1 Cor�narius argutus

AY669587.1 Cor�narius delibutusAY669612.1 Cor�narius rotundisporus

AF539730.1 Cor�narius austroturmalisOTU 1023

OTU 2153OTU 18981OTU 19822OTU 19034

OTU 26933OTU 2553

OTU 17207OTU 18659

OTU 26171OTU 26541

OTU 23750OTU 12474

OTU 2644OTU 19460

OTU 9147OTU 5013

OTU 9521FJ039611.1 Cor�narius duracinusGQ159908.1 Cor�narius pseudocandelaris

OTU 26194OTU 7849

GQ159860.1 Cor�narius velenovskyi OTU 17175OTU 19944

OTU 6677OTU 12976

AY669656.1 Cor�narius incisus AY669655.1 Cor�narius acutovelatus

FJ039606.1 Cor�narius ceraceusFJ157003.1 Cor�narius acutusOTU 3231

OTU 12440AY669651.1 Cor�narius cys�diocatenatus

OTU 8882OTU 6053

AF539727.1 Cor�narius pachynemeusOTU 13501

OTU 17073OTU 3195

OTU 18150OTU 15546

OTU 17985OTU 10720

OTU 16037OTU 9069

OTU 18957OTU 14593

OTU 22118OTU 24443

OTU 3600OTU 4725

OTU 11980OTU 15329

OTU 9940OTU 13013

OTU 8619OTU 12349

OTU 12484EU655655.1 Cor�narius rapaceotomentosus

DQ663411.1 Cor�narius sanc�-felicisEU655660.1 Cor�narius insignibulbus

EU057001.1 Cor�narius jardinensisEU056976.1 Cor�narius magnivelatus

OTU 26725OTU 12377

OTU 19253OTU 12260

OTU 752EU660953.1 Cor�narius cretax

OTU 5187OTU 1410

OTU 15101OTU 27038

OTU 14504OTU 14517

OTU 4929OTU 14916

OTU 14512OTU 9017

OTU 2674OTU 24339

OTU 18369OTU 15128

AF539736.1 Cor�narius olivaceobubalinus OTU 4232

OTU 6233OTU 3490OTU 10798OTU 2456

OTU 8407OTU 18770

OTU 857OTU 26840

OTU 3898AY669654.1 Cor�narius spilomeus

OTU 133FJ039604.1 Cor�narius ochrophyllus

OTU 7587OTU 3280

OTU 3693OTU 11718

OTU 5836AY669596.1 Cor�narius bolaris

OTU 8827AY669645.1 Cor�narius anomalus

EF420146.1 Cor�narius sericeolazulinusFJ039651.1 Cor�narius anomalovelatus

OTU 4234OTU 6434

GQ159887.1 Cor�narius anomalovelatusOTU 8855

OTU 10556AY669646.1 Cor�narius caninus

GQ159900.1 Cor�narius rigensAY669637.1 Cor�narius schlerophyllarum

OTU 406OTU 23850

GU234083 Hebeloma mesophaeum OTU 2041

OTU 15011OTU 8218

GQ159827.1 Cor�narius purpurascensAY669520.1 Cor�narius nanceiensisOTU 5338

AY669521.1 Cor�narius populinusAY174797.1 Cor�narius saginusOTU 9222

OTU 14845FJ039613.1 Cor�narius balteatus

OTU 1594GU233350.1 Cor�narius anauensis

OTU 9813OTU 27167

FJ039619.1 Cor�narius herpe�cusFJ039622.1 Cor�narius herpe�cus

FJ039621.1 Cor�narius herpe�cusEU821686.1 Cor�narius salor

GU233329.1 Cor�narius aerugineoconicusOTU 2542

AY033104.1 Cor�narius elaca�pusOTU 13359

OTU 1859OTU 15334

AY669677.1 Cor�narius fulvoconicusAJ889977.1 Cor�narius torvus

EU433388.1 Cor�narius agathosmusOTU 504

OTU 9276OTU 8686

OTU 116OTU 4229

OTU 1668OTU 3529OTU 10831

OTU 10378OTU 5366

EU821656.1 Cor�narius traganusFJ039542.1 Cor�narius ser�pes

FJ717541.1 Cor�narius saniosusOTU 1074

OTU 12791AY669658.1 Cor�narius umbrinolens

AY669667.1 Cor�narius helvolusFJ039572.1 Cor�narius venustus

AY669670.1 Cor�narius anthracinusAY669688.1 Cor�narius biformis

GQ159813.1 Cor�narius saturninus AY669687.1 Cor�narius umbilicatus

OTU 2471OTU 6606

OTU 12519OTU 1496OTU 26469

AY669681.1 Cor�narius malachius OTU 9572

OTU 601OTU 26865OTU 18561

OTU 25211OTU 10559

AM713178.1 Cor�narius assiduus OTU 16806

DQ499466.1 Cor�narius boulderensis GQ159914.1 Cor�narius fulvescens

AY669679.1 Cor�narius subser�pesOTU 5651

OTU 11821OTU 2564

AY669665.1 Cor�narius hinnuleusAY669673.1 Cor�narius rubricosus

GQ159883.1 Cor�narius candelarisOTU 17537OTU 11785

OTU 12030FJ039708.1 Cor�narius sp.

OTU 13067GQ159774.1 Cor�narius turibulosus

AY669686.1 Cor�narius everniusAF430292.1 Cor�narius brunneus

EU266692.1 Cor�narius gen�lisAJ236072.2 Cor�narius pholideus

OTU 9936AY669607.1 Cor�narius basipurpureus

AY669594.1 Cor�narius callisteusEU821663.1 Cor�narius huronensis

AY669630.1 Cor�narius cannariusOTU 12419

EU821696.1 Cor�narius vibra�lisAF539734.1 Cor�narius lividusOTU 416

GQ159773.1 Cor�narius valgusOTU 14041

EU821684.1 Cor�narius gen�lisGU233333.1 Cor�narius aegrotus

OTU 10420OTU 14591

OTU 18368OTU 11462

OTU 7233OTU 5948DQ097876.1 Cor�narius callisteus

OTU 6025OTU 5985

OTU 19AF112147.1 Dermocybe austrovenetaAY669632.1 Cor�narius walkeri

AF389160.1 Cor�narius amoenusAY669595.1 Cor�narius rubellusAJ236064.2 Cor�narius rubellusOTU 14254

OTU 3192OTU 14516

GU233342.1 Cor�narius picoidesOTU 13297

AY174782.1 Cor�narius infractusOTU 3972

OTU 14800OTU 26181

OTU 727OTU 14618

OTU 6126OTU 4057

AY669583.1 Cor�narius malicoriusFJ039596.1 Cor�narius humboldtensis

OTU 3916FJ039701.1 Cor�narius sp.

GQ159873.1 Cor�narius tubariusOTU 17187OTU 468

AY082608.1 Dermocybe cinnamomeaAY669585.1 Cor�narius olivaceofuscus

FJ157089.1 Cor�narius neosanguineusFJ157116.1 Cor�narius neosanguineus

OTU 580OTU 14542

OTU 4810OTU 27315OTU 197

OTU 11469OTU 4748

EU821667.1 Cor�narius phoeniceusFJ039591.1 Cor�narius smithiiFJ039594.1 Cor�narius idahoensisAY669581.1 Cor�narius palustris GQ159882.1 Cor�narius chrysolitus

Dicranaceae - Dicranoloma

0.02

DQ485176 D. leichhard�i (Australia)

e4002189779 D. havilandii

DQ462607 DQ464184 D. menziesii (New Zealand)

e4002189823 D. braunii

DQ462605 DQ485175 D. dicarpum (New Zealand)

AF135075 AF144116 Paraleucobryum enerve

e4002189802 D. assimile

DQ462599 DQ464189 D. robustum (New Zealand)

KF423978 KF423621 D. cylindrothecium (Russia)DQ462601 DQ464191 D. robustum (Chile)


DQ462604 DQ464186 D. chilense (Chile)

e4002189775 D. reflexum

AF146376 AF144112 Mesotus celatus

DQ462594 DQ464193 D. billarderi (New Zealand)

DQ462598 DQ464188 D. robustum (New Zealand)

MS9_e4002189739 D. brevisetum

e4002189763 D. brevisetum

DQ462596 DQ464196 D. billarderi (Rwanda)



L0856640 e4002189709 D. sp. (Indonesia)




DQ462597 DQ464197 D. billarderi (Reunion)

DQ462606 DQ464185 D. plurisetum (New Zealand)



KF423992 KF423638 Holomitrium crispulum


L0093198 e4002189685 D. reflexum (Indonesia)


e4002189776 D. blumii


e4002189814 D. blumii

L0092869 e4002189708 D. diaphanoneurum (Australia)


AF129587 AF144113 Dicranum polysetum

DQ462595 DQ464194 D. billarderi (Chile)

e4002189788 D. brevisetum





DQ485179 D. menziesii (Australia)

e4002189787_D_reflexum_Kinabalu_Sayap

DQ462600 DQ464190 D. robustum (Chile)

KF423995 KF423641 Holomitrium arboreum

DQ485178 D. fasciatum (Australia)


e4002189768 D. assimilee4002189791 D. assimile

L0856682 e4002189720 D. havilandii

AF135067 AF140696 Dicnemon calycinum

e4002189780 D. blumii

DQ485174 D. plurisetum (New Zealand)

1

0.84

1

0.73

1

1

0.84

1

1

0.91

1

1

0.86

0.96

0.82

0.65

0.66

1

1

0.76

0.990.99

1

0.99

0.57

1

0.95

1

0.97

1

1

1

Diptera - Diopsidae

0.02

RMNH.INS.557898 Teleopsis scapulistylus sp.n.

RMNH.INS.557894 Teleopsis nr. quinquegu�ata

RMNH.INS.555159 Teleopsis nr. sexgu�ata

RMNH.INS.555138 Teleopsis pallifacies

RMNH.INS.557902 Teleopsis sabah sp.2

Teleopsis discrepans XIII1


RMNH.INS.555156 Teleopsis longiconus sp.n.

RMNH.INS.555161 Cyrtodiopsis nr. dalmanni sp. 2

Teleopsis pallifacies IV5

RMNH.INS.555116 Teleopsis pseudogracilifemur sp.n.

RMNH.INS.557945 Teleopsis nr. discrepans sp. B

RMNH.INS.555127 Teleopsis gracilifemur sp.n.

RMNH.INS.555146 Teleopsis nr. discrepans sp. A

RMNH.INS.557959 Teleopsis nr breviscopium




Teleopsis_gracilifemur sp.n. V2

RMNH.INS.555141 Teleopsis nr. discrepans sp. C

RMNH.INS.555117 Teleopsis gracilifemur sp.n.

RMNH.INS.557960 Eurydiopsis sarawakensis

RMNH.INS.555162 Cyrtodiopsis nr. dalmanni sp. 1

Teleopsis longiconus sp.n.V1

RMNH.INS.557903 Teleopsis sp. 2

RMNH.INS.555160 Teleopsis sabah sp. 1

Cyrtodiopsis nr. dalmanni sp.1 IIX3

0.52

0.93

1

0.87

0.88

1

1

1

1

0.98

1

1

1

0.98

0.73

1

0.7

0.89

1

1

0.97

1

3500-4000m3000-3500m2500-3000m2000-2500m1500-2000m1000-1500m500-1000m0-500m

Extended Data Figure 2 | Phylogenetic reconstructions for Coleoptera, Dicranaceae, Diptera and Cortinariaceae. Details of materials and methods used aregiven in Supplementary Table 2.

RESEARCH LETTER


Ericaceae - Rhododendron

0.0070

SNP28518 R. sp.

SNP21944 R. praetervisum

SNP21930 R x. planecostatum

SNP28503 R. fallacinum

SNP21939 R. javanicum

SNP32306 R. buxifolium

SNP21928 R. fallacinum SNP21931 R. fallacinum

SNP21940 R. crassifolium

SNP32302 R. ericoides

SNP21943 R. suaveolens

SNP21910 R. stapfianum

SNP21912 R sp.SNP21918 R. cf. rugosum

SNP21927 R stenophyllum


SNP32303 R. ericoides

SNP32305 R. lowii




SNP21911 R. exuberans

SNP21925 R. simulans


SNP21917 R. sp.

SNP21932 R. praetervisum

SNP28504 R. rugosum

SNP21907 R. bagobonum

SNP28515 R. cuneifolium

SNP28509 R. acuminatum

GU176637 AF394258 R. camtscha�cum


SNP28508 R. rugosum


SNP21955 R. orbiculatum





SNP28517 R. stenophyllum


SNP21936 R. borneense

SNP21908 R. bur�i

SNP28510 R. rugosum


SNP21926 R. cf. stapfianum

SNP21906 R. rugosum

SNP28516 R. cf. cuneifolium




SNP21916 R p.


SNP21933 R. orbiculatum




SNP28513 R. lowii


SNP21909 R cf. rugosum




0.89

0.5

1

0.55

0.99

0.62

0.97

0.99

1

0.91

1

0.93

0.57

1

0.56

1

0.76

0.91

0.87

0.99

1

1

0.95

0.53

0.89

Gastropoda - Philalanka

0.05

SP13510 Philalanka sp.2

SP13515 Philalanka thienemanni


JX988066 Subulina octona




HQ171582-Zospeum exiguum (Slovenia)




HQ171529 Carychium mexicanum (USA)

SP13428 Ibycus rachelae








1

1

1

0.97

1

1

1

1

Glomeraceae - Glomus

0.3

OTU 20927

OTU 34472

AJ849467 Glomus xanthium

GU059549 Glomus indicum

OTU 20507

OTU 8741

OTU 5708

OTU 23633

FM992400 Glomus proliferum

OTU 21224

OTU 34383

FM865561 Glomus intraradices

AJ567764 Glomus intraradices


J437105 Glomus sinuosum

OTU 27453

OTU 24836

OTU 8898

OTU 6655

OTU 4196


OTU 27338

OTU 29948

OTU 33992

OTU 12813

OTU 20709

OTU 26364

AM713407 Glomus eburneum

OTU 8632

FJ009600 Glomus intraradices

OTU 35053

OTU 3458


OTU 16590

OTU 7081

OTU 29995

EF989103 Glomus intraradices

OTU 5441

AJ972462 Glomus diaphanum

OTU 21220


FJ009598 Glomus intraradices

OTU 5233

OTU 12112OTU 24971

OTU 18850

OTU 13666

OTU 8446


OTU 27919

OTU 17823

OTU 32979

OTU 5542

EF989105 Glomus intraradices

OTU 6588

OTU 21269

AJ437106 Glomus sinuosum

OTU 1606


OTU 21530

OTU 9223

OTU 29430

AM713409 Glomus eburneum

OTU 14114

OTU 10293

OTU 11936AY035654 Glomus clarum

OTU 13007

43100

92

100

60

31

10

2533

8866

98

46

30

100

79

32

18

90

56

96

59

3037

100

35

65

42

69

40

19

7322

14

87

57

55

64

27

68

60

80

93

57

33

94

77

70

44

51

24

25

87

98

37

37

48

89

36

45

99

56

56

74

12

23

95

100

100

Hirudinea - Tritetrabdella

0.04

SP13306 T. kinabaluensis kinabaluensis-

HQ322464 T. taiwana (Taiwan)


SP13382 T. kinabaluensis inobongensis

HQ203161 P. pungens (Australia)

HQ203195 T. taiwana (China)

SP13407 H. sp.


HQ203194 T. scandens (Thailand)



AB685259 T. taiwana (China)


SP13398 T. kinabaluensis kinabaluensis





0.58

1

1

1

1

0.99

1

1

1

0.54

Hemiptera - Rhagovelia

0.04

RMNH.INS.555251 R. samardaca

RMNH.INS.555228 R. silau

RMNH.INS.555449 R. simulata







RMNH.INS.555452 Mesovelia horvathi




RMNH.INS.555212 R. ranau



























1

1

0.95

0.54

0.99

0.91

1

0.97

0.85

0.62

0.97

1

1

0.73

0.97

3500-4000m3000-3500m2500-3000m2000-2500m1500-2000m1000-1500m500-1000m0-500m

Extended Data Figure 3 | Phylogenetic reconstructions for Ericaceae, Gastropoda, Glomeraceae, Hemiptera and Hirudinea. Details of materials andmethods used are given in Supplementary Table 2.

