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LETTERdoi:10.1038/nature14949
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)
C. spp. (2,040–2,100 m)
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C. collinitus (Germany)
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C. sp. (1,900 m)
C. vitreopileatus (New Zealand)
C. spp. (2,900 m)
C. sp. (2,040–2,100 m)
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)
Impatiens
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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.
RESEARCH LETTER
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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).
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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.
LETTER RESEARCH
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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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3 5 0 | N A T U R E | V O L 5 2 4 | 2 0 A U G U S T 2 0 1 5
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’.
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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
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0.06
RBINS.15995 K. sp.2
RBINS.15993 K. sp.2
RBINS.16009 K. baluensis
AB611921 Kalophrynus pleuros�gma
RBINS.16096 K. baluensis
RBINS.16010 K. baluensis
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.16023 K. baluensis
RBINS.15992 K. sp.1
RBINS.16043 K. baluensis
RBINS.16063 K. baluensis
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.PK174 M. mariaelenae
NMF2012-01399.PK020 M. mariaelenae
NMF2012-01399.PK254 M. malayana
NMF2012-01399.PK158 M. malayana
NMF2012-01399.PK069 M. malayana
NMF2012-01399.PK262 M. malayana
NMF2012-01399.PK098 M. malayana
NMF2012-01399.PK016 M. mariaelenae
NMF2012-01399.PK312 M. malayana
NMF2012-01399.PK305 M. malayana
NMF2012-01399.PK109 M. malayana
NMF2012-01399.PK373 M. cornutaAY297412 M. assimilis (Philippines)
NMF2012-01399.PK080 M. sp.5
NMF2012-01399.PK214 M. mariaelenae
NMF2012-01399.PK186 M. mariaelenae
NMF2012-01399.PK265 M. malayana
NMF2012-01399.PK306 M. malayana
NMF2012-01399.PK221 M. mariaelenae
NMF2012-01399.PK291 M. malayana
NMF2012-01399.PK210 M. hanoii
NMF2012-01399.PK003 M. mariaelenae
NMF2012-01399.PK023 M. mariaelenae
NMF2012-01399.PK191 M. mariaelenae
EU815599 Ligonipes sp. (Australia)
NMF2012-01399.PK260 M. nov.sp.2
NMF2012-01399.PK068 M. malayana
NMF2012-01399.PK249 M. malayana
NMF2012-01399.PK311 M. malayana
NMF2012-01399.PK285 M. malayana
NMF2012-01399.PK256 M. nov.sp.2
NMF2012-01399.PK222 M. mariaelenae
NMF2012-01399.PK263 M. malayana
NMF2012-01399.PK116 M. malayana
NMF2012-01399.PK008 M. mariaelenae
NMF2012-01399.PK052 M. sp.4
NMF2012-01399.PK017 M. mariaelenae
NMF2012-01399.PK198 M. mariaelenae
NMF2012-01399.PK244 M. mariaelenae
NMF2012-01399.PK097 M. malayana
NMF2012-01399.PK259 M. malayana
NMF2012-01399.PK175 M. mariaelenae
NMF2012-01399.PK252 M. mariaelenae
NMF2012-01399.PK266 M. malayana
NMF2012-01399.PK137 M. malayana
NMF2012-01399.PK226 M. mariaelenae
NMF2012-01399.PK083 M. malayana
NMF2012-01399.PK250 M. malayana
NMF2012-01399.PK298 M. malayana
NMF2012-01399.PK103 M. malayana
NMF2012-01399.PK308 M. malayana
NMF2012-01399.PK245 M. malayana
NMF2012-01399.PK264 M. nov.sp.2
