Philippine Journal of Science 142: 223-244, Special Issue ISSN 0031 - 7683 Date Received: ?? ???????? 2013

Diversity and Evolution of in the Philippines

Jun Wen1, Limin Lu2, and John K. Boggan1

1 Department of Botany, MRC-166, Smithsonian Institution, PO Box 37012, Washington, DC 20013-7012, U.S.A. 2 State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China

Vitaceae is economically well-known for grapes and ecologically important as major in tropical and temperate forests. This study analyzes the assembly of taxa of Vitaceae in the Philippines in the phylogenetic framework using five chloroplast DNA markers and enumerates the of Vitaceae in the Philippines. The results suggest active floristic exchanges between the Philippines and other parts of Asia and Australia in the and Pliocene with some elements in the Oligocene. The taxonomic study recognizes seven genera and 55 species. Two new species are validly published and four new combinations are made: sinuosa (Merr.) J. Wen & Boggan, comb. nov.; Causonis corniculata (Benth.) J. Wen & L.M. Lu, comb. nov.; Causonis pterita (Merr.) J. Wen & L.M. Lu, comb. nov.; simplicifolia (Merr.) J. Wen & Boggan, comb. nov.; Tetrastigma silvestrei Elmer ex J. Wen & Boggan, sp. nov., and coi J. Wen & Boggan, sp. nov. The last species is named in honor of Leonardo Co.

Key Words: Vitaceae, Philippines, phylogeny, taxonomy, Ampelocissus, Ampelopsis, Causonis, Cayratia, , , , Pterisanthes, , Nothocissus, Tetrastigma, Vitis

INTRODUCTION have suggested the need to redefine several genera. Cayratia Juss., Cyphostemma (Planch.) Alston and Tetrastigma were Vitaceae (the grape family) consists of about 14 genera and supported to form a clade by all these analyses and the 900 species primarily distributed in tropical regions (Wen monophyly of Cyphostemma and Tetrastigma was each 2007a). The family is well-known for grapes (species of Vitis strongly supported. Cayratia was, however, found to be L.) and is ecologically important because many species are paraphyletic with Tetrastigma and Cyphostemma nested major climbers in tropical and temperate forests. The within it (Lu et al. 2013), rendering the need for redefining Tetrastigma (Miq.) Planch. of Vitaceae is also famous in the generic limits of Cayratia. The phylogenetic analyses southeast Asia for being the host of the holoparasitic also supported a clade of the grape genus Vitis and its close , which includes Rafflesia R. Br. ex Gray, the relatives Ampelocissus Planch., Pterisanthes Blume and genus producing the largest flower of the world (Wen 2007a). Nothocissus (Miq.) Latiff from the tropics (hereafter referred Vitaceae has been subjected to recent phylogenetic studies to as the Vitis-Ampelocissus clade, see Ren et al. 2011). (e.g., Soejima & Wen 2006, Rossetto et al. 2007, Wen et al. Pterisanthes was found to be nested within Ampelocissus 2007, Ren et al. 2011, Nie et al. 2012, Trias-Blasi et al. 2012, (Ren et al. 2011). A clade of Ampelopsis and its close Lu et al. 2013, Wen et al. 2013). The phylogenetic analyses relatives (the African Rhoicissus Planch., the Australian Planch. and a small clade of about five species Corresponding Author: [email protected] of Cissus L. from South America) has been resolved as