LETTER RESEARCH


Nepenthaceae - Nepenthes

0.2

L.4149375-6 N. lowii

L.4149364 N. ampullaria

L.4149374 N. tentaculata

L.4149112 N. mirabilis

L.4149369 N. chaniana


L.4149384 N. gracilis

L.4149394 N. x kinabaluensis





L.4149389 N. villosa


L.4149117 N. edwardsiana






L.4149373 N. lowii





L.4149380 N. fusca L.4149381 N. fusca

L.4149355-6 N. x harryana

L.4149089 N. mirabilis L.4149365 N. gracilis


AF315885 N. pervillei (Seychelles)


L.4149359 N. x hookeriana







SNP34297 N. rajah

L.4149378 N. lowii








L.4149074-77 N. edwardsiana


L.4149124 N. lowii





L.4149391 N.villosa

L.4149382 N. fusca


L.4149368 N. fusca







L.4149370 N. chaniana







0.99

1

0.99

0.970.99

1

1

0.83

1

1

1

0.99

0.98

Odonata - Coeliccia

0.04

RMNH.INS.507662 C. sp.

RMNH.INS.503978 C. octogesima (Singapore)

RMNH.INS.500047 Calicnemia rectangulata (Pahang Malaysia)

RMNH.INS.501058 C. albicauda (Malaysia)

RMNH.INS.503926 C. didyma (Terengganu Malaysia)

RMNH.INS.509175 C. poungyi (Thailand)

RMNH.INS.507728 C. nemoricola







RMNH.INS.506487 C. cyanomelas (Hong Kong)

RMNH.INS.228944 C. octogesima (Malaysia)




RMNH.INS.503932 C. didyma (Terengganu Malaysia)







RMNH.INS.506836 C. octogesima (Malaysia)


RMNH.INS.506294 C. nemoricola (Sarawak Malaysia)

RMNH.INS.506304 C. nemoricola (Sarawak Malaysia)





1

1

1

1

0.98

0.9

0.98

0.72

0.83

1

0.74

10.95

0.99

1

0.55

1

1

1

Polygalaceae - Epirixanthes

0.04

CM011 (L) E. papuana

WJJO de Wilde & BEE de Wilde-Duy�es 21409 (L) Salomonia cantoniensis

CM006 (L) E. pallida

CM030 (L) E. elongata

CM001-2 (L) E. pallida

CM009 (L) E. pallida


CM013 (L) E. kinabaluensis


CM032 (L) E. elongata


K'tung 78 6390 Salomonia ciliata

0.69

1

1

1

0.951

Rubiaceae - Hedyotis

0.0020

bZ030 H. valetoniana (NewGuinea)

FL038 H. pulchella

FL035 H. pulchella

FL011 H. sp. (Philippensis )

FL039 H. pulchella

FL058A H. macrostegia

FL052 H. macrostegia

FL037 H. pulchella

FL068 H.pulchella


FL077 H. pulchella



FL042 H. pulchella

cB019 H. fissis�pula (Borneo)


cA092 H. schlechteri (NewGuinea)

FL031 H. sp. (Philippensis)

FL047 H. pulchella. var. magnis�pula

cB042 O. pubescens (NewGuinea)

FL046 H. pulchella

FL048 H. pulchella. var. magnis�pula

1

1

0.91

1

1

1

1

0.71

1

1

1

Trombidiformes - Hygrobates

0.2

RMNH.ACA.1362 H. acu�palpis

RMNH.ACA.1370 H. hamatus

RMNH.ACA.1353 H. hamatoides


RMNH.ACA.1429 Atrac�des cognatus


RMNH.ACA.1360 H. acu�palpis

RMNH.ACA.1438 Atrac�des cf. propatulus

RMNH.ACA.1364 H. hamatus




1

1

0.74

1

3500-4000m3000-3500m2500-3000m2000-2500m1500-2000m1000-1500m500-1000m0-500m

Extended Data Figure 4 | Phylogenetic reconstructions for Nepenthaceae, Odonata, Polygalaceae, Rubiaceae and Trombidiformes. Details of materials andmethods used are given in Supplementary Table 2.