NMF2012-01399.PK009 M. mariaelenae
NMF2012-01399.PK246 M. mariaelenae
NMF2012-01399.PK014 M. mariaelenae
NMF2012-01399.PK019 M. mariaelenae
NMF2012-01399.PK144 M. malayana
NMF2012-01399.PK179 M. mariaelenae
NMF2012-01399.PK290 M. malayana
NMF2012-01399.PK013 M. mariaelenae
JX145678 M. evidens (Gabon)
NMF2012-01399.PK286 M. malayana
NMF2012-01399.PK213 M. mariaelenaeNMF2012-01399.PK215 M. mariaelenae
NMF2012-01399.PK257 M. nov.sp.2
NMF2012-01399.PK011 M. mariaelenae
NMF2012-01399.PK081 M. sp.5
NMF2012-01399.PK352 M. mariaelenae
NMF2012-01399.PK015 M. mariaelenae
NMF2012-01399.PK050 M. nov.sp.1
NMF2012-01399.PK005 M. mariaelenae
NMF2012-01399.PK183 M. sp.3
NMF2012-01399.PK026 M. mariaelenae
NMF2012-01399.PK177 M. mariaelenae
NMF2012-01399.PK010 M. mariaelenae
NMF2012-01399.PK219 M. mariaelenae
NMF2012-01399.PK225 M. mariaelenae
NMF2012-01399.PK320 M. malayana
NMF2012-01399.PK022 M. mariaelenae
NMF2012-01399.PK247 M. mariaelenae
NMF2012-01399.PK224 M. mariaelenae
NMF2012-01399.PK206 M. sp.3
NMF2012-01399.PK227 M. mariaelenae
NMF2012-01399.PK036 M. christae
NMF2012-01399.PK248 M. malayana
NMF2012-01399.PK027 M. mariaelenae
NMF2012-01399.PK304 M. malayana
NMF2012-01399.PK073 M. sp.5
NMF2012-01399.PK317 M. malayana
NMF2012-01399.PK217 M. mariaelenae
JX145679 M. foenisex (Gabon)
NMF2012-01399.PK012 M. mariaelenae
NMF2012-01399.PK223 M. mariaelenae
NMF2012-01399.PK051 M. sp.4
NMF2012-01399.PK315 M. malayana
NMF2012-01399.PK255 M. malayana
NMF2012-01399.PK220 M. mariaelenae
NMF2012-01399.PK293 M. malayana
NMF2012-01399.PK018 M. mariaelenae
NMF2012-01399.PK261 M. malayana
NMF2012-01399.PK316 M. malayana
NMF2012-01399.PK216 M. mariaelenae
NMF2012-01399.PK303 M. malayana
NMF2012-01399.PK307 M. malayana
NMF2012-01399.PK301 M. malayana
NMF2012-01399.PK357 M.cornuta
NMF2012-01399.PK178 M. mariaelenae
NMF2012-01399.PK077 M. sp.5
NMF2012-01399.PK007 M. mariaelenae
JX145676 M. cf. gedongensis (Malaysia)
NMF2012-01399.PK200 M. mariaelenae
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
e4002185538 A. clemensiae
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)
e4002185515 A. amoenum
e4002185551 A. amoenum
EU329028 EU329072 A. clivicola (Japan)
e4002185691 A. amoenum
EU329038 EU329081 A. melanolepis (Japan)
EU329043 EU329086 A. oblitescens (Japan)
EU329034 EU329077 A. iseanum (Japan)
e4002185621 A. clemensiae
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)
e4002185526 A. clemensiae
e4002185631 A. amoenum
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)
FL033 I. kinabaluensis
CM0025 I. kinabaluensis
DQ147845 I. flaccida (India)
CM0028 I. kinabaluensis
DQ147839 I. eberhard�i (Vietnam)
DQ147832 I. cuspidata (India)
FL007 I. kinabaluensis
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.
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Coleoptera - Ptomaphaginus
0.08
RMNH.INS.549270 Ptomaphaginus bryan�oides
RMNH.INS.555626 Ptomaphaginus bryan�oides
RMNH.INS.555623 Ptomaphaginus ater
RMNH.INS.555603 Ptomaphaginus kinabaluensis
RMNH.INS.555629 Ptomaphaginus kinabaluensis
RMNH.INS.555633 Ptomaphaginus.nr. fagei
RMNH.INS.555610 Ptomaphaginus similipes
RMNH.INS.555611 Ptomaphaginus c.f. la�manus
RMNH.INS.555594 Ptomaphaginus bryan�oides
RMNH.INS.555592 Ptomaphaginus similipes
RMNH.INS.555619 Ptomaphaginus n.sp. bryan� complex