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the first diverged lineage within Vitaceae, but the genus the Philippines and Asia overall, with a global sampling Ampelopsis is paraphyletic (Nie et al. 2012). scheme (Appendix 1). Sequences of five plastid markers including atpB-rbcL, rps16, trnC-petN, trnH-psbA, and The Malesian region is an important center of diversity of trnL-F were mostly obtained from GenBank based on Vitaceae with nine of the 14 recognized genera and 165 studies from the senior author’s lab (e.g., Soejima & Wen of the 900 species in the family: Ampelocissus (35 spp.), 2006, Nie et al. 2010, 2012, Chen et al. 2011a, b, Ren et Ampelopsis (2 spp.), Cayratia (20 spp.), Cissus (31 spp.), al. 2011, Lu et al. 2012, 2013, Liu et al. 2013), with 52 Nothocissus (6 spp.), Parthenocissus (1 sp.), Pterisanthes new sequences (including nine atpB-rbcL, nine rps16, (20 spp.), Tetrastigma (50 spp.), and Vitis (1 sp. native 15 trnC-petN, 11 trnH-psbA, and eight trnL-F) generated and 1 sp. cultivated). for the current study following the protocols in Ren et al. Vitaceae of the Philippines was treated initially by Blanco (2011) and Lu et al. (2013). The program Sequencher 5.0 (1837, 1845, 1877), who recognized seven species of (Gene Codes Co., Ann Arbor, Michigan, USA) was used to Cissus, belonging to four currently recognized genera evaluate chromatograms for base confirmation and to edit of Vitaceae (Causonis Raf., Cayratia, Cissus and contiguous sequences. Sequences were initially aligned Tetrastigma), and one genus Melicope Forster & Forster f. using MUSCLE 3.8.31 (Edgar 2004), and the alignment of Rutaceae (Hartley 2001; also see Merrill 1905, 1918b). was then adjusted manually in Geneious 6.1.4 (created by Merrill treated Vitaceae in a series of publications (Merrill Biomatters, available from http://www.geneious.com/). 1907, 1912a, b, 1916, 1918a, 1920, 1923). Elmer (1915, 1939) described a few species of the family. Quisumbing Phylogenetic Analyses & Merrill (1924) and Quisumbing (1944) further Phylogenetic analyses were initially conducted for documented Philippine Vitaceae, described new taxa, and individual DNA regions with maximum likelihood made nomenclatural changes in the family. Nevertheless, (ML) method using RAxML 7.2.6 with 1,000 bootstrap the family has not been revised taxonomically and replicates (Stamatakis 2006). Preliminary analyses for systematically for the Philippines in many decades since. each region indicated no significant topological conflicts The objectives in this study are to: (1) discuss the (BS > 70%; Hillis & Bull 1993) among individual evolutionary assembly of Vitaceae in the Philippines based markers. Furthermore, because the chloroplast genome on a broad phylogenetic framework of the grape family; is generally considered as one unit without recombination and (2) enumerate the taxonomic diversity of Vitaceae in in angiosperms (Whitfeld & Bottomley 1983), sequences the Philippines. of the five plastid markers were thus combined and were analysed with maximum parsimony (MP) and Bayesian In memory of Leonardo Co, the senior author Jun Wen felt inference (BI). that it would be most meaningful to provide an updated treatment of Vitaceae in the Philippines and discuss its Maximum parsimony analyses were conducted using evolutionary assembly, in light of recent phylogenetic PAUP*4.0b10 (Swofford 2003) with a heuristic search evidence. Wen first met Leonardo Co in 1991 at the Harvard strategy followed by 100 random-addition-sequence University Herbaria where Wen was a postdoctoral fellow replicates with tree bisection and reconnection (TBR) and Co was visiting. Wen communicated with Leonardo branch swapping. Bootstrap values (BS) were estimated Co on Vitaceae, Leeaceae and Araliaceae treatments for with 1,000 bootstrap replicates (Felsenstein 1985) the Philippines in several different occasions. After the 6th using heuristic searched as described above, but with International Flora Malesiana Symposium in Los Baños branch swapping limited to 10,000,000 rearrangements in 2004, Wen joined the post-symposium excursion led by per replicate due to memory constraints. A partitioned Leonardo to various areas in Luzon. During the trip, Wen Bayesian analysis was conducted using MrBayes 3.2.1 and Co also discussed the taxonomy and identification of (Ronquist & Huelsenbeck 2003, 2012). The data set a number of Vitaceae specimens encountered during the was partitioned into five subsets corresponding to five trip and examined many images of Vitaceae specimens plastid regions with unlinked substitution models. Two they both had. independent runs were simultaneously performed with each run consisting of one cold chain and three heated chains. Each run was conducted with 15,000,000 generations and sampled one tree every 1,500 generations. MATERIALS AND METHODS Analyses were run until the average standard deviation of the split frequencies approached 0.01, indicating that two runs converged onto a stationary distribution. A 50% Taxon Sampling majority-rule consensus tree and the posterior probabilities The study sampled 163 accessions of Vitaceae and its (PP) were calculated after discarding the first 25% trees sister family Leeaceae (Ridsdale 1974), emphasizing as burn-in. The generalized time reversible model (GTR)

224 Philippine Journal of Science Wen et al.: Vitaceae in the Philippines… Vol. 142: Special Issue with a gamma-distributed rate variation was selected as the research. In this study, we excluded Leea from Vitaceae, best-fit models for each partition as well as the combined following Ridsdale (1974) and Wen (2007b). data set as determined by MrModeltest 2.3 (Nylander 2004) under the Akaike Information Criterion (AIC).