RESEARCH LETTER


Cornopteris_christenseniana

Athyrium_nikkoense

Athyrium_cuspidatum

Athyrium_x_akiense

Woodwardia_japonica

Athyrium_kuratae

Athyrium_oblitescens

Diplazium_simplicivenium

Athyrium_x_hisatsuanum

Athyrium_viridescentipe

Deparia_lobatocrenata

Athyrium_subrigescens

Athyrium_brevifrons

Woodsia_polystichoides

Diplazium_hachijoense

Athyrium_niponicum

Athyrium_rupestre

Athyrium_arisanense

Athyrium_setuligerum

Athyrium_amoenum

Athyrium_nakanoi

Deparia_conilii

Athyrium_atkinsonii

Homalosorus_pycnocarpos

Athyrium_otophorum

Thelypteris_palustris

Diplazium_nipponicum

Cornopteris_decurrenti

Athyrium_vidalii

Athyrium_sheareri

Athyrium_x_tokashikii

Deparia_petersenii

Cornopteris_crenulatoserrulata

Athyrium_eremicola

Blechnum_spicant

Hymenasplenium_unilaterale

Athyrium_yokoscense

Athyrium_strigillosum

Athyrium_clemensiae

Athyrium_clivicola

Athyrium_wardii

Asplenium_bulbiferum

Athyrium_kirisimaense

Athyrium_melanolepis

Athyrium_iseanum

Athyrium_tashiroi

Athyrium_filix_femina

Athyrium_deltoidofrons

Athyrium_tozanense

Athyrium_neglectum

1,28

9,4

12,01

0,88

34,28

5,25

29,06

4,19

7,13

75,08

2,64

11,86

58,63

0,96

16,83

51,58

21,17

6,55

2,7

53,85

10,22

0,44

0,7

1,58

8,99

67,59

1,41

0,43

7,42

2,15

13,5

17,95

40,21

14,53

81,52

4,23

10,72

14,96

18,79

2,32

99,02

8,51

9,68

62,42

0,53

2,39

0,23

1,27

71,74

Begonia_kingiana

PET_stevei07_01455

PET_spec1Borneo07_00575

PAR_elisabethae10_00618

Begonia_hatacoa

PET_spec1Papua08_02330

Begonia_robusta

PET_bonthainensis09_01403

Begonia_masoniana

Begonia_fenicis

FL071_Begonia_beryllae_concat_D345

DIP_demissa10_00617

Begonia_silletensis

Begonia_cleopatrae

PET_didyma08_02240

Begonia_burbidgei_D363_FL040_concat

PET_hispidissima_GLombulan09_01418PET_capituliformis09_01419

ALI_alicida10_00614

Begonia_sudjanae

Begonia_chloroneura

Begonia_brevirimosa

Begonia_symsanguinea

PET_torajana1109_01410

REI_hymenophylla10_00613

SPH_obovoidea10_00621

PET_prionota_spec809_01407

Begonia_pavonina

FL074_Begonia_cf_beryllae_cf_D347

Begonia_nigritarum

Begonia_doloisii_concat_D350_PH115

TET_polygonoides09_02176ndhA_ndhF_rpl32

Begonia_venusta

Begonia_multijugata

Begonia_varipeltata

Begonia_siccacaudata

Begonia_bracteata

PET_aff_congesta09_00060

PET_vermeulenii_07_01837

Begonia_sutherlandii

PET_hekensis08_02227

PET_macintyreana07_01454

BAR_boliviensis09_02161phu_ndhA_ndhF_rpl32

Begonia_burbidgei_DCT2012

PET_specSumbawa09_01015

Begonia_argenteomarginata

Begonia_spec_China2

Begonia_strigosa

Begonia_koordersii

PET_specKalimantan09_01013

Begonia_vaccinioides_D351_ndhA_concat

Begonia_laruei

PET_ozotothrix08_02845

PET_wrayi10_00642_extraction

Begonia_aptera

Begonia_cauliflora_aff_D364_FL066_concat

Begonia_palmata

Begonia_serratipetala

PET_weigallii09_01017

Begonia_aff_elisabethae

Begonia_goegoensis

DIP_spec_Thailand310_00624

BRA_specSumatra08_02319

PET_chiasmogyna07_00577

Begonia_poculifera

Begonia_socotrana

PET_rantemarioensis09_01405

PET_nobiaespec1709_01416

Begonia_sikkimensis

Begonia_verecunda

Begonia_longifolia

Begonia_dregeiSOL_radicans09_02177ndhA_ndhF_rpl32

Begonia_dipetala

Begonia_muricata

Begonia_amphioxus_concat_DCT2012

PET_mendumiae07_00578

PET_pseudolateralis08_02220ndhA_ndhF_rpl32

Begonia_multangula

PET_aff_mekonggensisspec1209_01411

Begonia_hernandioides

PET_specPhilippines08_03026

Begonia_versicolor

Begonia_spec_Sumbawa1

Begonia_guttapila

Begonia_tenuifolia

PET_harauensis09_01011

Begonia_sizemoreae

Begonia_malabarica

Begonia_watuwilensis

Begonia_imbricata_D365_FL075_concat

IGN_smithiae10_00615

Begonia_samahensis

Begonia_areolata

Begonia_negrosensis

Begonia_grandis

Begonia_poliloensis

PET_comestibilis_spec509_01402

PET_sanguineopilosaspec1809_01417

PAR_spec_Thailand110_00620

DIP_spec_Thailand210_00622

Begonia_aff_multangula

Begonia_decora

Begonia_chlorosticta

Begonia_oxyloba_ndhA_ndhF_rpl32

PET_flacca09_01009PET_lasioura_spec1309_01412

Begonia_acetosella

Begonia_corrugata

PET_specNewGuinea09_01016

Begonia_spec_China1

FL062_Begonia_inobongensis_concat_D342

Begonia_morsei

BRA_resecta08_00204

QUA_goudotii09_02167ndhA_ndhF_rpl32

REI_brandisiana10_00619

DIP_aceroides10_00623

GIR_nelumbifolia09_02171ndhA_ndhF_rpl32

Begonia_masarangensis

Begonia_roxburghii

Begonia_vaccinioides_D353_concat

FL065_Begonia_cauliflora_aff_concat_D344

Begonia_floccifera

Begonia_pendula_DCT2012

Begonia_flagellaris

2,09

11,67

0,78

0,29

2,23

0,56

0,68

3,22

0,99

0,74

2,83

6,63

1,73

1,04

1,35

7,18

2,69

1,41

5,6

13,9

0,43

15,11

3,67

3,74

6,2

2,2

2,15 1,12

1,15

0,15

2,73

2,03

0,93

4,26

3,44

10,01

13,02

5,61

3,37

0,34

22,41

6,6

1,45

4,61

1,6

0,47

5,16

9,04

6,03

0,58

3,18

2,15

0,58

0,7

0,24

3,17

2,53

2,09

0,51

1,12

0,16

4,67

3,55

9,27

4,38

9,58

2,4

2,48

6,5812,54

2,15

0,54

2,66

0,67

2,26

10,87

1,47

0,85

10,12

3,54

7,38

5,67

3,4

0,09

10,18

4,58

1,67

10,75

0,13

15,76

18,37

17,14

14,62

2,94

18,71

1,33

1,06

4

9,39

4,33

1,4

4,11

4,55

0,92

12,34

4,83

4,61

1,88

2,99

4,95

1,47

3,9

1,42

10,61

4,15

2,9

4,13

11,35

1,81

1,23

5,93

12,61

7,59

3,15

Prumnopitys_ferruginea

Dacrydium_balansae

Retrophyllum_minus

Podocarpus_spinulosus

Podocarpus_sp_PK_2012a

Prumnopitys_ladei

Afrocarpus_usambarensis

Phyllocladus_toatoa

Lagarostrobos_franklinii

Podocarpus_neriifolius

Saxegothaea_conspicua

Podocarpus_sp_PK_2012b

Dacrydium_beccarii

Dacrycarpus_kinabaluensis111

Podocarpus_parlatorei

Podocarpus_hallii

Phyllocladus_alpinus

Prumnopitys_taxifolia

Nageia_fleuryi

Podocarpus_pilgeri

Podocarpus_nubigenus

Podocarpus_brassii

Podocarpus_sylvestris

Podocarpus_bracteatus

Podocarpus_purdieanus

Dacrycarpus_imbricatus105

Podocarpus_angustifolius

Podocarpus_henkelii

Podocarpus_lucienii

Podocarpus_sellowii

Nageia_nagi

Podocarpus_latifolius

Podocarpus_oleifolius

Manoao_colensoi

Podocarpus_forrestii

Afrocarpus_falcatus

Podocarpus_elongatus

Podocarpus_drouynianus

Podocarpus_celatus

Dacrycarpus_dacrydioides

Podocarpus_ramosii

Podocarpus_totara

Podocarpus_milanjianus

Podocarpus_novae-caledoniae

Falcatifolium_papuanum

Podocarpus_polystachyus

Nageia_formosensis

Podocarpus_subtropicalis

Podocarpus_rubens


Dacrydium_elatum

Podocarpus_gnidioides

Podocarpus_lawrencei

Nageia_wallichiana

Falcatifolium_taxoides

Prumnopitys_andina

Podocarpus_madagascariensis

Microcachrys_tetragona

Podocarpus_dispermus

Phyllocladus_hypophyllus

Retrophyllum_rospigliosii

Podocarpus_lambertii

Prumnopitys_ferruginoides

Afrocarpus_gracilior

Podocarpus_nivalis

Dacrydium_guillauminii

Dacrycarpus_compactus

Podocarpus_acutifolius

Phyllocladus_trichomanoides


Phyllocladus_aspleniifolius

Podocarpus_degeneri

Podocarpus_costalis


Podocarpus_grayae

Podocarpus_sprucei

Podocarpus_chingianus

Podocarpus_elatus

Podocarpus_salignus

Podocarpus_cunninghamii

Podocarpus_guatemalensis

Sundacarpus_amarus

Afrocarpus_dawei

Podocarpus_matudae

Podocarpus_urbanii

Acmopyle_sahniana

Afrocarpus_mannii

Retrophyllum_comptonii

Podocarpus_affinis

Dacrydium_cupressinum

Podocarpus_macrophyllus

Podocarpus_pallidus

Pherosphaera_fitzgeraldii

Podocarpus_archboldii

Podocarpus_coriaceus

Podocarpus_longifoliolatus

Podocarpus_ledermannii

Podocarpus_smithii

Podocarpus_fasciculus

Dacrycarpus_expansus

Lepidothamnus_intermedius

Podocarpus_rumphii

3,19

5,22

4,84

61,25

118,87

2,25

76,41

87,55

6,91

8,64

12,91

29,01

24,03

9,5

0,66

1,06

68,15148,07

12,56

8,62

22,13

20,01

1,48

0,97

9,5

7,08

7,25

3,43

1,18

10,41

35,88

14,59

127,01

11,04

48,76

34,37

14,87

161,21

1,28

33,15

113,14

3,78

1,28

5,15

30,28

17,46

1,3

15,69

15,21

8,95

13,03

18,13

102,93

1,52

5,07

2,78

6,94

57,28

29,17

43,55

5,92

15,15

7,5825,45

10,32

0,76

0,54

7,83

43,96

1,79

19,89

4,85

3,02

1,57

9,41

1,36

93,46

1,62

60,82

21,65

6,54

11,73

4,07

1,18

14,93

15,66

9,51

6,19

5,65

3,59

8,87

1,17

0,93

6,27

1,63

2,65

10,43

19,19

0,55

0,83

24,99

111

Athyrium

Begonia

Dacrycarpus

Coeliccia_flavostriata

RMNH_INS_507675_C_species_1_500to750mASL_PHS

RMNH_INS_503926_C_didyma_Terengganu_Malaysia

RMNH_INS_509175_C_poungyi_Thailand

RMNH_INS_503977_C_octogesima_Singapore

Coeliccia_borneensis

RMNH_INS_507768_C_nemoricola_500to750mASL_IN

Coeliccia_cyaneothorax

Coeliccia_ryukyuensis

RMNH_INS_506468_C_cyanomelas_HongKong

RMNH_INS_500047_Calicnemia_rectangulata_Pahang_Malaysia_OUTGROUP

Calicnemia_sinensis

Calicnemia_chaseni

Coeliccia_flavicauda

RMNH_INS_501058_C_albicauda_Malaysia

RMNH_INS_507662_C_species_2_1500to1750mASL_KHQ

2,11

0,26

2,1

1,19

1,75

2,92

2,26

1,68

0,85

1,51

2,17

1,81

3,82

2,73

4,44

Microtis

Platythelys

Maxillaria

Musella_lasiocarpa

Neuwiedia

Elaeis_oleifera

Dendrobium_officinale

Platanthera

Epipactis

Masdevallia

Cymbidium

Selenipedium

Mormodes

Listera

Ludisia

Wullschlaegelia

ChloraeaPterostylisMegastylis

Lanaria

Vitekorchis

Pogonia

Orchis

Hypoxis

DiurisChiloglottis

Dendrobium_kingianum

Agrostophyllum_majus

Cypripedium

Cleistes

Codonorchis

Phalaenopsis

Bifrenaria

Phragmipedium

Palmorchis

Pachyplectron

Nypa_fruticans

Eria

Vanilla

Altenstenia

Apostasia

Eriochilus

StanhopeaGongora

Astelia

Pleione

Sarcoglottis

Nervilia

Exalaria

Paphiopedilum

Empodium

Zygopetalum

Rhodohypoxis

Ponthieva

Habenaria

Cattleya

Earina_autumnalis

Monadenia

Spiranthes

Sobralia

Goodyera

Galeandra

Anigozanthos_flavidus

Dossinia

Earina_valida

Dendrochilum_glumaceumDendrochilum_scriptum

DisperisDisa

Dendrobium_crystallinum

Lycaste

Dendrochilum_cobbianum

Calanthe

Coelogyne_cristata

22,78

33,32

77,54

39,33

54,79

31,03

7,64

37,86

32,69

25,34

58,84

23,91

32,87

32,34

40,35

60,29

50,15

18,6123,13

23,76

8,73

12,7

9,21

35,81

66,17

16,71

35,08

41,37

71,7

34,34

107,11

14,69

33,36

13,61

83,63

32

49,86

49,73

17,13

45,06

42,67

36,41

25,83

23,48

32,84

27,82

7,66

42,7

57,81

38,38

25,37

35,38

11,63

45,44

16,38

23,14

2,18

19

17,05

102,79

34,71

29,37

24,46

19,66

13,87

16,46

27,06

14,35

40,12

69,66

62,55

40,24

19,91

Dendrochilum_cobbianum

Dendrochilum_alatum

Dendrochilum_glumaceum

Dendrochilum_ovatum

Dendrochilum_exasperatum

Dendrochilum_haslamii

Dendrochilum_graciliscapum

Dendrochilum_trusmadiense

Dendrochilum_arachnites

Dendrochilum_grandiflorum

Dendrochilum_muluense

Dendrochilum_pterogyne

Dendrochilum_stachyodes

Dendrochilum_dewindtianum

Dendrochilum_gibbsiae

Dendrochilum_dewildei

Dendrochilum_alpinum

Dendrochilum_pseudoscriptum

Dendrochilum_kamborangense

Dendrochilum_karoense

Dendrochilum_cupulatum

Dendrochilum_corrugatum

Dendrochilum_scriptum

Dendrochilum_tenompokense

Dendrochilum_transversum

Dendrochilum_joclemensii

Coelogyne_cristata

5,38

0,43

0,05

0,42

0,94

0,51

2,65

2,92

0,6

8,83

4,93

1,29

0,35

1,09

1,31

0,4

1,6

0,1

0,71

2,31

0,3

3,51

0,4

1,71

0,53

2,69

Polygala_paniculata

Epirixanthes_elongata

Salomonia_cantoniensis

Polygala_californica

Muraltia_karroica

Polygala_senega

Carpolobia_alba

Xanthophyllum_sp

Polygala_myrtifolia

Monnina_xalapensis

Polygala_vulgaris

Bredemeyera_floribunda

Eriandra_fragrans

Nylandtia_spinosa

Atroxima_afzeliana

Polygala_amara

Salomonia_ciliata

Comesperma_esulifolium

Polygala_longicaulis

Polygala_comosa

Polygala_alpicola

Epirixanthes_kinabaluensis

Polygala_costaricensis

Polygala_paucifolia

Securidaca_retusa

Securidaca_diversifolia

Polygala_chamaebuxus

Epirixanthes_pallida

Epirixanthes_papuana

Securidaca_longepedunculata

Polygala_violacea

19,67

35,75

31,93

18,16

7,23

30,7

46,78

9,72

0,83

3,07

36,67

25,68

16,02

16,03

28,69

13,7

29,97

11,1

13,34

24,5

29,43

33,53

9,54

7,22

2,11

42,16

54,58

5,6815,87

28,09

Coeliccia

Orchidaceae – Dendrochilum

Dendrochilum

Epirixanthes

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

MS48_e4002189827_D_assimile_Kinabalu_Gunung_Alab

MS3_e4002189762_D_brevisetum_Kinabalu_Sayap

DQ462598_DQ464188_D_robustum_New_Zealand

L0856682_e4002189720_D_havilandii_Kinabalu_Layang_Laya

MS53_e4002189792_D_sp_Kinabalu_Gunung_Alab


KF423978_KF423621_D_cylindrothecium_Russia

DQ485179_D_menziesii_Australia

MS29_e4002189789_D_assimile_Kinabalu_KNP

MS44_e4002189779_D_havilandii_Kinabalu_Laban_Rata

MS52_e4002189780_D_blumii_Kinabalu_Gunung_Alab

L0092869_e4002189708_D_diaphanoneurum_Australia

MS17_e4002189740_D_braunii_Kinabalu_Sayap


DQ462594_DQ464193_D_billarderi_New_Zealand

DQ462596_DQ464196_D_billarderi_Rwanda

L0856640_e4002189709_D_sp_Indonesia_Java


DQ462595_DQ464194_D_billarderi_Chile

AF135075_AF144116_Paraleucobryum_enerve


KF423995_KF423641_Holomitrium_arboreumKF423992_KF423638_Holomitrium_crispulum



DQ462606_DQ464185_D_plurisetum_New_Zealand


DQ462607_DQ464184_D_menziesii_New_Zealand


DQ485174_D_plurisetum_New_ZealandMS25_e4002189741_D_assimile_Kinabalu_Sayap

MS12_e4002189775_D_reflexum_Kinabalu_Sayap



DQ462600_DQ464190_D_robustum_Chile

DQ462605_DQ485175_D_dicarpum_New_Zealand

MS20_e4002189776_D_blumii_Kinabalu_L2

MS38_e4002189802_D_assimile_Kinabalu_Laban_Rata

MS27_e4002189765_D_assimile_Kinabalu_Sayap

AF146376_AF144112_Mesotus_celatus_OUTGROUP

DQ485176_D_leichhardtii_Australia

MS35_e4002189766_D_braunii_Kinabalu_L2MS19_e4002189764_D_braunii_Kinabalu_Sayap


AF129587_AF144113_Dicranum_polysetum

L0093198_e4002189685_D_reflexum_Indonesia_Sumatra

DQ462604_DQ464186_D_chilense_Chile

MS39_e4002189814_D_blumii_Kinabalu_Laban_Rata

MS37_e4002189790_D_assimile_Kinabalu_L2MS33_e4002189742_D_assimile_Kinabalu_KNP

AF135067_AF140696_Dicnemon_calycinum


DQ485178_D_fasciatum_Australia

DQ462597_DQ464197_D_billarderi_Reunion





0,08

0,210,07

2,33

1,790,68

0,39

0,04

3,26

0,05

0,12

0,71

0,04

0,38

1,49

0,23

0,13

0,18

3,51

0,67

0,16

0,03

2,69

1,1

0,05

1,81

0,32

1,25

0,81

0,08

0,13

0,93

0,53

0,27

0,95

0,03

0,42

0,05

0,09

0,74

0,33

0,04

0,03

0,03

0,38

0,8

0,1

0,2

0,05

0,61

0,38

1,83

0,61

0,12

0,1

0,03

1,14


AF135067_AF140696_Dicnemon_calycinum

DQ485176_D_leichhardtii_Australia


DQ462604_DQ464186_D_chilense_Chile



MS53_e4002189792_D_sp_Kinabalu_Gunung_Alab

MS38_e4002189802_D_assimile_Kinabalu_Laban_Rata

DQ462607_DQ464184_D_menziesii_New_Zealand

MS35_e4002189766_D_braunii_Kinabalu_L2

DQ462597_DQ464197_D_billarderi_Reunion

DQ485178_D_fasciatum_Australia

MS39_e4002189814_D_blumii_Kinabalu_Laban_Rata

DQ485179_D_menziesii_Australia


DQ462595_DQ464194_D_billarderi_Chile


MS27_e4002189765_D_assimile_Kinabalu_Sayap

KF423995_KF423641_Holomitrium_arboreum

AF146376_AF144112_Mesotus_celatus_OUTGROUP

KF423992_KF423638_Holomitrium_crispulum

L0856682_e4002189720_D_havilandii_Kinabalu_Layang_Laya




AF135075_AF144116_Paraleucobryum_enerve




L0856640_e4002189709_D_sp_Indonesia_JavaL0093198_e4002189685_D_reflexum_Indonesia_Sumatra



DQ462596_DQ464196_D_billarderi_Rwanda


DQ462606_DQ464185_D_plurisetum_New_Zealand


MS25_e4002189741_D_assimile_Kinabalu_SayapDQ485174_D_plurisetum_New_Zealand


L0092869_e4002189708_D_diaphanoneurum_Australia


DQ462605_DQ485175_D_dicarpum_New_Zealand

MS52_e4002189780_D_blumii_Kinabalu_Gunung_Alab


MS20_e4002189776_D_blumii_Kinabalu_L2





DQ462594_DQ464193_D_billarderi_New_Zealand


MS37_e4002189790_D_assimile_Kinabalu_L2


KF423978_KF423621_D_cylindrothecium_Russia

AF129587_AF144113_Dicranum_polysetum


0,79

0,2

0,05

3,53

0,93

0,05

2,69

0,33

0,14

0,35

1,13

0,05

0,57

2,78

0,12

0,18

2,75

0,58

1,18

0,14

0,08

0,21

0,08

0,49

0,1

0,07

1,08

0,06

1,45

1,88

4,98

0,28

1,03

0,12

0,93

1,66

0,20,07

0,08

2,26

1,41

0,15

1,04

0,24

0,59

1,67

0,42

0,64

0,08

0,050,3

1,23

0,06

5,38

0,57

4,08

0,52

Dicranoloma (ITS rate fi xed)

Dicranoloma (trnL rate fi xed)