RMNH.INS.555604 Ptomaphaginus kinabaluensis
RMNH.INS.549243 Prionochaeta harmandi (Japan)
RMNH.INS.555625 Ptomaphaginus bryan�oides
RMNH.INS.555591 Ptomaphaginus bryan�oides
RMNH.INS.555593 Ptomaphaginus similipes
RMNH.INS.555681 Ptomaphaginus similipes
RMNH.INS.555645 Ptomaphaginus bryan�oides
RMNH.INS.549271 Ptomaphaginus bryan�oides
RMNH.INS.555618 Ptomaphaginus n.sp. bryan� complex
RMNH.INS.555607 Ptomaphaginus latescens
RMNH.INS.555631 Ptomaphaginus kinabaluensisRMNH.INS.555601 Ptomaphaginus kinabaluensis
RMNH.INS.549263 Ptomaphaginus similipes
RMNH.INS.555602 Ptomaphaginus kinabaluensis
RMNH.INS.555680 Ptomaphaginus similipes
RMNH.INS.555598 Ptomaphaginus bryan�oides
RMNH.INS.555628 Ptomaphaginus bryan�oides
RMNH.INS.555621 Ptomaphaginus n.sp. bryan� complex
RMNH.INS.555616 Ptomaphaginus similipes
RMNH.INS.555677 Ptomaphaginus bryan�oides
RMNH.INS.555644 Ptomaphaginus bryan�oides
RMNH.INS.555599 Ptomaphaginus bryan�oides
RMNH.INS.555606 Ptomaphaginus kinabaluensis
RMNH.INS.555637 Ptomaphaginus bryan�oides
RMNH.INS.549268 Ptomaphaginus bryan�oides
RMNH.INS.555612 Ptomaphaginus similipes
RMNH.INS.555632 Ptomaphaginus nr. fagei
RMNH.INS.555627 Ptomaphaginus bryan�oides
RMNH.INS.555685 Ptomaphaginus similipes
RMNH.INS.549269 Ptomaphaginus bryan�oides
RMNH.INS.555620 Ptomaphaginus n.sp. bryan� complex
RMNH.INS.555600 Ptomaphaginus bryan�oides
RMNH.INS.555609 Ptomaphaginus similipes
RMNH.INS.555624-Ptomaphaginus ater
RMNH.INS.555622 Ptomaphaginus n.sp. bryan� complex
RMNH.INS.555605 Ptomaphaginus kinabaluensis
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)
e4002189764 D. braunii
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)
MS2_e4002189750 D. brevisetum
e4002189755 D. havilandii
L0856640 e4002189709 D. sp. (Indonesia)
e4002189740 D. braunii
e4002189743 D. havilandii
e4002189789 D. assimile
DQ462597 DQ464197 D. billarderi (Reunion)
DQ462606 DQ464185 D. plurisetum (New Zealand)
e4002189742 D. assimile
e4002189777 D. assimile
KF423992 KF423638 Holomitrium crispulum
e4002189766 D. braunii
L0093198 e4002189685 D. reflexum (Indonesia)
e4002189767 D. havilandii
e4002189776 D. blumii
e4002189765 D. assimile
e4002189814 D. blumii
L0092869 e4002189708 D. diaphanoneurum (Australia)
e4002189790 D. assimile
AF129587 AF144113 Dicranum polysetum
DQ462595 DQ464194 D. billarderi (Chile)
e4002189788 D. brevisetum
e4002189815 D. assimile
MS4_e4002189774 D. brevisetum
MS3_e4002189762 D. brevisetum
e4002189827 D. assimile
DQ485179 D. menziesii (Australia)
e4002189787_D_reflexum_Kinabalu_Sayap
DQ462600 DQ464190 D. robustum (Chile)
KF423995 KF423641 Holomitrium arboreum
DQ485178 D. fasciatum (Australia)
e4002189741 D. assimile
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.557968 Teleopsis pallifacies
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
RMNH.INS.555128 Teleopsis pallifacies
RMNH.INS.557932 Teleopsis pseudogracilifemur sp.n.
RMNH.INS.555122 Teleopsis pseudogracilifemur sp.n.
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
G2015 Macmillan Publishers Limited. All rights reserved
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
SNP21938 R. crassifolium
SNP32303 R. ericoides
SNP32305 R. lowii
SNP32304 R. buxifolium
SNP21935 R. javanicum
SNP21914 R. suaveolens
SNP21911 R. exuberans
SNP21925 R. simulans
SNP21941 R. crassifolium
SNP21917 R. sp.