Divergence Time Estimation RESULTS We estimated the divergence time of clades using The parsimony analysis generated more than 100,000 most the combined 5-marker data set with BEAST 1.7.4 parsimonious trees (MPTs) with a length of 4,025 steps, a (Drummond & Rambaut 2007). The strict molecular consistency index (CI) of 0.65, and a retention index (RI) clock model was rejected from our dataset based on a of 0.89, with gaps treated as missing data. The BI and likelihood ratio test, and we thus estimated divergence MP analyses produced a similar topology and the strict time using a Bayesian relaxed clock model with rate consensus tree of the combined data set with the Bayesian variation across branches uncorrelated and lognormally posterior probabilities, and bootstrap support values from distributed. The data set was partitioned into five subsets the MP analyses, was presented in Figures 1, 2A & 2B. corresponding to five plastid regions with unlinked substitution models using BEAUti 1.7.4. To satisfy the The divergence time estimates for Vitaceae, with the temporal and topological constraints, we specified a clades from the Philippines marked, are presented in starting tree derived from the majority-rule consensus tree Figure 3. of the BI analyses. The BI runs relied on the GTR + G DISCUSSION model (with four rate categories) and a Yule process tree prior. Two separate Markov chain Monte Carlo (MCMC) Evolutionary Assembly of Vitaceae in the Philippines analyses were run, each with 100,000,000 generations and Our phylogenetic analyses supports six major clades sampling every 10,000 generations. The program Tracer with Vitaceae using Leea of Leeaceae as the outgroup 1.5 (Rambaut & Drummond 2007) was used to check that (Figure 1): (I) Ampelopsis-Rhoicissus-Clematicissus- effective sample size (ESS) for all relevant parameters Cissus striata complex, (II) a small Australasian Cissus were well above 200 and that stationarity probably had clade (Cissus II), (III) the Ampelocissus-Pterisanthes- been reached. The maximum clade credibility tree was Nothocissus-Vitis clade, (IV) the Parthenocissus-Yua then built using TreeAnnotator 1.7.4 (beast.bio.ed.ac.uk/ clade, (V) the core Cissus clade, and (VI) the Cayratia- TreeAnnotator) with the initial 25% of trees discarded as Causonis-Cyphostemma-Tetrastigma clade. Ampelopsis, burn-in. Final tree was edited in FigTree 1.3.1 (Rambaut Ampelocissus and Cayratia are each supported to be 2009) to view dates of all nodes and variation of rate of paraphyletic (Figures 2A & 2B), as reported in our earlier substitutions on branches. studies (Ren et al. 2011; Nie et al. 2012; Lu et al. 2013), Vitaceae has well preserved seed records in the suggesting the need for generic re-circumscription. Tertiary (e.g., Reid & Chandler 1933, Kirchheimer 1939, Taxa of Philippine Vitaceae are found scattered throughout Miki 1956, Tiffney & Barghoorn 1976). Ampelocissus the Vitaceae phylogeny (Figures 1, 2A, 2B). The dated parvisemina Chen & Manchester from the Beicegala phylogeny (Figure 3) suggests very active exchanges Creek locality in North Dakota in U.S.A. (56.8-62.0 Ma) of Philippine taxa with other parts of Asia (see Figures was well preserved and can be accurately assigned to 2A, 2B & 3), especially during Miocene and Pliocene. genus Ampelocissus (Chen and Manchester 2007). We This is consistent with the discussion by Merrill (1926) used Ampelocissus parvisemina to constrain the crown concerning the evolution of the Philippine flora based of the Ampelocissus-Nothocissus-Pterisanthes-Vitis clade on floristic and geologic evidence. Tetrastigma has with a lognormal distribution (mean: 58.5 Ma; log (stdev): the highest endemism in the Philippines, with about 0.03; offset: 0 Ma; mean in real space) to include the 15 endemic species. Species of the genus exhibited 95% HPD interval of the fossil date (56.8-62.0 Ma). We divergences with other Asian species at different times, also constrained the stem of Vitaceae as 90.7 ± 1.0 with ranging from Oligocene to Pliocene (Figure 3). The a normal prior distribution based on the estimated age of Philippines are thus important for the evolutionary 90.65-90.82 Ma by Magallón & Castillo (2009). diversification of Tetrastigma. Few endemic taxa were included in sampling for other genera. Nevertheless, Taxonomic Enumeration the Philippine Vitaceae shows close affinities with taxa Taxonomic enumeration of Vitaceae in the Philippines of other parts of Asia as well as Australia (Figures 2A was based on herbarium studies of the specimens kept & 2B). Future studies with more sampling of endemic in the following herbaria: A, BO, F, K, KEP, L, LAE, Philippine taxa are needed to analyze the detailed patterns LBC, PNH, SING and US, as well as extensive literature of biogeographic assembly of Philippine Vitaceae.