Iteitensis

Iuliginosa

Iraphidotrix

Imildbraediitelekii

Ihochstetteri_hochstetteri

Itrichosepala

Ikeillii_pubescens

Iflanaganae

Iethiopica

Ifenghwaiana

Iarachnoides

Ihochstetteri_jacquesii

Ipurpurea

Ikamerunensis_obanensis

Ihawkeri

Ipoilanei

Icampanulata

Iengleri_engleri

Ibalsamina

Icongolensis

Imeruensis_meruensis

Inana

Imengtseana

Ikamerunensis_kamerunensis

Iaureliana

Iclavicornu

Ibururiensis

Ihamata

Iviscida

Imannii_WestAfrica

Isiculifer

Iirvingii

Iinaperta

Iassurgens

Inolitangere

Iaquatilis

Isakerana

Irubromaculata_rubromaculata

Ibalfourii

Iscabrida

Ihoehnellii

Iedgeworthii

Ihians

Imannii_CentralAFrica

Ilukwangulensis

Ifurcata

Imacroptera

Ikinabaluensis

Iomeiana

Iyingjiangensis

Irubrostriata

Ipalpebrata

Icapensis

Ibombycina

Isodeni

Imackeyana_zenkeri

Isylvicola

Ichungtienensis

Itinctoria_elegantissima

Ihydrogetonoides

Ieberhardtii

Iuniflora

Ikeillii_keillii

Iniamniamensis

Inapoensis

Iplatypetala

Ibegonifolia

Iforrestii

Ibequartii

Ifaberi

Itaronensis

Ikilimanjari

Imazumbaensis

Iimbicilla

Iglandulifera

Ipseudohamata

Ipseudomacroptera

Iusambarensis

Ihenslowiana

Ikerriae

Ituberosa

Iconchibracteata

Ieryaleia

Iauricoma

Imeruensis_cruciata

Irectangula

Ithamnoidea

Icyanantha

Ichinensis

Ibriartii

Iulugurensis

Imonticola

Icatati

Ibicaudata

Icuspidata

Ipalliderosae_palliderosae

Idesmantha

Ipallida

Imackeyana_claeri

Ipercordata_percordata

Iwalleriana

Irubromaculata_imagiensis

Ioumina

Ipseudoviola

Ilatifolia

Ipseudozombensis

Iconfusa_longicornu

Iparviflora

Igordoni

Icecilii

Iburtoniiburtonii

Iflaccida

Iioides

Idigitata_digitata

Ivolkensii

Idavidi

Ichevalieri

Icolumbaria

Iserpens

Iparasitica

Ifischeri

Ipalliderosae_lupangaensis

Itweediae

12,67

1,77

2,66

1,36

2,53

3,6

3,21

6,49

1,22

4,1

1,86

1,92

9,1

12,81

1,13

3,14

1,38

3,05

0,58

4,46

0,81

0,47

0,51

3,24

4,27

0,83

3,79

2,48

1,77

0,36

4,5

0,61

2,87

0,96

1,74

2,97

0,42

1,94

0,353,27

5,21

0,98

4,54

2,06

4,63

0,73

3,49

13,69

0,62

3,51

3,35

1,29

1,89

0,19

4,15

1,31

3,69

2,31

1,05

5,92

4,03

1,47

0,93

2,16

0,44

2,69

1,96

1,02

0,73

4,62

2,28

3,43

4,83

4,36

5,31

3,46

2,43

1,64

0,56

3,13

1,37

3,24

6,68

17,38

6,18

1,18

2,27

1,81

4,24

0,724,79

5,21

19,22 0,52

0,77

5,62

1,01

5,5

1,65

1,77

0,63

4,13

0,33

5,5

1,12

1,08

3,465,76

0,23

9,69

0,92

1,02

9,7

1

8,32

3,18

1,34

0,71

7,82

0,58

0,18

8,47

Impatiens

posterior1

0

posterior1

0

posterior1

0

posterior1

0

Cordylostigma_virgata_cB032

Hedyotis_lindleyana_cB023_Asia

Oldenlandia_paridifolia_TM3167_Chin

Hedyotis_membranacea_SN78_India

Hedyotis_quinquenervia_aT97_Ceylon

Oldenlandia_erecta_SN2_Asia

Neanotis_indica_cC030_India

Kadua_affinis

Hedyotis_schlechteri_cA092_NewGuine

Hedyotis_lawsoniae_aT98_Ceylon

Agathisanthemum_quadricostatum_cC03

Oldenlandia_angolensis

Oldenlandia_tenelliflora_cA089_Asia

Oldenlandia_pulcherrima

Hedyotis_sp_bZ020_Vietnam

Leptopetalum_foetidum_cB044_Pacific

Oldenlandia_benguetensis_cB026_SEAs

Agathisanthemum_chlorophyllum_cB031

Oldenlandia_assimilis

Oldenlandia_contracta_bZ008

Agathisanthemum_globosum

Hedyotis_capitellata_bR98_SEAsia

Oldenlandia_bracteosa_cB004_China

Oldenlandia_nervosa

Houstonia_caerulea

Spermacoce_hispida

Hedyotis_rhinophylla_aT90_Ceylon

Oldenlandia_galioides

Hedyotis_trichoclada_cB029_NewGuine

Hedyotis_purpurascens_cC032_India

Oldenlandia_chrysotricha_bR100_Chin

Oldenlandia_cherevensis_cA054_SEAsi

Oldenlandia_brachypoda_SN88_Asia

Oldenlandia_assimilis_cB055_China

Hedyotis_cantoniensis_cA056_China

Oldenlandia_herbacea_var_herbacea

Oldenlandia_taborensis

Oldenlandia_cryptantha_TM3202_China

Hedyotis_megalantha_aT99

Hedyotis_acutangula_cB002_Asia

Oldenlandia_ovatifolia_bZ044_SEAsia

Hedyotis_caudatifolia

Oldenlandia_mitrasacmoides

Neanotis_nummulariformis_cC029_Ceyl

Hedyotis_swertioides_India

Hedyotis_pachycarpa_bZ011_Asia

Hedyotis_sp_cB011

Hedyotis_effusa

Oldenlandia_angustifolia_cB014_Indi

Hedyotis_ampliflora_cB056_China

Oldenlandia_ternata_bZ027_SEAsia

Oldenlandia_microtheca

Oldenlandia_pinifolia_bS031_IndiaSE

Neanotis_wightiana_bS011_IndiaSEAsi

Oldenlandia_gracilipes_cA061_Thaila

Hedyotis_sp_SKP946

Oldenlandia_lancea_cB003_China

Hedyotis_tridentata_bS36_Ceylon


Oldenlandia_umbellata_SN84_Asia

Hedyotis_ovata_VH048

Pleiocraterium_verticillare_bS39_In

Hedyotis_trimenii_bS37_India

Oldenlandia_lancifolia

Oldenlandia_echinulosa

Arcytophyllum_muticum

Oldenlandia_densa_robinsonii_bB92


Hedyotis_yangchunensis

Hedyotis_effusa_aT081_Asia

Lelya_osteocarpa

Hedyotis_communis_cA055_China

Oldenlandia_bodinieri_cB012_China

Hedyotis_nodulosa_bS26_Ceylon

Oldenlandia_uniflora

Lelya_prostrata_cB039

Leptopetalum_grayii_bZ041_Japan

Neanotis_hirsuta_bS009_Asia

Hedyotis_lessertiana_var_lessertian

Oldenlandia_capensis_var_capensis

Neanotis_gracilis_SN89_Asia

Hedyotis_lessertiana_var_marginata

Kadua_parvula

Hedyotis_fruticosa_aT85_India

Oldenlandia_stocksii_cC022_India

Hedyotis_cathayana_TM3214_China

Hedyotis_costata_bS41_Asia

Hedyotis_minutopuberula_TM3219_Chin

Stenaria_nigricans

Houstonia_longifolia

Oldenlandia_auricularia_cC021_Fiji

Oldenlandia_scabra_cB025_IndiaSEAsi

Metabolos_decipiens_cC017_Ceylon

Oldenlandia_tenuipes

Oldenlandia_tenelliflora

Oldenlandia_umbellata_cC025_Asia


Oldenlandia_tetrangularis_cB052_SEA

Hedyotis_coprosmoides_bS14_Ceylon

Hedyotis_cantoniensis

Hedyotis_scandens_cA052_Asia

Oldenlandia_herbacea_cB058

Oldenlandia_lineata_cB010_IndiaSEAs

Synaptantha_tillaeacea

Oldenlandia_erecta_cB024_Asia

Hedyotis_novoguinensis_cA093_NewGui

Arcytophyllum_thymifolium

Hedyotis_pitardiana_bZ023

Agathisanthemum_bojeri

Dibrachionostylus_kaessneri

Hedyotis_shiuyingiae_cB018_China

Oldenlandia_hainanensis_cB028_China

Oldenlandia_vachellii_cB007_China

Oldenlandia_consanguinea_aT79_China

Pentanopsis_fragrans

Paraknoxia_parviflora

Oldenlandia_salzmannii

Kohautia_cynanchica

Oldenlandia_biflora_bB067

Oldenlandia_diffusa_bS3_Asia


Hedythyrsus_spermacocinus

Amphiasma_merenkyanum_cB033

Hedyotis_decora_cB005_Thailand

Oldenlandia_wauensis

Oldenlandia_affinis_bZ039

Hedyotis_cheniana_TM3190_China

Hedyotis_coronaria_cB040_Asia

Oldenlandia_corymbosa_bB72

Oldenlandia_verticillata_SN5_Asia

Carphalea_madagascariensis

Oldenlandia_paridifolia_TM3165_Chin

Oldenlandia_chrysotricha_bR99_China

Hedyotis_stylosa_bS34_India

Hedyotis_obliquinervis_bZ009_Asia

Oldenlandia_lineata_bS6_IndiaSEAsia

Oldenlandia_trinervia_cC024_Asia

Hedyotis_caudatifolia_cB027_China

Hedyotis_sp_cA050

Hedyotis_philippensis_cB043_Asia

Mitrasacmopsis_quadrivalvis


Oldenlandia_auricularia_cB013_Asia

Oldenlandia_wiedemannii

Hedyotis_uncinella_bZ038_China

Hedyotis_macrostegia_Borneo

Batopedina_pulvinellata

Hedyotis_dianxiensis_bZ048_China

Pentodon_pentandrus_Zambia

Oldenlandia_tenuipes_bZ037_China

Cordylostigma_microcala

Hedyotis_valetoniana_bZ030_NewGuine

Hedyotis_shenzhenensis

Oldenlandia_stricta_cC023_India

Oldenlandia_rupicola_cC034_Africa

Oldenlandia_prostrata_bZ043_SEAsia

Neanotis_calycina_SN13_Asia

Neanotis_monosperma_cC027_India

Hedyotis_rigida_bZ042_SEAsia

Metabolos_rugosus_bS7_SEAsia

Hedyotis_pulchella_bZ029_Borneo

Hedyotis_articularis_bP48_India

Oldenlandia_melli_bZ036_China


Kohautia_coccinea

Oldenlandia_lapeyrousii_cA097_NewGu

Oldenlandia_hedyotidea_cB006_Asia

Hedyotis_fissistipula_cB019_Borneo

Oldenlandia_biflora

Neanotis_nummularia_cC028_Ceylon

Oldenlandia_auricularia_cC020_Ceylo

Hedyotis_thwaitesii_SN79_India

Oldenlandia_affinis

Kadua_fluviatilis

Hedyotis_vestita_bS40_Asia

Hedyotis_parryi_cB057_China

Neanotis_formosana_cC026_Taiwan

Hedyotis_pahompokae_bS042_Thailand

Oldenlandia_pterita

Oldenlandia_geophila

Hedyotis_nana_cA094_NewGuinea

Hedyotis_obscura_bS29_Ceylon

Oldenlandia_tenuis

Dentella_repens

Hedyotis_xinyiensis

Oldenlandia_duemmeri_cB041

Oldenlandia_rosulata

Oldenlandia_pubescens_cB042_NewGuin

Metabolos_decipiens_cC016_Ceylon

Oldenlandia_chrysotricha_bZ040_Chin

Oldenlandia_fastigiata

Hedyotis_marginata_SN72_Ceylon

Hedyotis_capitellata_bZ031_SEAsia

Hedyotis_dendroides_bS2_Ceylon

Hedyotis_korrorensis_aT83_Pacific

Hedyotis_rivalis_bZ035_Malaysia

Oldenlandia_consanguinea

Hedyotis_flavescens_SN61_Ceylon

Hedyotis_caudatifolia_cB037_China

Oldenlandia_nutans_bS27_NewGuinea

Oldenlandia_nematocaulis

Oldenlandia_goreensis

Hedyotis_gardnerii_bS016_Ceylon0,4

2,48

11,49

1,83

1,511,35

27,33

5,66

11,17

15,31

3,97

3,1

1,74

2,38

0,88

2,47

0,6

7,01

13,54

6,72

5,85

1,11

1,31

8,93

1,41

2,05

0,81

39,46

2,49

3,53

12,25

2,83

11,6

1,68

0,95

13,38

1,29

5,83

4,37

0,25

1,3

5,17

0,58

3,89

6,62

0,32

23,43

7,42

4,86

8,56

1,12

0,8

18,65

22,09

6,92

4,81

4,11

2,84

0,18

5,34

8,78

2,42

0,95

0,54

3,59

0,76

0,97

4,66

4,39

8,93

4,09

0,43

4,29

2,69

0,12

1,01

9,94

1,15

3,24

2,22

2,63

0,37

2,79

20,83

9,64

0,49

11,28

0,95

0,88

3,22

0,31

4,08

15,98

4,57

2,51

1,01

9,01

8,35

23,36

0,9

1,34

0,79

6,48

3,31

27,61

6,48

4,77

2,07

1,55

10,18

5,99

6,17

3,63

16,7

3,522,85

0,68

4,64

2,93

0,27

31,18

1,52

6,51

2,95

8,12

10,04

0,28

6,66

5,64

4,92

9,25

1,26

1,88

3,98

2,28

18,9

2,44

22,75

0,92

1,64

4,09

7,34

0,57

2,36

9,33

2,51

4,69

0,21

2,85

2,51

1,66

0,56

19,89

5,14

1,36

7,12

0,76

9,89

6,71

0,08

1,96

2,39

17,77

0,74

17,77

7,8

1,41

1,9

5,69

16,89

4,22

2,04

5,14

14,34

7,75

11,01

5,84

1,66

25,98

15,31

0,48

0,43

3,15

2,7

1,56

2,35

2,04

1,68

4,81

28,53

1,04

24,39

18,76

0,42

3,15

5,95

5,25

4,09

14,44

0,5

8,01

2,65

Hedyotis

Hygrobates_hamatoides

Hygrobates_hamatus

Hygrobates_hamatusH20

Atractides_cf_propatulus

Hygrobates_sp_2

Atractides_cognatus

2,9

1,23

3,72

9,55

1,71

Hygrobates

Ilex_zygophylla

Ilex_latifolia

Ilex_cissoidea

Ilex_wallichii2

Ilex_dumosa

Ilex_opaca

Ilex_rotunda2

Ilex_verticillata

Ilex_shennongjiaensis

Ilex_oppositifolia

Ilex_yunnanensis2

Ilex_ciliospinosa

Ilex_dugesii

Ilex_integra2

Ilex_spicata

Ilex_collina

Ilex_wallichii1

Ilex_chamaedryfolia

Ilex_theizans2

Ilex_leucoclada

Ilex_bioritsensis

Ilex_kusanoi

Ilex_teratopis

Helwingia_japonica

Ilex_maingayi

Ilex_geniculata

Ilex_hookeri

Ilex_montana

Ilex_micrococca

Ilex_canariensis

Ilex_repanda

Ilex_brevicuspis

Ilex_rugosa

Ilex_mitis

Ilex_integra1

Ilex_tolucana

Ilex_theizans1

Ilex_coriacea

Ilex_arnhemensis

Ilex_serrata

Ilex_discolor

Ilex_sebertii

Ilex_zhejiangensis

Ilex_decidua_var_curtissii

Ilex_pedunculosa

Ilex_warburgii

Ilex_liebmannii

Ilex_corallina

Ilex_georgei

Ilex_cornuta2

Ilex_purpurea1

Ilex_pubescensIlex_longipes

Ilex_sp_Craven

Ilex_colchica

Ilex_nothofagifolia

Ilex_tsoi

Ilex_laurina

Ilex_anomala

Ilex_cyrtura

Ilex_aquifolium

Ilex_pseudobuxus

Ilex_glabra

Ilex_integerrima

Ilex_perado

Ilex_argentina

Ilex_amelanchier

Ilex_cornuta1

Ilex_fragilis

Ilex_guianensis

Ilex_cassine

Ilex_vomitoria

Ilex_fargesii1

Ilex_nervulosa

Ilex_quercetorum

Ilex_paraguariensisIlex_hylonoma

Ilex_wilsonii

Ilex_amara

Ilex_goshiensis

Ilex_maximowicziana

Ilex_havilandii

Ilex_cymosaIlex_kinabaluensis

Ilex_mucronata

Ilex_yunnanensis1

Ilex_decidua

Ilex_kingianaIlex_ficoidea

Ilex_matanoana

Ilex_sugerokii

Ilex_liukiuensis

Ilex_brasiliensis

Ilex_triflora

Ilex_spinigera

Ilex_nitida

Ilex_x_makinoi

Ilex_rotunda1

Ilex_laevigata

Ilex_mutchagara

Ilex_macrocarpa

Ilex_dimorphophylla

Ilex_microdonta

Ilex_crenata

Ilex_macropoda

Ilex_pernyi

Ilex_aculeolata

Ilex_mertensii

Ilex_x_kiusiana

Ilex_rubra

Ilex_purpurea2

Ilex_asperula

Ilex_cumulicola

Ilex_fargesii2

Ilex_buergeri

Ilex_revoluta

Ilex_x_attenuata

3,17

6,33

13,32

4,681,22

14

18,16

18,98

15,38

17,08

24,11

5,3

22,97

2,45

30,54

6,84

12

3,6

5,74

5,79

2,82

7,37

10,8

3,57

27,02

3,68

15,13

17,55

3,24

15,53

4,29

8,79

5,56

9,79

4,77

5,09

36,19

7,37

24,3

3,51

3,39

1,44

8,62

10,72

6,17

7,71

9,45

45,88

8,91

2,52

37,86

13,93

13,15

14,3524,18

3,85

11,22

21,23

9,2

26,87

21,35

8,29

14,58

5,97

5,73

8,97

10,48

3,72

15,47

8,61

12,25

5,17

5,68

35,56

13,33

36,34

10,98

16,85

22,97

26,52

5,69

18,64

6,83

5,21

1,69

32,54

10,38

3,44

4,68

17,52

3,91

3,6

2,22

33,57

4,26

6,91

4,66

7,34

24

10,27

1,37

13,12

16,24

5,19

73,98

8,67

9,58

13,06

4,05

6,21

17,72

8,23

1,82

18,81

7,31

13,37

Ilex (plastid data)

Ilex_spicata

Ilex_rotunda2

Ilex_liukiuensis

Ilex_macrocarpa

Ilex_fragilis

Ilex_chamaedryfolia

Ilex_cyrtura

Helwingia_japonica

Ilex_paraguariensis

Ilex_dumosa

Ilex_purpurea2

Ilex_geniculata

Ilex_aculeolata

Ilex_zhejiangensis

Ilex_x_attenuataIlex_cassine

Ilex_micrococca

Ilex_integra2

Ilex_kingiana

Ilex_argentina

Ilex_spinigera

Ilex_ficoidea

Ilex_pseudobuxus

Ilex_latifolia

Ilex_cornuta1

Ilex_mutchagara

Ilex_tolucana

Ilex_laurina

Ilex_cumulicola

Ilex_discolor

Ilex_wallichii2

Ilex_aquifolium

Ilex_integerrima

Ilex_pernyi

Ilex_crenata

Ilex_fargesii1

Ilex_theizans2

Ilex_sugerokii

Ilex_havilandii

Ilex_wilsonii

Ilex_nervulosa

Ilex_cornuta2

Ilex_teratopis

Ilex_theizans1

Ilex_buergeri

Ilex_matanoana

Ilex_maingayi

Ilex_serrata

Ilex_dugesii

Ilex_x_kiusiana

Ilex_yunnanensis1

Ilex_amelanchier

Ilex_cymosa

Ilex_canariensis

Ilex_perado

Ilex_zygophylla

Ilex_asperula

Ilex_vomitoria

Ilex_sebertii

Ilex_nothofagifolia

Ilex_arnhemensis

Ilex_maximowicziana

Ilex_leucoclada

Ilex_tsoi

Ilex_brasiliensis

Ilex_triflora

Ilex_collinaIlex_decidua

Ilex_anomala

Ilex_pubescens

Ilex_x_makinoi

Ilex_pedunculosa

Ilex_ciliospinosa

Ilex_integra1

Ilex_laevigata

Ilex_amara

Ilex_goshiensis

Ilex_verticillata

Ilex_georgei

Ilex_wallichii1

Ilex_brevicuspis

Ilex_yunnanensis2Ilex_mitis

Ilex_longipes

Ilex_guianensis

Ilex_warburgii

Ilex_rugosa

Ilex_hookeri

Ilex_hylonoma

Ilex_decidua_var_curtissii

Ilex_mertensii

Ilex_oppositifolia

Ilex_opaca

Ilex_fargesii2

Ilex_rotunda1

Ilex_kusanoi

Ilex_purpurea1

Ilex_sp_Craven

Ilex_macropoda

Ilex_montana

Ilex_colchica

Ilex_quercetorum

Ilex_corallina

Ilex_cissoidea

Ilex_rubra

Ilex_glabra

Ilex_dimorphophylla

Ilex_liebmannii

Ilex_revoluta

Ilex_bioritsensis

Ilex_mucronata

Ilex_nitidaIlex_repanda

Ilex_microdonta

Ilex_shennongjiaensis

Ilex_kinabaluensis

Ilex_coriacea

36,43

4,63

5,57

5,42

21,13

6,25

10,39

6,83

1,72

27,19

7,2

12,77

12,37

31,84

23,13

5,13

7,745,14

1,43

3,79

4,36

17,08

1,58

5,52

7,669,71

6,61

20,56

8,21

7,915,61

5,17

9,6

1,26

2,08

25,55

1,01

19,07

2,9

8,04

0,65

2,18

8,4

6,04

2,58

8,08

6,83

7,72

3,14

18,95

14,78

4,68

3,78

28,57

1,51

2,82

2,41

10,1

14,59

13,375,72

11,44

20,71

8,79

13,28

17,35

2,92

10,85

10,61

15,42

9,49

7,92

5,13

17,5

5,05

4,35

2,52

73,8

1,94

21,25

11,4

6,02

15,9

14,64

1,19

17,02

19,57

24,12

11,8

24,36

8,16

10,37

2,18

4,54

16,06

10,6

0,5

7,12

7,57

3,2

2,69

3,01

29,2

22,58

10,23

17,55

24,63

3,76

1,34

9,63

5,64

22,41

14,77

8,85

2,94

3,87

Ilex (nuclear data)

JX145678_M_evidens_Gabon

PK373_M_cornuta_500to750mASL_PHS

Myrmarachne_sp_MRB113

Myrmarachne_sp_F_FSC_2006

PK210_M_hanoii_1000to1250mASL_MA

PK036_M_christae_2500to2750mASL_L2

PK183_M_sp_3_1750to2000mASL_GA

PK051_M_sp_4_500to750mASL_PHS

AY297412_M_assimilis_Philippines

Myrmarachne_japonica

Myrmarachne_sp_B_FSC_2006

Myrmarachne_sp_D_FSC_2006

PK068_M_malayana_500to750mASL_PHS

EU815599_Ligonipes_sp_Australia_OUTGROUP

Myrmarachne_plataleoides

PK256_M_nov_sp_2_1000to1250mASL_MA

Myrmarachne_sp_WPM_2008

JX145679_M_foenisex_Gabon

JX145676_M_cf_gedongensis_Malaysia

PK050_M_nov_sp_1_500to750mASL_PHS

Myrmarachne_sp_A_FSC_2006

PK003_M_mariaelenae_1500to1750mASL_KHQ

PK073_M_sp_5_500to750mASL_PHS

0,01

0,25

0,33

0,49

0,32

1,05

0,83

0,33

1,32

0,12

0,63

0,58

0,23

0,88

0,88

0,38

0,76

0,01

0,51

0,1

0,17

0,64

Myrmarachne

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

Extended Data Figure 5 | Chronograms for Athyrium, Begonia, Coeliccia,Dacrycarpus, Epirixanthes, Dendrochilum, Dicranoloma, Impatiens,Hygrobates, Hedyotis, Ilex and Myrmarachne. See Supplementary Table 5

for more details. Node colours represent posterior probabilities. Endemicspecies printed in red.