SNP21932 R. praetervisum
SNP28504 R. rugosum
SNP21907 R. bagobonum
SNP28515 R. cuneifolium
SNP28509 R. acuminatum
GU176637 AF394258 R. camtscha�cum
SNP21919 R. javanicum
SNP28508 R. rugosum
SNP21956 R. fallacinum
SNP21955 R. orbiculatum
SNP21937 R. crassifolium
SNP32307 R. buxifolium
SNP21929 R. suaveolens
SNP28514 R. cuneifolium
SNP28517 R. stenophyllum
SNP21915 R. stenophyllum
SNP21936 R. borneense
SNP21908 R. bur�i
SNP28510 R. rugosum
SNP21920 R. fallacinum
SNP21926 R. cf. stapfianum
SNP21906 R. rugosum
SNP28516 R. cf. cuneifolium
SNP32301 R. buxifolium
SNP21905 R. suaveolens
SNP28505 R. stenophyllum
SNP21916 R p.
SNP21913 R. fallacinum
SNP21933 R. orbiculatum
SNP21921 R. cuneifolium
SNP21945 R. fallacinum
SNP28502 R. fallacinum
SNP28513 R. lowii
SNP21923 R. crassifolium
SNP21909 R cf. rugosum
SNP21934 R. suaveolens
SNP21904 R. crassifolium
SNP21922 R. crassifolium
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
SP13522 Philalanka thienemanni
JX988066 Subulina octona
SP13508 Philalanka sp.1
SP13518 Philalanka thienemanni
SP13520 Philalanka thienemanni
HQ171582-Zospeum exiguum (Slovenia)
SP13523 Philalanka thienemanni
SP13517 Philalanka thienemanni
SP13506 Philalanka sp.1
HQ171529 Carychium mexicanum (USA)
SP13428 Ibycus rachelae
SP13507 Philalanka sp.1
SP13511 Philalanka sp.3
SP13521 Philalanka thienemanni
SP13519 Philalanka thienemanni
SP13509 Philalanka sp.1
SP13492 Philalanka sp.3
SP13516 Philalanka thienemanni
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
AJ557009 Glomus intraradices
J437105 Glomus sinuosum
OTU 27453
OTU 24836
OTU 8898
OTU 6655
OTU 4196
FM865562 Glomus intraradices
OTU 27338
OTU 29948
OTU 33992
OTU 12813
OTU 20709
OTU 26364
AM713407 Glomus eburneum
OTU 8632
FJ009600 Glomus intraradices
OTU 35053
OTU 3458
FM865573 Glomus intraradices
OTU 16590
OTU 7081
OTU 29995
EF989103 Glomus intraradices
OTU 5441
AJ972462 Glomus diaphanum
OTU 21220
FM865568 Glomus intraradices
FJ009598 Glomus intraradices
OTU 5233
OTU 12112OTU 24971
OTU 18850
OTU 13666
OTU 8446
FM865572 Glomus intraradices
OTU 27919
OTU 17823
OTU 32979
OTU 5542
EF989105 Glomus intraradices
OTU 6588
OTU 21269
AJ437106 Glomus sinuosum
OTU 1606
AJ567351 Glomus intraradices
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)
HQ322469 T. taiwana (Taiwan)
SP13382 T. kinabaluensis inobongensis
HQ203161 P. pungens (Australia)
HQ203195 T. taiwana (China)
SP13407 H. sp.
SP13380 T. kinabaluensis inobongensis
HQ203194 T. scandens (Thailand)
HQ322466 T. taiwana (Taiwan)
HQ322468 T. taiwana (Taiwan)
AB685259 T. taiwana (China)
HQ322465 T. taiwana (Taiwan)
SP13398 T. kinabaluensis kinabaluensis
SP13381 T. kinabaluensis inobongensis
HQ322463 T. taiwana (Taiwan)
HQ322467 T. taiwana (Taiwan)
SP13383 T. kinabaluensis inobongensis
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.555471 R. simulata
RMNH.INS.555473 R. simulata
RMNH.INS.555444 R. simulata
RMNH.INS.555221 R. silau
RMNH.INS.555260 R. samardaca
RMNH.INS.555246 R. samardaca
RMNH.INS.555452 Mesovelia horvathi
RMNH.INS.555248 R. samardaca
RMNH.INS.555250 R. samardaca
RMNH.INS.555466 R. simulata