LETTER RESEARCH


Euphrasia_micrantha

Euphrasia_borneensis

Euphrasia_tarokoana

Euphrasia_maximowicziiEuphrasia_kerneriEuphrasia_hirtella

Euphrasia_tricuspidata

Euphrasia_fennica

Euphrasia_picta

Euphrasia_insignis

Euphrasia_regelii

Euphrasia_tauricaEuphrasia_chrysantha

Euphrasia_cockayniana

Euphrasia_oakesii

Euphrasia_confusa

Euphrasia_revoluta

Euphrasia_stricta

Euphrasia_alpina

Euphrasia_randii

Euphrasia_hookeriBartsia

Euphrasia_glabrescens

Euphrasia_nankotaizanensis

Euphrasia_disperma

Euphrasia_scabra

Euphrasia_cuneata

Euphrasia_minima

Euphrasia_antarctica

Euphrasia_phragmostoma

Euphrasia_vigursii

Euphrasia_subarctica

Euphrasia_collina

Euphrasia_petiolaris

Euphrasia_inopinata

Euphrasia_dinarica

Euphrasia_kashmiriana

Euphrasia_parviflora

Euphrasia_mirabilis

Euphrasia_nemorosa

Euphrasia_philippinensis

Euphrasia_marchesettii

Euphrasia_alsa

Euphrasia_foulaensis

Euphrasia_christii

Euphrasia_wettsteinii

Euphrasia_durietziana

Euphrasia_rostkoviana

Euphrasia_formosissima

Euphrasia_callosa

Euphrasia_tetraquetra

Euphrasia_fedtschenkoana

Euphrasia_spectabilis

Euphrasia_officinalis

Euphrasia_salisburgensis

Euphrasia_arctica

Euphrasia_disjuncta

Euphrasia_pectinata

Euphrasia_cisalpina

Euphrasia_grandiflora

Euphrasia_transmorrisonensis

1,5

5,33

0,72

6,54

1,07

0,18

4,03

1,31

3,89

0,29

0,62

0,44

7,38

0,57

1,3

0,86

3,29

2,72

1,31

7,61

5,38

0,19

1,21

1,4

2,71

2,15

0,89

0,66

4,88

0,42

1,75

2,87

2,62

1,87

8,22

6,3

5,96

1,62

1,23

26,5

6,41

0,36

3,55

0,37

4,43

8,49

2,36

0,42

0,97

3,12

1,03

2,25

0,21

3,34

0,95

0,36

6,73

0,46

4,45

1,72

Euphrasia

MJ81_K_baluensis_1750to2000mASL_GA

Elachistocleis_ovalis

Glyphoglossus_molossus

Dermatonotus_muelleri

Anodonthyla_boulengerii

Phrynomantis_bifasciatus

Platypelis_grandis

Scaphiophryne_calcarata

Otophryne_pyburni

Calluella_guttulata

Ramanella_cf_obscura_MNHN_2000_628

Rhombophryne_alluaudi

Cophixalus_sp_SJR3271

Dyscophus_antongilii

AB611939_Metaphrynella_sundana

Breviceps_mossambicus

MJ65_K_sp_2_500to750mASL_IN

Gastrophryne_carolinensis

Asterophrys_turpicola

MJ64_K_sp_1_500to750mASL_IN

AB611931_Metaphrynella_pollicaris

Callulina_kreffti

Hypopachus_variolosus

Stumpffia_psologlossa

Microhyla_heymonsi

Dyscophus_insularis

Paradoxophyla_palmata

Plethodontohyla_brevipes

Chiasmocleis_shudikarensis

Breviceps_fuscus

AB611913_K_interlineatus

Anodonthyla_montana

Microhyla_pulchra

Syncope_hudsoni

Stumpffia_pygmaea

Kaloula_pulchra

Microhyla_butleri

AB611921_Kalophrynus_pleurostigma

Rhombophryne_testudo

AB611973_Phrynella_pulchra

Phrynomantis_annectens

Micryletta_inornata

24,02

29,94

16,39

29,63

10,68

18,93

24,54

18,33

28,48

60,18

48,85

37,99

42,56

20,77

49,42

27,07

41,53

47,32

21,61

35,78

29,24

19,96

16,95

14,47

46,69

32,43

34,74

58,5130,17

68,65

29,56

53,15

55,9

34,89

25,88

27,46

47,98

54,22

14,32

40,37

38,18

Nepenthes_fusca

Nepenthes_northiana

Nepenthes_reinwardtiana

Dioncophyllum_thollonii

Nepenthes_singalana

Nepenthes_macfarlanei

Nepenthes_villosa

Nepenthes_rajah

Nepenthes_sanguinea

Nepenthes_mirabilis

Nepenthes_specatabilis

Nepenthes_bongso

Nepenthes_bokorensis

Nepenthes_hurrelliana

Nepenthes_lowii

Nepenthes_truncata

Nepenthes_kamotiana

Nepenthes_ventricosa

Nepenthes_x_kinabaluensis

Ancistrocladus_tectorius

Nepenthes_ramispina

Nepenthes_campanulata

Nepenthes_vieillardii

Nepenthes_villosa3

Nepenthes_gymnamphora

Nepenthes_edwardsiana2

Nepenthes_boschiana

Nepenthes_vogelii

Nepenthes_gracilis2

Nepenthes_peltata

Nepenthes_villosa2

Nepenthes_chaniana

Nepenthes_hemsleyana

Nepenthes_rafflesiana

Nepenthes_glandulifera

Nepenthes_kongkandana


Nepenthes_alba

Drosophyllum_lusitanicum

Nepenthes_alata

Nepenthes_albomarginata

Nepenthes_rajah2

Nepenthes_andamana

Triphyophyllum_peltatum

Nepenthes_insignis

Nepenthes_maxima

Nepenthes_ampullaria

Nepenthes_benstonei

Nepenthes_tentaculata

Nepenthes_gracillima2,4

19,97 9,05

1,36

1,34

18,29

11,5

1,29

5,15

7,56

3,34

2,73

27,06

3,22

1,844,23

3,22

4,23

8,84

1,24

3,79

14,03

3,29

13,52

1,63

2,28

14,6

53,45

10,84

36,21

1,48

5,47

5,93

2,52

8,14

2,17

5,58

27,11

5,71

1,27

15,89

22,16

32,1

3,07

2,9317,91

1,55

24,97

6,36

Nepenthes_muluensis

Nepenthes_ampullaria

Nepenthes_talangensis

Nepenthes_clipeata

Nepenthes_aristolochioides

Nepenthes_petiolata

Nepenthes_boschiana

Drosophyllum_lusitanicum

Nepenthes_truncata

Nepenthes_sibuyanensis

Nepenthes_diatas

Nepenthes_spectabilis

Nepenthes_eymae

Nepenthes_villosa_2


Nepenthes_singalana

Nepenthes_mira

Nepenthes_mikei

Nepenthes_merrilliana

Nepenthes_dubia


Nepenthes_villosa2

Nepenthes_sanguinea


Nepenthes_inermis

Nepenthes_rafflesiana

Nepenthes_densiflora

Nepenthes_hamata

Nepenthes_mapuluensis

Ancistrocladus_abbreviatus

Nepenthes_veitchii

Nepenthes_alata

Nepenthes_sumatrana

Nepenthes_hirsuta

Nepenthes_burkei

Nepenthes_maxima

Nepenthes_rajah

Nepenthes_gracillima

Triphyophyllum_peltatum

Nepenthes_murudensis

Nepenthes_rajah2

Nepenthes_tobaica

Habropetalum_dawei

Nepenthes_villosa3

Nepenthes_lowii

Nepenthes_thorelii

Nepenthes_stenophylla

Nepenthes_ephippiata

Nepenthes_adnataNepenthes_longifolia


Nepenthes_insignis

Nepenthes_glabrata

Nepenthes_bicalcarata

Nepenthes_fusca

Dioncophyllum_thollonii

Nepenthes_burbidgeae

Nepenthes_macrovulgaris


Nepenthes_mirabilis


Nepenthes_bongso

Nepenthes_pilosa

Nepenthes_spathulata

Nepenthes_ramispina

Nepenthes_ovata

Nepenthes_gracilis2

Nepenthes_lavicola


Nepenthes_rhombicaulis

Nepenthes_eustachya

Nepenthes_bellii

Nepenthes_faizaliana

7,87

8,55

6,71

5,01

3,64

11,37

4,13

53,03

1,75

3,53

0,722,14

5,6

2,69

7,58

30,88

6,46

0,47

2,96

5,27

7,36

4,37

2,45

0,66

1,7

5,06

1,45

0,96

2,51

3,5

2,88

2,43

1,62

3,21

4,03

3,01

0,68

4,46

8,57

4,81

1,5

1,17

3,89

20,42

2,65

6,24

4,44

2,13

10,027,1

1,18

14,34

1,59

13,87

2,79

2,51

3,59

0,63

7,38

6,66

6,11

1,02

5,09

12,36

1,95

4,52

3,62

24,32

15,85

38,11

9,91

1,05

Kalophrynus

Nepenthes (ITS)

Nepenthes (matK)