RMNH.INS.555212 R. ranau
RMNH.INS.555254 R. samardaca
RMNH.INS.555259 R. samardaca
RMNH.INS.555219 R. ranau
RMNH.INS.555217 R. ranau
RMNH.INS.555238 R. silau
RMNH.INS.555255 R. samardaca
RMNH.INS.555243 R. samardaca
RMNH.INS.555242 R. samardaca
RMNH.INS.555256 R. samardaca
RMNH.INS.555247 R. samardaca
RMNH.INS.555244 R. samardaca
RMNH.INS.555440 R. simulata
RMNH.INS.555249 R. samardaca
RMNH.INS.555224 R. silau
RMNH.INS.555215 R. ranau
RMNH.INS.555214 R. ranau
RMNH.INS.555245 R. samardaca
RMNH.INS.555225 R. silau
RMNH.INS.555474 R. simulata
RMNH.INS.555253 R. samardaca
RMNH.INS.555252 R. samardaca
RMNH.INS.555241 R. samardaca
RMNH.INS.555258 R. samardaca
RMNH.INS.555257 R. samardaca
RMNH.INS.555445 R. simulata
RMNH.INS.555270 R. simulata
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
G2015 Macmillan Publishers Limited. All rights reserved
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.4149358 N. tentaculata
L.4149384 N. gracilis
L.4149394 N. x kinabaluensis
L.4149083 N. mirabilis
L.4149079 N. mirabilis
L.4149113 N. mirabilis
L.4149109 N. gracilis
L.4149389 N. villosa
L.4149386 N. tentaculata
L.4149117 N. edwardsiana
L.4149385 N. tentaculata
L.4149093 N. edwardsiana
L.4149383 N. mirabilis
L.4149120 N. tentaculata
L.4149395 N. tentaculata
L.4149373 N. lowii
L.4149078 N. gracilis
L.4149396 N. gracilis
L.4149107 N. villosa
L.4149101 N. tentaculata
L.4149380 N. fusca L.4149381 N. fusca
L.4149355-6 N. x harryana
L.4149089 N. mirabilis L.4149365 N. gracilis
L.4149106 N. edwardsiana
AF315885 N. pervillei (Seychelles)
L.4149096 N. tentaculata
L.4149359 N. x hookeriana
L.4149388 N. tentaculata
L.4149097 N. tentaculata
L.4149390 N. tentaculata
L.4149371 N. tentaculata
L.4149354 N. villosa
L.4149087 N. tentaculata
SNP34297 N. rajah
L.4149378 N. lowii
L.4149084 N. mirabilis
L.4149392 N. x kinabaluensis
L.4149115 N. tentaculata
L.4149357 N. x kinabaluensis
L.4149091 N. tentaculata
L.4149114 N. mirabilis
L.4149084 N. mirabilis
L.4149074-77 N. edwardsiana
L.4149082 N. mirabilis
L.4149124 N. lowii
L.4149110 N. tentaculata
L.4149367 N. ampullaria
L.4149080 N. gracilis
L.4149361 N. ampullaria
L.4149391 N.villosa
L.4149382 N. fusca
L.4149090 N. gracilis
L.4149368 N. fusca
L.4149088 N. tentaculata
L.4149377 N. gracilis
L.4149393 N. villosa
L.4149098 N. tentaculata
L.4149379 N. tentaculata
L.4149099 N. tentaculata
L.4149370 N. chaniana
L.4149362 N. ampullaria
L.4149081 N. mirabilis
L.4149095 N. tentaculata
L.4149086 N. tentaculata
L.4149387 N. tentaculata
L.4149092 N. gracilis
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.507677 C. sp.
RMNH.INS.501072 C. albicauda (Malaysia)
RMNH.INS.507735 C. nemoricola
RMNH.INS.501021 C. octogesima (Singapore)
RMNH.INS.503977 C. octogesima (Singapore)
RMNH.INS.506849 C. albicauda (Malaysia)
RMNH.INS.506487 C. cyanomelas (Hong Kong)
RMNH.INS.228944 C. octogesima (Malaysia)
RMNH.INS.507675 C. sp.
RMNH.INS.507734 C. nemoricola
RMNH.INS.506468 C. cyanomelas (Hong Kong)
RMNH.INS.503932 C. didyma (Terengganu Malaysia)
RMNH.INS.509174 C. poungyi (Thailand)
RMNH.INS.506475 C. cyanomelas (Hong Kong)
RMNH.INS.507678 C. sp.