Paphiopedilum_niveum

Paphiopedilum_philippinense

Paphiopedilum_micranthum

Paphiopedilum_sukhakulii

Paphiopedilum_glaucophyllum

Paphiopedilum_victoria-regina

Paphiopedilum_barbatum

Paphiopedilum_ciliolare

Paphiopedilum_wardii

Paphiopedilum_hennisianum

Paphiopedilum_glanduliferum

Paphiopedilum_druryi

Paphiopedilum_mastersianum

Paphiopedilum_kolopakingii

Paphiopedilum_liemianum

Paphiopedilum_primulinum

Paphiopedilum_charlesworthii

Paphiopedilum_gratrixianum

Paphiopedilum_besseae

Paphiopedilum_sangii

Paphiopedilum_stonei

Paphiopedilum_appletonianum

Paphiopedilum_lowii

Paphiopedilum_parishii

Paphiopedilum_randsii

Paphiopedilum_dianthum

Paphiopedilum_callosum

Phragmipedium_schlimii

Paphiopedilum_javanicum_var_virens

Paphiopedilum_spicerianum

Paphiopedilum_wilhelminiae

Paphiopedilum_villosum_var_boxallii

Paphiopedilum_lawrenceanum

Paphiopedilum_insigne

Paphiopedilum_delenatii

Paphiopedilum_haynaldianum

Paphiopedilum_concolor

Paphiopedilum_tonsum

Paphiopedilum_violascens

Paphiopedilum_malipoense

Paphiopedilum_superbiens_var_curtisii

Paphiopedilum_rothschildianum

Paphiopedilum_fowliei

Paphiopedilum_exul

Phragmipedium_longifolium

Paphiopedilum_adductum

5,73

1,16

1,24

0,61

2,63

3,57

4,16

1,32

2,13

0,14

0,46

3,78

0,79

1,8

2,46

9,14

2,04

1,06

4,79

12,77

0,93

9,16

5

0,64

22,6

2,76

6,65

1,48

2,01

0,533,67

6,07

1,8

1,94

3,56

1,19

2,75

0,32

7,22

1,56 0,85

5,35

0,71

2,77

0,07

Paphiopedilum

Anigozanthos_flavidus

Elaeis_oleifera

Listera

Paphiopedilum_haynaldianumPaphiopedilum_delenatii

Monadenia

Eriochilus

Empodium

Platythelys

Earina_autumnalis

Lanaria

Epipactis

Microtis

Masdevallia

Maxillaria

Pleione

Astelia

Wullschlaegelia

Diuris

Palmorchis

Phragmipedium

Gongora

Musella_lasiocarpa

Ponthieva

Mormodes

Pogonia

Bifrenaria

Neuwiedia

Dendrobium_crystallinum

Cymbidium

Cleistes

Ludisia

Dendrobium_officinale

Calanthe

Apostasia

Hypoxis

Eria

Codonorchis

Nervilia

Disa

Galeandra

Nypa_fruticans

Zygopetalum

Agrostophyllum_majus

Platanthera

Vitekorchis

Phalaenopsis

Dendrobium_kingianum

Altenstenia

Dossinia

Habenaria

Goodyera

Cattleya

Sobralia

Lycaste

Sarcoglottis

Cypripedium

Chiloglottis

Orchis

Vanilla

Chloraea

Exalaria

Stanhopea

Megastylis

Disperis

Pterostylis

Earina_valida

Pachyplectron

Rhodohypoxis

Spiranthes

Selenipedium

39,8233,16

50,11

61,82

77,19

34,07

40,81

18,43

22,56

106,78

40,79

13,75

8,73

32,6

12,1

33,18

68,88

16,72

49,39

38,74

83,97

39,36

31,92

19,41

26,22

32,21

14,01

60,75

26,52

24,5

33,24

11,76

16,11

18,9623,48

29,23

25,59

16,55

34,3837,1

38,32

13,71

48,92

59,36

24,327,67

14,0722,84

25,45

7,68

16,47

23,38

45,63

102,14

8,57

32,9

38,57

53,91

37,24

32,58

42,76

34,93

44,13

28,07

13,82

35,98

71,34

43,03

57,72

66,83

Orchidaceae – Paphiopedilum

Zospeum_pretneri

Zospeum_spelaeum

Zospeum_alpestre

SP13510_Philalanka_spec2_1000to1250mASL_MA

SP13507_Philalanka_spec1_250to500mASL_IN

SP13492_Philalanka_spec3_500to750mASL_PHS

HQ171529_Carychium_mexicanum_USA

Zospeum_frauenfeldi

Zospeum_subobesum

SP13519_Philalanka_thienemanni_1000to1250mASL_M

Carychium_ibazoricum

HQ171582_Zospeum_exiguum_Slovenia

Carychium_minimum

Zospeum_isselianum

Carychium_exile

Carychium_clappi

Carychium_exiguum

SP13428_Ibycus_rachelae_1000to1250mASL_MA

JX988066_Subulina_octona_unknown

Carychium_floridanum

15,37

0,63

21,4

2,21

8,12

4,59

9,01

0,64

1,04

19,72

6,66

1,53

2,67

1,4

3,56

12,01

3,08

5,73

1,26

RMNH_INS_555592_Ptomaphaginus_similipes_250to500mASL_Sugud

RMNH_INS_555607_Ptomaphaginus_latescens_1500to1750mASL_KHQ

RMNH_INS_555598_Ptomaphaginus_bryantioides_250to500mASL_Kiansom

RMNH_INS_555632_Ptomaphaginus_nr_fagei_1750to2000mASL_GA

Carabus_lewisianus

Dytiscus_alaskanus

Carabus_japonicus

RMNH_INS_555618_Ptomaphaginus_n_sp_bryanti_complex_2750to3000mASL_

RMNH_INS_555603_Ptomaphaginus_kinabaluensis_1500to1750mASL_KHQ

Dytiscus_dauricus

RMNH_INS_555623_Ptomaphaginus_ater_3000to3250mASL_L2

RMNH_INS_555611_Ptomaphaginus_cf_latimanus_750to1000mASL_SA

RMNH_INS_549243_Prionochaeta_harmandi_Japan

Carabus_kimurai

Carabus_albrechti

Carabus_yamato

0,88

0,66

0,06

0,1

0,42

1,6

2,14

1,55 0,15

0,7

0,06

0,12

0,98

1,05

1,99

Philalanka

Ptomaphaginus

Madhuca_pachyphylla_EDNA09_01386

Payena_leerii_EDNA09_02317

Diploknema_oligomera

Mimusops_obovata

Manilkara_obovata

Palaquium_sericeum_EDNA09_02304

Madhuca_utilis

Labramia_mayottensis

Palaquium_dasyphyllum_EDNA07_01936

Payena_obscura_EDNA09_02313

Sideroxylon_angustum

Sideroxylon_persimile

Vitellariopsis_cuneata

Madhuca_malaccensis_EDNA09_00958

Sideroxylon_wightianum

Sideroxylon_occidentale

Sideroxylon_picardae

Palaquium_maingayi_EDNA09_00957

Madhuca_elmeri_EDNA09_02183

Omphalocarpum_pachysteloides

Madhuca_sarawakensis_EDNA09_01136

Pouteria_maclayana_EDNA09_00990

Madhuca_microphylla

e4003829348_P_kinabaluensis_844mASL_OL

Sideroxylon_marginatum

Diploknema_siamensis_EDNA09_2330

Sideroxylon_inerme

Sideroxylon_oxyacanthum

Manilkara_kauki

Sideroxylon_americanum

Palaquium_microphyllum

Eberhardtia_aurata

Payena_lucida_EDNA09_02306

e4003829276_P_gigas_830mASL_OL

Englerophytum_natalense

Palaquium_microphyllum_EDNA09_02312

Payena_acuminata

Neolemonniera_clitandrifolia

Sideroxylon_mascatense

Sideroxylon_foetidissimumSideroxylon_capiri

Mimusops_caffra

Palaquium_pseudorostratum_EDNA09_01131

Mimusops_zeyheri

Palaquium_sumatranum_EDNA09_00955

Faucherea_parvifolia

Palaquium_oxleyanum_EDNA09_00719

Madhuca_laurifolia_EDNA09_02184

Sideroxylon_tepicense

Madhuca_curtisii_EDNA09_02307

Palaquium_amboinense

Madhuca_oblongifolia_EDNA09_01379

Burckella_polymera_EDNA09_01388

Labourdonnaisia_revoluta

Isonandra_perakensis_EDNA09_00998

Pouteria_sp_EDNA09_02322

Capurodendron_androyense

Palaquium_gutta_EDNA09_00717

Palaquium_sp_EDNA09_02320

Madhuca_korthalsii_EDNA09_01380

Sideroxylon_marmulano

Letestua_durissima

Palaquium_eriocalyx_EDNA09_00993

Sideroxylon_obovatum

Inhambanella_henriquesii

Palaquium_sericeum_EDNA09_01452

Sideroxylon_horridum

Autranella_congolensis

Palaquium_sp_EDNA09_00989

Labramia_costata

Palaquium_beccarianum_EDNA09_00952

Northia_seychellana

Palaquium_rigidum_EDNA09_01129

Palaquium_clarkeanum_EDNA09_00716

Madhuca_hainanensis

Palaquium_ridleyi_EDNA09_01132

Madhuca_lancifolia_EDNA09_01382

Madhuca_palida_EDNA09_01139

Palaquium_formosanum_EDNA09_00722

Xantolis_siamensis

Tieghemella_heckelii

Sideroxylon_floribundum

Mimusops_elengi

Burckella_macropoda_EDNA09_00995

Sarcosperma_laurinum

Madhuca_sericea_EDNA09_01378

Mimusops_comorensis

Palaquium_quercifolium_EDNA09_01451

Palaquium_lobbianum_EDNA09_02319

Vitellaria_paradoxa

Madhuca_longifolia_EDNA09_02311

Palaquium_rostratum_EDNA09_00956

Manilkara_discolor

Palaquium_cryptocarifolium_EDNA09_01144

Isonandra_sp

Sideroxylon_betsimisarakum

Palaquium_galactoxylon_EDNA09_00987

Lecomtedoxa_klaineana

Palaquium_brassii_EDNA09_01385

Vitellariopsis_marginata

Sideroxylon_reclinatum

Poutera_firma_EDNA09_01389

Sideroxylon_cubense

Madhuca_kunstleri_EDNA09_01138b

Argania_spinosa

Madhuca_palembanica

Manilkara_concolor

Palaquium_walsurifolium_EDNA09_01135

Madhuca_fulva_EDNA09_00985

Labourdonnaisia_calophylloides

Madhuca_prolixa_EDNA09_01381

Payena_maingayi_EDNA09_00988

Nesoluma_polynesiacum

Sideroxylon_lanuginosum

Manilkara_hexandra

Sideroxylon_saxorum

Madhuca_kuchingensis_EDNA09_01134

Sideroxylon_tenax

Diploknema_butyracea

Palaquium_calophyllum_EDNA09_00992

Isonandra_lanceolata_EDNA09_02321

e4003829322_P_microphylla_627mASL_IN

Madhuca_kingiana_EDNA09_00954

Sideroxylon_majus

Burckella_sp_EDNA09_02325

Madhuca_barbata_EDNA09_01143

Madhuca_erythrophylla_EDNA09_01137

Sideroxylon_repens

Madhuca_crassipes

Manilkara_zapota

Isonandra_compta_EDNA09_00984

Madhuca_motleyana

Palaquium_leiocarpum_EDNA09_01130

Sideroxylon_confertum

Palaquium_impressionervium_EDNA09_00720

Sideroxylon_salicifolium

Palaquium_rufolanigerum_EDNA09_01133

Madhuca_leucodermis

Aulandra_longifolia

Vitellariopsis_dispar

Payena_ferruginea_EDNA09_02314

Sideroxylon_lycioides

Palaquium_hexandrum_EDNA09_01142

Baillonella_toxisperma_EDNA09_01453

Palaquium_herveyi_EDNA09_01141

Pouteria_sp

Madhuca_sp_nov_EDNA09_01383

Palaquium_obovatum_EDNA09_00718

Sideroxylon_obtusifolium

Palaquium_xanthochymum_EDNA09_00721

7,39

9,27

6,05

3,88

3,93

24,31

38,68

20,87

39,04

3,56

7,17

3,35

36,4711,14

22,99

3,41

0,57

14,21

68,99

15,87

5,02

11,2

3,07

7,59

2,11

14,46

40,95

11,44

16,15

2,07

5,49

2,14

31,72

10,5648,14

1,23

1,09

4,67

4,2

25,31

7,8 4,75

7,47

35,28

34,27

1,56

0,99

35,84

11,51

0,42

31,84

0,67

0,83

0,59

5,35

4,62

29

1,69

3,072,49

10,45

16,2

0,31

3,3

6,63

5,14

10,51

3,64

6,39

1,25

6,74

5,13

22,5

20,06

7,72

6,88

8,66

25,76

4,02

9,15

17,35

10,39

18,39

4,23

8,63

0,84

31,42

47,79

40,08

1,04

7,1

6,19

17,92

6,28

17,9

1,33

34,47

20,72

7,56

4,56

31,37

53,33

10,03

22,12

4,89

22,54

5,2

20,82

5,59

1

2,49

21,03

75,68

25,54

8,5

15,52

3,49

8,82

9,49

29,74

1,72

23,61

5,15

11,14

0,42

14,91

2,89

22,48

19,5

0,85

3,58

2,58

6,04

2,06

0,56

15,98

0,97

18,08

4,44

4,12

0,67

3,8

26,01

18,62

0,75

3,06

4,64

0,28

18,53

Rhododendron_ericoides

Cassiope_tetragona

Rhododendron_ferrugineum

Rhododendron_buxifolium

Rhododendron_dilatatum

Rhododendron_hongkongense

Rhododendron_transiens

Rhododendron_anthosphaerum

Rhododendron_leptothrium

Rhododendron_reticulatum

Rhododendron_agastum

Rhododendron_forrestii

Rhododendron_weyrichii

Rhododendron_yakuinsulare

Rhododendron_sanctum

Rhododendron_edgeworthii

Rhododendron_farrerae

Rhododendron_albrechtii

Rhododendron_floccigerum

Rhododendron_cavaleriei

Rhododendron_stenophyllum

Rhododendron_amanoi

Rhododendron_fastigiatum

Rhododendron_beesianum

Rhododendron_tashiroi

Rhododendron_cyanocarpum

Rhododendron_auriculatum

Rhododendron_feddei

Rhododendron_irroratum

Rhododendron_pachypodum

Rhododendron_spinuliferum

Rhododendron_championiae

Rhododendron_mollicomum

Rhododendron_semibarbatum

Rhododendron_bagobonum

Rhododendron_williamsianum

Rhododendron_mackenzianum

Rhododendron_aureum

Rhododendron_wardii

Rhododendron_moulmainense

Rhododendron_bureavii

Rhododendron_mayebarae

Rhododendron_santapaui

Rhododendron_dimitrum

Rhododendron_crassifolium

Rhododendron_cuneifolium

Rhododendron_lukiangense

Rhododendron_tsusiophyllum

Rhododendron_lowii

Rhododendron_campylocarpum

Rhododendron_glischrum

Rhododendron_tutcherae

Rhododendron_macrosepalum

Rhododendron_scabrifolium

Rhododendron_maximum

Rhododendron_micranthum

Rhododendron_fortunei

Rhododendron_spiciferum

Rhododendron_selense

Rhododendron_ovatum

Rhododendron_formosanum

Rhododendron_luteumRhododendron_molle

Rhododendron_rubropunctatum

Rhododendron_praetervisum

Rhododendron_stamineum

Rhododendron_orbiculatum

Rhododendron_trichocladum

Rhododendron_occidentale

Rhododendron_stapfianum

Rhododendron_racemosum

Rhododendron_scabrum

Rhododendron_suaveolensRhododendron_canadense

Rhododendron_kiusianum

Rhododendron_rubiginosum

Rhododendron_acuminatum

Cassiope_fastigiata

Rhododendron_rugosum

Rhododendron_latoucheae

Rhododendron_fulvum

Rhododendron_mucronulatum

Rhododendron_borneense

Rhododendron_rex_subsp_fictolacteum

Rhododendron_indicum

Rhododendron_camtschaticum

Rhododendron_mariesii

Rhododendron_sinogrande

Rhododendron_neriiflorum

Rhododendron_hancockii

Rhododendron_alutaceum

Rhododendron_lagopus

Rhododendron_grande

Rhododendron_schlippenbachii

Rhododendron_wadanum

Rhododendron_aganniphum

Rhododendron_tosaense

Rhododendron_argyrophyllum

Rhododendron_aberconwayi

Rhododendron_boninense

Rhododendron_bachii

Rhododendron_duclouxii

Rhododendron_vernicosum

Rhododendron_eriocarpum

Rhododendron_mitriforme

Rhododendron_tomentosum_subsp_tomentosum

Rhododendron_kiyosumense

Rhododendron_subestipitatum

Rhododendron_dayaoshanense

Rhododendron_decorum

Rhododendron_westlandii

Rhododendron_albiflorum

Rhododendron_vialii

Rhododendron_delavayi

Rhododendron_exuberans

Rhododendron_redowskianum

Rhododendron_henryi

Rhododendron_nipponicum

Rhododendron_virgatum

Rhododendron_fallacinum

Rhododendron_serpyllifolium

Rhododendron_javanicum

Rhododendron_primuliflorum

Rhododendron_viscistylum

Rhododendron_ponticum

Rhododendron_kaempferi

4,83

7,56

12,91

14,19

4,39

3,22

6,31

2,68

2,81

9,18

2,34

6,37

6,29

4,07

22,46

4,21

10,57

6,33

5,63

0,59

19,61

4,29

0,7

6,42

10,65

7,25

2,92

1,81

5,63

5,4

6,25

2,57

2,52

17,79

9,72

7,08

14,96

3,19

16,62

7,69

2,11

1,98

14,6

10,16

16,62

5,92

5,2

9,13

7,37

1,21

7,35

2,49

2,19

11,74

4,12

13,64

19,66

3,42

5,72

3,4

3,94

14,44

10,32

3,81

4,15

5,16

5,02

4,15

22,13

8,57

12,4

2,61

1,89

5,91

7,39

6,7

17,08

3,28

4,61

3,88

25,14

2,59

4,83

2,87

0,61

3,98

4,24

12,05

4,09

42

3,02

5,1

8,78

2,89

4,01

2,86

8,74

4,04

1,09

6,31

5,01

3,63

7,16

18,93

36,25

7,82

9,5

11,69

1,89

2,83

5,13

11,72

4,23

1,98

3,67

4,69

2,62

10,91

9,58

27,9

3,48

2,43

8,11

9,26

2,71

Payena

Rhododendron

Weinmannia_samoensis

Pancheria_brunhesii

Weinmannia_parviflora

Weinmannia_sp_Bradford_655


Weinmannia_raiateensis


Weinmannia_vescoi

Brunellia

Weinmannia_marojejyensis

Karrabina_benthamiana

Vesselowskya

Pancheria_hirsuta

Weinmannia_bangii

Pancheria_phylliraeoides

Weinmannia_tinctoria

Weinmannia_fraxinea

Pullea_glabra

Weinmannia_auriculata

Weinmannia_bojeriana

Cephalotus_follicularis

Spiraeopsis_celebica

Weinmannia_stenostachya

Caldcluvia_paniculata

Weinmannia_exigua

Weinmannia_myrtifolia

Eucryphia_moorei

Weinmannia_silvicola

Cunonia_montana

Cunonia_capensis

Ackama_rosifolia

Weinmannia_racemosa

Pancheria_engleriana

Weinmannia_tomentosa

Weinmannia_trichosperma