RMNH.INS.507775 C. nemoricola
RMNH.INS.507721 C. nemoricola
RMNH.INS.505734 C. poungyi (Thailand)
RMNH.INS.506836 C. octogesima (Malaysia)
RMNH.INS.507768 C. nemoricola
RMNH.INS.506294 C. nemoricola (Sarawak Malaysia)
RMNH.INS.506304 C. nemoricola (Sarawak Malaysia)
RMNH.INS.507780 C. nemoricola
RMNH.INS.507792 C. nemoricola
RMNH.INS.507782 C. nemoricola
RMNH.INS.507793 C. nemoricola
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
CM002-1 (L) E. pallida
CM013 (L) E. kinabaluensis
CM002-2 (L) E. pallida
CM032 (L) E. elongata
CM001-1 (L) E. pallida
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
FL050 H. macrostegia
FL077 H. pulchella
FL055 H. macrostegia
FL054 H. macrostegia
FL042 H. pulchella
cB019 H. fissis�pula (Borneo)
FL049 H. macrostegia
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.1351 H. hamatoides
RMNH.ACA.1429 Atrac�des cognatus
RMNH.ACA.1355 H. hamatoides
RMNH.ACA.1360 H. acu�palpis
RMNH.ACA.1438 Atrac�des cf. propatulus
RMNH.ACA.1364 H. hamatus
RMNH.ACA.1354 H. hamatoides
RMNH.ACA.1352 H. hamatoides
RMNH.ACA.1357 H. hamatoides
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
G2015 Macmillan Publishers Limited. All rights reserved
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
Dacrycarpus_kinabaluensis913
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
Dacrycarpus_imbricatus102
Phyllocladus_aspleniifolius
Podocarpus_degeneri
Podocarpus_costalis
Dacrycarpus_imbricatus106
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
DQ462599_DQ464189_D_robustum_New_Zealand
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
MS2_e4002189750_D_brevisetum_Kinabalu_Sayap
DQ462594_DQ464193_D_billarderi_New_Zealand
DQ462596_DQ464196_D_billarderi_Rwanda
L0856640_e4002189709_D_sp_Indonesia_Java
MS47_e4002189815_D_assimile_Kinabalu_Gunung_Alab
DQ462595_DQ464194_D_billarderi_Chile
AF135075_AF144116_Paraleucobryum_enerve
MS43_e4002189767_D_havilandii_Kinabalu_Laban_Rata
KF423995_KF423641_Holomitrium_arboreumKF423992_KF423638_Holomitrium_crispulum
MS41_e4002189743_D_havilandii_Kinabalu_Laban_Rata
MS42_e4002189755_D_havilandii_Kinabalu_Laban_Rata
DQ462606_DQ464185_D_plurisetum_New_Zealand
MS9_e4002189739_D_brevisetum_Kinabalu_Sayap
DQ462607_DQ464184_D_menziesii_New_Zealand
MS16_e4002189823_D_braunii_Kinabalu_Sayap
DQ485174_D_plurisetum_New_ZealandMS25_e4002189741_D_assimile_Kinabalu_Sayap
MS12_e4002189775_D_reflexum_Kinabalu_Sayap
MS45_e4002189791_D_assimile_Kinabalu_Gunung_Alab
MS28_e4002189777_D_assimile_Kinabalu_KNP
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
MS21_e4002189788_D_brevisetum_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
MS4_e4002189774_D_brevisetum_Kinabalu_Sayap
DQ485178_D_fasciatum_Australia
DQ462597_DQ464197_D_billarderi_Reunion
MS11_e4002189763_D_brevisetum_Kinabalu_Sayap
DQ462601_DQ464191_D_robustum_Chile
MS51_e4002189768_D_assimile_Kinabalu_Gunung_Alab
MS13_e4002189787_D_reflexum_Kinabalu_Sayap
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
MS12_e4002189775_D_reflexum_Kinabalu_Sayap
AF135067_AF140696_Dicnemon_calycinum
DQ485176_D_leichhardtii_Australia
MS51_e4002189768_D_assimile_Kinabalu_Gunung_Alab
DQ462604_DQ464186_D_chilense_Chile
MS3_e4002189762_D_brevisetum_Kinabalu_Sayap
MS44_e4002189779_D_havilandii_Kinabalu_Laban_Rata
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
MS4_e4002189774_D_brevisetum_Kinabalu_Sayap
DQ462595_DQ464194_D_billarderi_Chile
MS13_e4002189787_D_reflexum_Kinabalu_Sayap
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
MS45_e4002189791_D_assimile_Kinabalu_Gunung_Alab
MS9_e4002189739_D_brevisetum_Kinabalu_Sayap
MS43_e4002189767_D_havilandii_Kinabalu_Laban_Rata
AF135075_AF144116_Paraleucobryum_enerve
MS19_e4002189764_D_braunii_Kinabalu_Sayap
MS29_e4002189789_D_assimile_Kinabalu_KNP
MS47_e4002189815_D_assimile_Kinabalu_Gunung_Alab