Weinmannia_clemensiae

Cunonia_macrophylla

Weinmannia_sanguisugarum

Caldcluvia_paniculosa

Weinmannia_serrata

Cunonia_pulchella

Codia_discolor

Cunonia_balansae

Weinmannia_richii

Weinmannia_hooglandii

Weinmannia_humbertiana

Weinmannia_rutenbergii

Weinmannia_dichotoma

Callicoma_serratifolia

Weinmannia_arguta

Weinmannia_madagascariensis

Pancheria_reticulata

Cunonia_atrorubens

Weinmannia_vitiensis

Weinmannia_minutiflora

Weinmannia_elattantha

Eucryphia_cordifolia

87,63

18,73

5,27

32,41

3,14

25

8,79

2,02

9,23

6,79

3,94

1,786,09

63,37

34,24

20,18

28,67

39,86

2,45

48,62

4,57

22,88

24,56

3,91

22,7

2,05

16,94

10,81

10,02

9,41

59,78

27,91

3,45

6,66

6,78

5,89

15,97

42,57

2,62

10,6

14,25

6,34

11,8

1,64

12,63

50,63

5,1

63,68

2

2,02

9,55

16,98

2,97

2,06

0,96

0,94

Weinmannia

Arcteranthis_cooleyae

Ranunculus_cardiophyllus

Ranunculus_natans

Ranunculus_multifidus

Ranunculus_marschlinsii

Ranunculus_adoneus

Ranunculus_hydrophilus

Ranunculus_tembensis

Ranunculus_macounii

Ranunculus_gregarius

Ranunculus_sericeus

Ranunculus_chinensis

Ranunculus_flagelliformis

Ranunculus_saruwagedicus

Ranunculus_rufosepalus

Ranunculus_sceleratus

Coptis_lutescens

Ranunculus_polyanthemos

Ranunculus_basilobatus

Ranunculus_millefolius

Beckwithia_andersonii

Ranunculus_kotschyi

Kumlienia_hystricula

Ranunculus_seguieri

Ranunculus_bonariensis

Ranunculus_pinnatus

Ranunculus_heterorrhizus

Anemone_quinquefolia

Halerpestes_uniflora

Ranunculus_meyeri

Coptis_deltoidea

Ranunculus_hispidus

Ceratocephala_orthoceras

Ranunculus_regelianus

Ranunculus_japonicus

Ranunculus_rumelicus

Ranunculus_venetus

Ranunculus_pseudomillefoliatus

Ranunculus_cf_pseudopygmaeus_KE2011

Ranunculus_sartorianus

Ranunculus_peduncularis

Ranunculus_brutius

Ranunculus_pseudohirculus

Ranunculus_millefoliatus

Ranunculus_longicaulis

Ranunculus_cortusifolius

Ranunculus_gouanii

Coptidium_lapponicum

Ranunculus_kuepferi_subsp_orientalis

Isopyrum_thalictroides

Ranunculus_peltatus_subsp_peltatus

Ranunculus_hybridus

Ranunculus_garganicus

Ranunculus_paludosus

Ranunculus_psilostachys

Ranunculus_crenatus

Ranunculus_argyreus

Ranunculus_amblyolobus

Ranunculus_papulentus

Ranunculus_pulchellus

Ranunculus_ollissiponensis

Ranunculus_traunfellneri

Ranunculus_bulbosus_subsp_bulbosus

Coptis_quinquefolia

Halerpestes_cymbalaria

Ranunculus_gramineus

Ranunculus_glabriusculus

Ranunculus_macropodoides

Ranunculus_alismifolius

Ranunculus_damascenus

Ranunculus_lowii

Ranunculus_acris

Ranunculus_repens

Ranunculus_reptans

Ranunculus_arvensis

Ranunculus_sprunerianus

Ranunculus_leptorrhynchus

Cyrtorhyncha_ranunculina

Ranunculus_spicatus

Ranunculus_ficariifolius

Ranunculus_caprarum

Ranunculus_uncinatus

Hamadryas_delfinii

Ranunculus_caucasicus

Ranunculus_linearilobus

Ranunculus_pinardi

Ranunculus_asiaticusRanunculus_oxyspermus

Ranunculus_afghanicus

Ranunculus_brevifolius_subsp_brevifolius

Ranunculus_amerophyllus

Ranunculus_cassius

Ranunculus_carpaticus

Ranunculus_bilobus

Ranunculus_taisanensis

Ranunculus_sphaerospermus

Krapfia_clypeata

Ranunculus_septentrionalis

Peltocalathos_baurii

Ranunculus_bullatus

Ranunculus_trilobus

Coptis_chinensis

Ranunculus_cantoniensis

Ranunculus_nipponicus

Ranunculus_petiolaris

Ranunculus_aduncus

Callianthemoides_semiverticillatus

Ranunculus_hyperboreus

Ranunculus_silerifolius

Ranunculus_amplexicaulis

Coptis_teeta

Coptis_trifolia

Ranunculus_cornutus

Ranunculus_sojakii

Ranunculus_brotherusii

Ranunculus_mauiensis

Ranunculus_membranaceus

Ranunculus_carinthiacus

Ranunculus_gmelinii

Ranunculus_hirtellus

Ranunculus_diffusus

Ranunculus_pyrenaeus

Ranunculus_cheirophyllus

Ranunculus_serbicus

Ranunculus_grandiflorus

Ranunculus_micranthus

Ranunculus_lateriflorus

Ranunculus_pygmaeus

Ranunculus_volkensii

Ranunculus_neapolitanus

Ranunculus_montanus

Ranunculus_radicans

Ranunculus_nivalis

Trautvetteria_grandis

Ranunculus_pseudomontanus

Ranunculus_magellensis

Ranunculus_platanifolius

Ranunculus_cassubicifolius

Ranunculus_hawaiensis

Ranunculus_parviflorus

Coptis_omeiensis

Ranunculus_laetus

Ranunculus_alpestris

Ranunculus_dissectus_subsp_napellifolius

Oxygraphis_polypetala

Ranunculus_aucheri

Myosurus_minimus

Ranunculus_sardous

Ranunculus_glacialis

Ranunculus_stagnalis

Ranunculus_gracilis

Ranunculus_pseudolowii

Ranunculus_makaluensis

Ranunculus_brachylobus

Ranunculus_parnassiifolius_subsp_parnassiifolius

Ranunculus_constantinopolitanus

Ranunculus_penicillatus_subsp_pseudofluitans

Ranunculus_brassii

Ranunculus_occidentalis

Ficaria_fascicularis

Ranunculus_pedatifidus

Ranunculus_maclovianus

Ranunculus_elbrusensis

Ranunculus_tenuirostris

Ranunculus_ophioglossifolius

Ranunculus_chius

Ranunculus_thora

Ranunculus_illyricus

Ranunculus_punctatus

Ficaria_verna

Ranunculus_lingua

Ranunculus_gelidus

Ranunculus_orthorhynchus

Ranunculus_cacuminis

Ranunculus_acetosellifolius

Ranunculus_oreophytus

Ranunculus_muricatus

Ranunculus_cicutarius

Laccopetalum_giganteum

Ranunculus_apiifolius

Ranunculus_carpaticola

Ranunculus_trichophyllus

Coptidium_pallasii

Ranunculus_macrorrhynchus

Ranunculus_submarginatus

Ranunculus_papyrocarpus

Ranunculus_cappadocicus

Ranunculus_lyallii

Ranunculus_serpens_subsp_nemorosus

Ranunculus_fascicularis

Ranunculus_hierosolymitanus

Ranunculus_flammula

Coptis_japonica

Ranunculus_termei

Ranunculus_acriformis

Ranunculus_collinus

Ranunculus_pensylvanicus

Ranunculus_marginatus

Ranunculus_aconitifolius

Ranunculus_buhsei

Ranunculus_granatensis

Ranunculus_aquatilis

Ranunculus_villarsii

Ranunculus_nephelogenes

Ranunculus_baldshuanicus

Ranunculus_notabilis

Ranunculus_velutinus

Ranunculus_pollinensis

Ranunculus_strigillosus

Ranunculus_apenninus

Ranunculus_breyninus

Ceratocephala_falcata

Ranunculus_raeae

Ranunculus_lanuginosus

Ranunculus_fuegianus

4,39

0,64

0,11

0,69

0,59

0,75

1,45

0,19

0,33

0,59

0,5

0,61

0,42

0,3

0,99

1,73

0,38

0,92

1,42

1,61

0,31

0,92

0,88

0,39

3,41

0,61

0,66

1,19

0,41

0,37

0,63

0,59

5,17

0,69

1,07

1,33

0,81

0,99

0,95

0,54

1,1

1,01

0,33

9,27

0,57

0,42

1,27

0,74

0,19

0,69

0,43

0,65

0,33

1,05

0,73

0,55

1,25

0,74

0,8

0,62

2,13

0,82

0,63

2,97

0,34

0,88

0,73

1,97

0,25

3,57

0,48

1,31

1,45

6,52

1,02

0,08

0,95

0,36

0,57

0,34

2,56

5,65

0,681,58

1,2

0,2

1,81

0,91

0,64

0,18

0,15

0,15

2,08

0,46

0,69

0,25

0,22

2,41

1,22

0,5

1,01

0,94

1,78

2,93

1,66

0,17

0,89

0,76

2,43

4,61

1,26

2,45

1,93

0,51

0,32

0,9

0,9

0,81

0,75

0,45

0,24

7,54

1,39

0,48

0,644,03

0,66

0,96

1,01

1,23

0,23

1,01

1,17

2,19

1,37

0,33

7,03

0,55

0,61

3,1

1,63

0,75

0,25

0,46

2,85

0,68

2,26

0,86

0,37

1,31

1,55

0,71

0,49

0,39

0,3

0,16

0,47

0,33

1,2

1,58

0,97

0,8

0,1

0,38

1,85

2,27

0,27

1,05

1,63

1,05

3,77

1,88

0,43

0,68

0,29

2,41

1,66

0,75

1,34

3,22

0,29

1,97

2,18

0,94

0,48

0,92

5,33

0,34

0,43

1,46

1,08

0,48

1,42

0,56

1,3

0,37

0,530,81

2,25

1,02

0,2

1,34

1,06

0,53

0,83

1,77

0,49

0,97

0,71

2,87

4,24

1,2

0,33

0,24

Luffa_echinata

Trichosanthes_intermedia

Trichosanthes_truncata3

Trichosanthes_elmeri

Trichosanthes_borneensis

Luffa_aegyptiaca

Trichosanthes_laceribractea1

Gymnopetalum_tubiflorum1

Trichosanthes_phonsenae2

Trichosanthes_wallichiana

Trichosanthes_phonsenae1

Gymnopetalum_chinense

Trichosanthes_rosthornii2

Trichosanthes_schlechteri

Trichosanthes_odontosperma3

Trichosanthes_pendula

Trichosanthes_obscura

Nothoalsomitra_suberosa

Gymnopetalum_scabrum1

Trichosanthes_pubera_var._rubriflos1

Trichosanthes_subvelutina1

Trichosanthes_pallida2



Trichosanthes_laeoica2

Echinocystis_lobata

Trichosanthes_lepiniana3


Trichosanthes_pilosa_var._roseipulpa

Trichosanthes_edulis

Trichosanthes_miyagii

Trichosanthes_pedata

Trichosanthes_homophylla

Trichosanthes_inthanonensis1

Trichosanthes_reticulinervis

Trichosanthes_montana_subsp._crassipes

Trichosanthes_cucumerina2

Trichosanthes_villosa5

Trichosanthes_pentaphylla1


Luffa_graveolens

Trichosanthes_multiloba2

Trichosanthes_pallida1

Cyclanthera_pedata


Trichosanthes_dioica

Trichosanthes_quinquefolia


Momordica_charantia

Trichosanthes_postarii2

Trichosanthes_papuana

Trichosanthes_dentifera

Gymnopetalum_orientale

Trichosanthes_pentaphylla2

Trichosanthes_pilosa2

Trichosanthes_tricuspidata_subsp._tricuspidata

Trichosanthes_kirilowii_var._japonica1

Trichosanthes_tricuspidata_subsp._javanica



Trichosanthes_sepilokensis

Linnaeosicyos_amara


Trichosanthes_celebica

Sicyos_angulatus


Trichosanthes_wawrae

Trichosanthes_postarii1

Trichosanthes_mucronata

Trichosanthes_laeoica1

Luffa_operculata

Trichosanthes_montana_subsp._montana

Trichosanthes_kerrii

Lagenaria_siceraria


Trichosanthes_quinquangulata2

Trichosanthes_pubera_var._fissisepala2

Ecballium_elaterium


Marah_macrocarpus



Trichosanthes_fissibracteata


Austrobryonia_micrantha

Trichosanthes_beccariana

Trichosanthes_pubera_var._fissisepala1

Trichosanthes_quinquangulata1

Trichosanthes_cucumerina1

Trichosanthes_smilacifolia

Trichosanthes_rosthornii1

Trichosanthes_multiloba1



Trichosanthes_auriculata


Trichosanthes_adhaerens




Trichosanthes_pubera_var._rubriflos2

Hodgsonia_heteroclita


Luffa_acutangula

Bryonia_dioica


Trichosanthes_inthanonensis2

Trichosanthes_kinabaluensis

Trichosanthes_baviensis

Trichosanthes_holtzei

Trichosanthes_hylonoma

Trichosanthes_bracteata

Trichosanthes_nervifolia

Trichosanthes_globosa

13,19

5,04

13,99

13,26

2,46

7,11

3,22

29

6,47

5,21

2,91

6,57

2,31

1,33

23,87

1,3

3,55

18,03

7,19

0,95

12,33

1,18

8,55

6,13

14,38

4,2

20,76

5,89

3,06

4,73

0,61

1,43

9,97

1,23

5,55

37,17

2,98

3,41

8,76

2,63

6,21

10,18

35,7

16,07

8,5

17,89

1,63

5,57

6,54

12,36

9,35

2,62

4,15

5,5

4,54

7,58

31,8

29,34

7,2

1,69

15,38

10,95

15,66

24,52

9,33

6,16

8,38

8,17

39,46

2,48

3,74

7,34

3,75

56,82

7,31

3,76

8,55

4,87

3,42

6,12

3,38

5,81

34,51

1,48

8,08

13,28

21,1

5,44

3,82

17,73

0,79

53,03

23,57

4,47

23,81

0,49

21,26

16,52

19,92

6,95

13,22

4,97

4,49

2,87

44,34

4,79

39,86

12,16

13,52

5,98

7,8950,28

3,39

Ranunculus

Trichosanthes Domanibdella_palmyrae

Phytobdella_meyeri

Chtonobdella_whitmani

Philaemon_sp_green_EB_2010

Hirudinaria_manillensis

Haemadipsa_hainana

Nesophilaemon_skottsbergii

Tritetrabdella_taiwana_C

Haemadipsa_sylvestris

Idiobdella_seychellensis

Neoterrabdella_australis

Domanibdella_EB_2010

Malagabdella_niarchosorum

Philaemon_pungens

Haemadipsa_cavatuses

Philaemon_mediorubra

Tritetrabdella_taiwana_A

Haemadipsa_picta

Tritetrabdella_scandens

Leiobdella_sp_EB_2010

Malagabdella_vagans

Haemadipsa_ornata

Haemadipsa_rjukjuana

Haemadipsa_zeylanica_montivindicis

SP13398_T_kinabalensis_kinabalensis_1750to2000mASL_G

Tritetrabdella_taiwana_B

Philaemon_cf_grandis_EB_2010

Philaemon_sp_brown_EB_2010

Haemadipsa_japonica_japonica

Philaemon_gloriosi

SP13383_T_kinabalensis_inobongensis_250to500mASL_IN

Malagabdella_fallax

Chtonobdella_bilineata

Planobdella_sp_EB_2010

4,47

3,22

2,01

16,67

0,87

3,96

3,37

3,49

5,35

3,01

3,33

4,38

2,68

2,76

7,23

6,39

2,89

3,48

2,89

8,64

4,41

3,26

4,75

3,33

0,73

3,26

3,81

2,686,25

4,83

2,71

4,21

5,76

Tritetrabdella

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

posterior1

0

Extended Data Figure 6 | Chronograms for Euphrasia, Kalophrynus,Nepenthes, Paphiopedilum, Philalanka, Ptomaphaginus, Payena,Trichosanthes, Rhododendron, Ranunculus, Tritetrabdella and

Weinmannia. See Supplementary Table 5 for more details. Node coloursrepresent posterior probabilities. Endemic species printed in red.

RESEARCH LETTER


Athyrium_arisanense

Athyrium_iseanum

Athyrium_x_hisatsuanum

Athyrium_strigillosum

Athyrium_clivicola

Athyrium_setuligerum

Athyrium_x_tokashikii

Athyrium_kuratae

Athyrium_subrigescens

Athyrium_oblitescens

Athyrium_otophorum

Athyrium_x_akiense

Athyrium_viridescentipe

Athyrium_eremicola

Athyrium_wardii

Athyrium_amoenum

Athyrium_deltoidofrons

Athyrium_neglectum

Athyrium_vidalii

Athyrium_nakanoi

Athyrium_tozanense

Athyrium_clemensiae

Athyrium_atkinsonii

Athyrium_kirisimaense

Athyrium_tashiroi

Athyrium_yokoscense

Athyrium_nikkoense

Athyrium_rupestre

Athyrium_brevifrons

Athyrium_melanolepis

Athyrium_filix_femina

35 30 25 20 15 10 5 0Millions of years ago

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

K

W

W

W

W

W

K

W

W

W

W

W

W

W

W

W

BioGeoBEARS DEC+J on Athyrium M0_unconstrainedancstates: global optim, 2 areas max. d=0; e=0; j=0.0344; LnL=−8.42

W

W

W

K

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

K

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

Begonia_cauliflora

Begonia_vaccinioides

Begonia_burbidgei

Begonia_imbricata

Begonia_cf_beryllae

Begonia_pendula

Begonia_amphioxus

2.5 2 1.5 1 0.5 0Millions of years ago

K

K

K

K

K

KB

KB

K

K

KB

KB

KB

KB

BioGeoBEARS DEC+J on Begonia M0_unconstrainedancstates: global optim, 2 areas max. d=0.8061; e=0; j=0; LnL=−3.54

K

K

K

K

K

K

K

K

K

K

K

K

Coeliccia_cyaneothorax

C_nemoricola

C_octogesima

C_species_1

C_species_2

C_albicauda

C_poungyi

C_cyanomelas

RMNH_INS_503926_C_didyma

Coeliccia_ryukyuensis

3 2.5 2 1.5 1 0.5 0Millions of years ago

W

W

K

K

B

KB

W

W

W

B

KB

BW

K

K

W

W

W

W

W

BioGeoBEARS DEC+J on Coeliccia M0_unconstrainedancstates: global optim, 3 areas max. d=0.0615; e=0; j=0.0588; LnL=−14.15

W

W

K

K

B

KB

W

W

W

W

K

K

B

B

B

W

W

W

Dacrycarpus_compactus

Dacrycarpus_expansus






Dacrycarpus_dacrydioides

60 50 40 30 20 10 0Millions of years ago

W

KBW

W

W

W

KBW

KBW

W

W

KBW

K

KBW

K

KBW

W

BioGeoBEARS DEC+J on Dacrycarpus M0_unconstrainedancstates: global optim, 3 areas max. d=0.0479; e=0; j=0; LnL=−11.60