L0856640_e4002189709_D_sp_Indonesia_JavaL0093198_e4002189685_D_reflexum_Indonesia_Sumatra
DQ462599_DQ464189_D_robustum_New_Zealand
MS21_e4002189788_D_brevisetum_Kinabalu_Sayap
DQ462596_DQ464196_D_billarderi_Rwanda
DQ462601_DQ464191_D_robustum_Chile
DQ462606_DQ464185_D_plurisetum_New_Zealand
DQ462598_DQ464188_D_robustum_New_Zealand
MS25_e4002189741_D_assimile_Kinabalu_SayapDQ485174_D_plurisetum_New_Zealand
MS42_e4002189755_D_havilandii_Kinabalu_Laban_Rata
L0092869_e4002189708_D_diaphanoneurum_Australia
MS33_e4002189742_D_assimile_Kinabalu_KNP
DQ462605_DQ485175_D_dicarpum_New_Zealand
MS52_e4002189780_D_blumii_Kinabalu_Gunung_Alab
MS41_e4002189743_D_havilandii_Kinabalu_Laban_Rata
MS20_e4002189776_D_blumii_Kinabalu_L2
MS2_e4002189750_D_brevisetum_Kinabalu_Sayap
DQ462600_DQ464190_D_robustum_Chile
MS17_e4002189740_D_braunii_Kinabalu_Sayap
MS48_e4002189827_D_assimile_Kinabalu_Gunung_Alab
DQ462594_DQ464193_D_billarderi_New_Zealand
MS16_e4002189823_D_braunii_Kinabalu_Sayap
MS37_e4002189790_D_assimile_Kinabalu_L2
MS11_e4002189763_D_brevisetum_Kinabalu_Sayap
KF423978_KF423621_D_cylindrothecium_Russia
AF129587_AF144113_Dicranum_polysetum
MS28_e4002189777_D_assimile_Kinabalu_KNP
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
Hedyotis_sp_bZ022_Vietnam
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_sp_bZ018_Vietnam
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
Hedyotis_sp_bZ021_Vietnam
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
Hedyotis_sp_bZ024_Vietnam
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
Hedyotis_sp_bZ013_Vietnam
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
Hedyotis_sp_bZ025_Vietnam
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
G2015 Macmillan Publishers Limited. All rights reserved
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_edwardsiana3
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_edwardsiana2
Nepenthes_singalana
Nepenthes_mira
Nepenthes_mikei
Nepenthes_merrilliana
Nepenthes_dubia
Nepenthes_reinwardtiana
Nepenthes_villosa2
Nepenthes_sanguinea
Nepenthes_gymnamphora
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_ventricosa
Nepenthes_insignis
Nepenthes_glabrata
Nepenthes_bicalcarata
Nepenthes_fusca
Dioncophyllum_thollonii
Nepenthes_burbidgeae
Nepenthes_macrovulgaris
Nepenthes_edwardsiana3
Nepenthes_mirabilis
Nepenthes_albomarginata
Nepenthes_bongso
Nepenthes_pilosa
Nepenthes_spathulata
Nepenthes_ramispina
Nepenthes_ovata
Nepenthes_gracilis2
Nepenthes_lavicola
Nepenthes_macfarlanei
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_sp_Bradford_435
Weinmannia_raiateensis
Weinmannia_sp_Bradford_837
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_truncata1
Trichosanthes_laceribractea3
Trichosanthes_laeoica2
Echinocystis_lobata
Trichosanthes_lepiniana3
Trichosanthes_odontosperma2
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
Trichosanthes_lepiniana2
Luffa_graveolens
Trichosanthes_multiloba2
Trichosanthes_pallida1
Cyclanthera_pedata
Trichosanthes_subvelutina3
Trichosanthes_dioica
Trichosanthes_quinquefolia
Gymnopetalum_scabrum3
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_truncata2
Trichosanthes_kirilowii_var._japonica2
Trichosanthes_sepilokensis
Linnaeosicyos_amara
Trichosanthes_laceribractea2
Trichosanthes_celebica
Sicyos_angulatus
Trichosanthes_villosa1
Trichosanthes_wawrae
Trichosanthes_postarii1
Trichosanthes_mucronata
Trichosanthes_laeoica1
Luffa_operculata
Trichosanthes_montana_subsp._montana
Trichosanthes_kerrii
Lagenaria_siceraria
Trichosanthes_pilosa4
Trichosanthes_quinquangulata2
Trichosanthes_pubera_var._fissisepala2
Ecballium_elaterium
Trichosanthes_kirilowii_var._japonica3
Marah_macrocarpus
Trichosanthes_villosa2
Gymnopetalum_tubiflorum3
Trichosanthes_fissibracteata
Trichosanthes_subvelutina2
Austrobryonia_micrantha
Trichosanthes_beccariana
Trichosanthes_pubera_var._fissisepala1
Trichosanthes_quinquangulata1
Trichosanthes_cucumerina1
Trichosanthes_smilacifolia
Trichosanthes_rosthornii1
Trichosanthes_multiloba1
Gymnopetalum_scabrum2
Trichosanthes_villosa3
Trichosanthes_auriculata
Trichosanthes_lepiniana1
Trichosanthes_adhaerens
Gymnopetalum_tubiflorum2
Trichosanthes_odontosperma1
Trichosanthes_villosa4
Trichosanthes_pubera_var._rubriflos2
Hodgsonia_heteroclita
Trichosanthes_pilosa3
Luffa_acutangula
Bryonia_dioica
Trichosanthes_pilosa1
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
G2015 Macmillan Publishers Limited. All rights reserved
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_imbricatus102
Dacrycarpus_kinabaluensis111
Dacrycarpus_imbricatus106
Dacrycarpus_kinabaluensis913
Dacrycarpus_imbricatus105
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
5 4 3 2 1 0Millions of years ago
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
25 20 15 10 5 0Millions of years ago
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
10 8 6 4 2 0Millions of years ago
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
5 4 3 2 1 0Millions of years ago
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
G2015 Macmillan Publishers Limited. All rights reserved
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
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
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
4 3 2 1 0Millions of years ago
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
50 40 30 20 10 0Millions of years ago
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_albomarginata
Nepenthes_bokorensis
Nepenthes_vieillardii
Nepenthes_andamana
Nepenthes_kongkandana
Nepenthes_singalana
Nepenthes_macfarlanei
Nepenthes_benstonei
Nepenthes_reinwardtiana
Nepenthes_specatabilis
Nepenthes_insignis
Nepenthes_bongso
Nepenthes_gymnamphora
Nepenthes_ventricosa
Nepenthes_alata
Nepenthes_campanulata
Nepenthes_truncata
Nepenthes_peltata
Nepenthes_edwardsiana2
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
25 20 15 10 5 0Millions of years ago
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
7 6 5 4 3 2 1 0Millions of years ago
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
SP13510_Philalanka_spec2_10
SP13492_Philalanka_spec3_50
4 3 2 1 0Millions of years ago
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
G2015 Macmillan Publishers Limited. All rights reserved
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
15 10 5 0Millions of years ago
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
15 10 5 0Millions of years ago
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
10 8 6 4 2 0Millions of years ago
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
6 4 2 0Millions of years ago
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_sp_Bradford_837
Weinmannia_racemosa
Weinmannia_silvicola
Weinmannia_arguta
Weinmannia_sanguisugarum
Weinmannia_stenostachya
Weinmannia_bojeriana
Weinmannia_humbertiana
Weinmannia_minutiflora
Weinmannia_marojejyensis
Weinmannia_madagascariensi
Weinmannia_rutenbergii
Weinmannia_sp_Bradford_655
Weinmannia_clemensiae
Weinmannia_exigua
Weinmannia_richii
Weinmannia_fraxinea
Weinmannia_hooglandii
35 30 25 20 15 10 5 0Millions of years ago
KBW
W
W
W
W
W
W
W
W
W
W
KBW
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
KBW
KBW
W
KBW
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
KBW
W
BioGeoBEARS DEC+J on Weinmannia M0_unconstrainedancstates: global optim, 3 areas max. d=6e−04; e=0; j=0; LnL=−12.42
KBW
W
W
W
W
W
W
W
W
W
W
KBW
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
W
KBW
KBW
W
W
W
W
W
W
WW
W
W
WW
W
W
W
WW
W
W
WW
W
W
W
W
W
W
WW
WW
WW
K
WW
KBW
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
G2015 Macmillan Publishers Limited. All rights reserved
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
G2015 Macmillan Publishers Limited. All rights reserved