W

W

W

W

W

K

K

W

W

W

W

W

KBW

KBW

Dendrochilum_alatum

Dendrochilum_corrugatum

Dendrochilum_kamborangense

Dendrochilum_stachyodes

Dendrochilum_trusmadiense

Dendrochilum_muluense

Dendrochilum_grandiflorum

Dendrochilum_alpinum

Dendrochilum_pterogyne

Dendrochilum_pseudoscriptum

Dendrochilum_scriptum

Dendrochilum_transversum

Dendrochilum_dewindtianum

Dendrochilum_tenompokense

Dendrochilum_haslamii

Dendrochilum_exasperatum

Dendrochilum_gibbsiae

Dendrochilum_joclemensii

Dendrochilum_cupulatum

5 4 3 2 1 0Millions of years ago

KB

K

K

K

KB

K

B

K

K

K

K

K

KB

KB

KB

KB

KB

KB

K

K

K

K

B

B

K

K

K

K

K

K

KB

KB

KB

KB

KB

K

B

BioGeoBEARS DEC+J on Dendrochilum M0_unconstrainedancstates: global optim, 3 areas max. d=0.4531; e=0; j=0.0092; LnL=−15.86

K

K

K

K

B

K

B

K

K

K

K

K

B

KB

B

K

B

K

K

K

K

K

K

K

B

K

K

K

K

K

B

KB

B

B

B

KB

D_billarderi

D_blumii

D_robustum

D_chilense

D_dicarpum

D_sp_Indonesia

D_reflexum

D_fasciatum

D_plurisetum

D_assimile

D_havilandii

D_leichhardtii

D_braunii

D_brevisetum

D_menziesii

D_diaphanoneurum

D_cylindrothecium

D_sp_Kinabalu_Gunung_Alab


W

W

KBW

W

W

W

W

W

W

W

W

W

W

KW

KBW

W

KBW

KBW

KBW

W

W

W

W

KBW

W

W

KBW

K

W

KBW

KBW

W

W

W

KB

BioGeoBEARS DEC+J on Dicranoloma M0_unconstrainedancstates: global optim, 3 areas max. d=0.2702; e=0; j=0; LnL=−30.34

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

KB

W

W

W

W

W

W

W

W

W

W

W

W

W

K

W

W

W

Epirixanthes_elongata

Epirixanthes_papuana

Epirixanthes_kinabaluensis

Epirixanthes_pallida

Salomonia_ciliata

Salomonia_cantoniensis


K

KBW

KBW

K

KBW

KBW

KBW

K

KBW

KBW

KBW

BioGeoBEARS DEC+J on Epirixanthes M0_unconstrainedancstates: global optim, 3 areas max. d=0.149; e=0; j=0; LnL=−2.69

K

K

K

K

KBW

K

K

K

K

KBW

Euphrasia_taurica

Euphrasia_petiolaris

Euphrasia_borneensis

Euphrasia_durietziana

Euphrasia_transmorrisonensis

Euphrasia_tarokoana

Euphrasia_nankotaizanensis

Euphrasia_philippinensis


W

W

W

W

W

W

W

W

W

K

W

W

W

W

W

BioGeoBEARS DEC+J on Euphrasia M0_unconstrainedancstates: global optim, 2 areas max. d=0; e=0; j=0.071; LnL=−2.96

W

W

W

W

W

W

W

W

W

K

W

W

W

W

Hedyotis_fissistipula_cB019_B

Hedyotis_philippensis_cB043_A

Hedyotis_rigida_bZ042_SEAsia

Hedyotis_macrostegia_Borneo

Hedyotis_pulchella_bZ029_Bor

Hedyotis_schlechteri_cA092_N

Hedyotis_valetoniana_bZ030_N

Oldenlandia_nutans_bS27_New

Oldenlandia_pubescens_cB042


W

W

B

KBW

W

K

W

W

B

KBW

BW

K

KB

W

W

W

W

BioGeoBEARS DEC+J on Hedyotis M0_unconstrainedancstates: global optim, 3 areas max. d=0.1289; e=0; j=0.0812; LnL=−15.92

W

W

B

B

W

K

W

W

W

B

B

KBW

K

K

W

W

Extended Data Figure 7 | Ancestral range estimations for Athyrium, Begonia, Coeliccia, Dacrycarpus, Dendrochilum, Dicranoloma, Epirixanthes,Euphrasia and Hedyotis. See Methods for further details. The trees are alphabetically ordered by genus name.

LETTER RESEARCH


Hygrobates_hamatoides

Hygrobates_hamatus

Hygrobates_sp_2

8 6 4 2 0Millions of years ago

K

KB

K

KB

KB

BioGeoBEARS DEC+J on Hygrobates M0_unconstrainedancstates: global optim, 2 areas max. d=0.433; e=0; j=0.1711; LnL=−1.20

K

KB

K

K

Ilex_aquifolium

Ilex_perado

Ilex_colchica

Ilex_spinigera

Ilex_bioritsensis

Ilex_georgei

Ilex_kingiana

Ilex_pernyi

Ilex_buergeri

Ilex_warburgii

Ilex_cyrtura

Ilex_ficoidea

Ilex_corallina

Ilex_fargesii2

Ilex_ciliospinosa

Ilex_fargesii1

Ilex_hookeri

Ilex_nothofagifolia

Ilex_latifolia

Ilex_hylonoma

Ilex_integra2

Ilex_liukiuensis

Ilex_matanoana

Ilex_mertensii

Ilex_x_kiusiana

Ilex_leucoclada

Ilex_x_makinoi

Ilex_cornuta1

Ilex_cornuta2

Ilex_dimorphophylla

Ilex_zhejiangensis

Ilex_integra1

Ilex_kinabaluensis

Ilex_maximowicziana

Ilex_rugosa

30 25 20 15 10 5 0Millions of years ago

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

K

W

W

BioGeoBEARS DEC+J on Ilex M0_unconstrainedancstates: global optim, 2 areas max. d=0; e=0; j=0.0109; LnL=−4.85

W

W

W

W

W

K

W

W

W

W

W

W

W

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W

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W

W

W

W

W

W

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W

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W

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W

W

W

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WW

W

W

W

W

W

W

W

W

W

WW

WW

W

W

WW

W

W

W

WW

W

WW

W

WW

W

W

Ichevalieri

Ikinabaluensis

Ieberhardtii

Ipoilanei

Iaureliana


W

W

W

W

W

K

W

W

W

BioGeoBEARS DEC+J on Impatiens M0_unconstrainedancstates: global optim, 2 areas max. d=0; e=0; j=0.2671; LnL=−2.56

W

W

K

W

W

W

W

W

K_baluensis

K_sp_1

K_interlineatus

K_sp_2

K_pleurostigma


KBW

K

K

K

K

K

W

K

KBW

BioGeoBEARS DEC+J on Kalophrynus M0_unconstrainedancstates: global optim, 3 areas max. d=0.0057; e=0; j=0.1631; LnL=−5.42

KBW

K

K

W

K

K

K

K

PK210_M_hanoii_1000to1250m

PK003_M_mariaelenae_1500to

PK073_M_sp_5_500to750mAS

PK256_M_nov_sp_2_1000to12

PK036_M_christae_2500to275

PK183_M_sp_3_1750to2000m

PK050_M_nov_sp_1_500to750

0.8 0.6 0.4 0.2 0Millions of years ago

KB

K

KB

KB

KB

KB

KB

K

KB

KB

KB

KB

KB

BioGeoBEARS DEC+J on Myrmarachne M0_unconstrainedancstates: global optim, 2 areas max. d=5; e=0; j=1.9; LnL=−1.49

K

B

B

B

KB

B

B

K

K

K

K

B

Nepenthes_gracillima

Nepenthes_sanguinea

Nepenthes_alba

Nepenthes_ramispina


Nepenthes_bokorensis

Nepenthes_vieillardii

Nepenthes_andamana

Nepenthes_kongkandana

Nepenthes_singalana


Nepenthes_benstonei


Nepenthes_specatabilis

Nepenthes_insignis

Nepenthes_bongso



Nepenthes_alata

Nepenthes_campanulata

Nepenthes_truncata

Nepenthes_peltata


Nepenthes_villosa

Nepenthes_hurrelliana

Nepenthes_fusca

Nepenthes_vogelii

Nepenthes_boschiana

Nepenthes_maxima

Nepenthes_glandulifera

Nepenthes_chaniana

Nepenthes_lowii

Nepenthes_rajah

Nepenthes_x_kinabaluensis

Nepenthes_northiana


KBW

KBW

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

BW

W

K

K

K

KB

B

B

B

B

KB

B

K

W

W

W

W

BW

W

W

W

W

W

W

W

BW

W

W

W

W

W

W

B

W

W

K

K

B

B

B

B

W

B

B

KB

K

K

B

BioGeoBEARS DEC+J on Nepenthes M0_unconstrainedancstates: global optim, 3 areas max. d=0.0074; e=0; j=0.0156; LnL=−29.45

B

K

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

W

WW

K

K

K

B

W

B

BB

KB

B

K

KBW

W

W

W

W

W

W

W

WW

WW

W

W

W

WW

W

WW

W

B

W

KK

B

B

B

B

B

B

BB

K

Paphiopedilum_randsii

Paphiopedilum_philippinense

Paphiopedilum_stonei

Paphiopedilum_kolopakingii

Paphiopedilum_rothschildianum

Paphiopedilum_glaucophyllum

Paphiopedilum_liemianum

Paphiopedilum_primulinum


BW

BW

BW

B

B

W

W

W

BW

B

B

K

W

W

W

BioGeoBEARS DEC+J on Paphiopedilum M0_unconstrainedancstates: global optim, 3 areas max. d=0.0149; e=0; j=0.0848; LnL=−8.51

W

B

BW

K

B

W

W

BW

BW

W

B

B

W

W

e4003829276_P_gigas_830mA

e4003829322_P_microphylla_6

e4003829348_P_kinabaluensis

Payena_obscura_EDNA09_023

Payena_ferruginea_EDNA09_0

Payena_leerii_EDNA09_02317

Payena_lucida_EDNA09_0230

15 10 5 0Millions of years ago

B

B

B

B

B

B

B

KB

K

BW

B

BW

BW

BioGeoBEARS DEC+J on Payena M0_unconstrainedancstates: global optim, 3 areas max. d=0.0575; e=0; j=0.1107; LnL=−12.85

B

B

B

B

B

B

B

B

B

B

B

K

SP13507_Philalanka_spec1_25

SP13519_Philalanka_thienema




B

K

K

B

B

K

B

BioGeoBEARS DEC+J on Philalanka M0_unconstrainedancstates: global optim, 2 areas max. d=0; e=0; j=3; LnL=−1.17

K

B

K

B

K

B

Extended Data Figure 8 | Ancestral range estimations for Hygrobates, Ilex, Impatiens, Kalophrynus, Myrmarachne, Nepenthes, Paphiopedilum, Payenaand Philalanka. See Methods for further details. The trees are alphabetically ordered by genus name.

RESEARCH LETTER


Ptomaphaginus_latescens

Ptomaphaginus_similipes

Ptomaphaginus_kinabaluensis

Ptomaphaginus_cf_latimanus

Ptomaphaginus_bryantioides

2 1.5 1 0.5 0Millions of years ago

B

B

K

K

KB

B

K

KB

B

BioGeoBEARS DEC+J on Ptomaphaginus M0_unconstrainedancstates: global optim, 3 areas max. d=0.5097; e=0; j=0.6198; LnL=−4.20

B

B

K

B

K

K

B

K

Ranunculus_amerophyllus

Ranunculus_basilobatus

Ranunculus_pseudolowii

Ranunculus_brassii

Ranunculus_saruwagedicus

Ranunculus_lowii

Ranunculus_cantoniensis

Ranunculus_silerifolius

Ranunculus_chinensis

1 0.8 0.6 0.4 0.2 0Millions of years ago

W

W

W

W

W

W

W

W

W

W

W

W

W

B

W

W

W

BioGeoBEARS DEC+J on Ranunculus M0_unconstrainedancstates: global optim, 2 areas max. d=0; e=0; j=0.0742; LnL=−3.50

W

W

B

W

W

W

W

W

W

W

W

W

W

W

W

W

Rhododendron_acuminatum

Rhododendron_fallacinum

Rhododendron_bagobonum

Rhododendron_stenophyllum

Rhododendron_crassifolium

Rhododendron_exuberans

Rhododendron_javanicum

Rhododendron_lowii

Rhododendron_orbiculatum

Rhododendron_suaveolens

Rhododendron_stapfianum

Rhododendron_praetervisum

Rhododendron_rugosum

Rhododendron_buxifolium

Rhododendron_borneense

Rhododendron_cuneifolium

Rhododendron_ericoides

Rhododendron_santapaui


K

K

KB

K

K

K

KBW

KB

KBW

K

K

K

KB

K

KW

KB

K

K

KB

KBW

KB

KB

KB

KBW

KB

KBW

KB

KB

KB

KB

K

KB

KB

K

W

BioGeoBEARS DEC+J on Rhododendron M0_unconstrainedancstates: global optim, 3 areas max. d=0.1291; e=0.0222; j=0; LnL=−38.63

K

K

K

K

K

K

K

K

K

K

K

K

K

K

W

K

K

K

K

K

K

K

K

K

K

KBW

K

K

K

K

K

K

K

K

Teleopsis_sabah

Teleopsis_trichophora

Teleopsis_rubicunda

Teleopsis_breviscopium

Teleopsis_pallifacies

Teleopsis_scapulistylus

Teleopsis_pseudogracilifemur

Teleopsis_gracilifemur

Teleopsis_nr_discrepans

Teleopsis_sp_2

Teleopsis_nr_discrepans_sp_B

Teleopsis_sabah_sp_2

Teleopsis_discrepans

Teleopsis_nr_discrepans_sp_C

Teleopsis_adjacens


B

B

W

W

W

B

B

K

B

B

B

B

B

B

B

W

W

W

KB

B

K

KB

B

B

B

B

KB

KB

W

BioGeoBEARS DEC+J on Teleopsis M0_unconstrainedancstates: global optim, 3 areas max. d=0.0173; e=0; j=0.1103; LnL=−24.05

B

B

W

W

B

B

B

K

B

B

B

B

B

W

B

W

W

W

W

B

B

K

B

B

B

B

B

KB

Trichosanthes_borneensis

Trichosanthes_globosa

Trichosanthes_kinabaluensis

Trichosanthes_obscura

Trichosanthes_intermedia

Trichosanthes_montana

Trichosanthes_sepilokensis


BW

B

W

B

B

BW

BW

W

K

B

B

BW

B

BioGeoBEARS DEC+J on Trichosanthes M0_unconstrainedancstates: global optim, 3 areas max. d=0.019; e=0; j=0.2724; LnL=−9.98

B

B

W

B

B

B

B

W

W

B

K

BW

SP13398_T_kinabalensis_kina

Tritetrabdella_taiwana_A

Tritetrabdella_scandens

Haemadipsa_cavatuses


W

K

W

K

W

W

W

BioGeoBEARS DEC+J on Tritetrabdella M0_unconstrainedancstates: global optim, 2 areas max. d=0; e=0; j=0.3937; LnL=−1.83

W

W

W

K

K

W

Cunonia_atrorubens

Cunonia_pulchella

Cunonia_balansae

Cunonia_montana

Cunonia_macrophylla

Cunonia_capensis

Pancheria_brunhesii

Pancheria_hirsuta

Pancheria_reticulata

Pancheria_engleriana

Pancheria_phylliraeoides

Weinmannia_dichotoma

Weinmannia_serrata

Weinmannia_parviflora

Weinmannia_samoensis

Weinmannia_raiateensis

Weinmannia_vescoi

Weinmannia_vitiensis


Weinmannia_racemosa

Weinmannia_silvicola

Weinmannia_arguta

Weinmannia_sanguisugarum

Weinmannia_stenostachya

Weinmannia_bojeriana

Weinmannia_humbertiana

Weinmannia_minutiflora

Weinmannia_marojejyensis

Weinmannia_madagascariensi

Weinmannia_rutenbergii


Weinmannia_clemensiae

Weinmannia_exigua

Weinmannia_richii

Weinmannia_fraxinea

Weinmannia_hooglandii


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BioGeoBEARS DEC+J on Weinmannia M0_unconstrainedancstates: global optim, 3 areas max. d=6e−04; e=0; j=0; LnL=−12.42

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Extended Data Figure 9 | Ancestral range estimations for Ptomaphaginus, Ranunculus, Rhododendron, Teleopsis, Trichosanthes, Tritetrabdella andWeinmannia. See Methods for further details. The trees are alphabetically ordered by genus name.

LETTER RESEARCH


Extended Data Table 1 | Details for all studied endemic taxa

Plants are highlighted in green, fungi in blue, and animals in brown.*For clade numbering in Cortinarius, see Supplementary Table 2.**Centric/eccentric status determined via biogeographic analysis (see Extended Data 4).*** For definitions and sources, see Supplementary Table 3.****K, Mount Kinabalu; T, Mount Tambuyukon; C, Crocker Range; TM, Mount Trus Madi.*****Manuscript name.

RESEARCH LETTER


evolutionofendemismonayoungtropicalmountain vincent s. f. t. merckx1,2, kasper p. hendriks1,3, kevin...

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