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Figure 1. Strict consensus tree of Vitaceae using maximum parsimony analysis of the combined plastid data set. Posterior probabilities are displayed below the branches and parsimony bootstrap values are shown above branches with only major clades labeled. All taxa from the Philippines were marked with a star. Specimen details are listed in Appendix 1.

226 Philippine Journal of Science Wen et al.: Vitaceae in the Philippines… Vol. 142: Special Issue

A

B

Figure 2. Strict consensus tree of Vitaceae using maximum parsimony analysis of the combined plastid data set. Posterior probabilities are displayed below the branches and parsimony bootstrap values are shown above branches with geographic origins of the taxa and support values of all clades labeled. A. The Cayratia-Cyphostemma- Tetrastigma clade, and the Cissus clade; B. the remaining clades of Vitaceae and the outgroup taxa.

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Figure 3. Maximum clade credibility tree of Vitaceae inferred from combined atpB-rbcL, rps16, trnC-petN, trnH-psbA, and trnLF data using BEAST. Gray bars indicate 95% highest posterior density intervals. Clade constraints are shown with black stars. Nodes of Philippine Vitaceae were indicated as 1 to 29.

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Overall Taxonomy of Vitaceae in the Philippines 1. Ampelopsis Michx. Seven genera and 55 species are distributed in the Ampelopsis Michx., Fl. Bor.-Amer. 1: 159. 1803. Philippines, including two cultivated species. They are Deciduous woody climbers; hermaphrodite; tendrils usually Ampelopsis (1 sp.), Ampelocissus (9 spp., including 1 sp. few and scattered, bifurcate, without adhesive discs. Leaves of Pterisanthes), Vitis (1 native sp. and 1 cultivated sp.), membranaceous, simple to variously lobed, or trifoliate. Cissus (9 native spp. and 1 cultivated sp.), Cayratia (5 spp.), Inflorescence a loose thyrse, few to many flowered, leaf- Causonis (4 spp.), and Tetrastigma (24 spp.). Pterisanthes opposite, often repeatedly bifurcate and appearing loosely has been recently shown to be nested within the Asian corymbose, rarely elogate and racemose, the branches Ampelocissus (Wen et al. 2007, Ren et al. 2011) and is ending with 3-flowered dichasia, sometimes showing herein treated as a synonym of the latter. Causonis is herein transitions to tendrils, with the peduncle or a branch recognized as a segregate genus of Cayratia based on recent coiled, functioning as a tendril for climbing. Flowers phylogenetic evidence to maintain the monophyly of each 5-merous; calyx small, saucer-like; corolla spreading at genus (Lu et al. 2013, also see Figure 2A). anthesis; stamens 5, erect; floral disc cupular, lower part adnate to the ovary, slightly lobed; ovary 2-locular, style Key to the genera of Vitaceae in the Philippines slender, stigma small, simple. a berry, subglobose to obovoid, blue, black or green, 1-4-seeded; seeds obovate to 1. Stigma of the bisexual flowers 4-lobed or 4-parted, broadly so, convex on abaxial side, angular on adaxial side, enlarged, wider than the tip of style, styles very short . . chalazal knot spatulate, raphe narrowly linear, 2 adaxial ...... 7. Tetrastigma grooves oblanceolate, present in the lower half of the seed; 1. Stigma not 4-parted or 4-lobed, usually never wider endosperm T-shaped in cross section. 2n = 40. than the end of style ...... 2 In this paper, Ampelopsis is more narrowly defined 2. Petals united into a cap-like structure (calyptra) that based on the phylogenetic results of Nie et al. (2012) drops off as a unit at anthesis; bark on older twigs by excluding members of section Leeaceifoliae (with usually loose, peeling off in stripes ...... 3. Vitis pinnately to bipinnately compound leaves) to ensure the monophyly of Ampelopsis. The genus was shown 2. Petals not forming a calyptra; bark intact, not peeling to be paraphyletic with the African Rhoicissus, the off in stripes ...... 3 Australian Clematicissus and a small clade of Cissus 3. Inflorescence with a tendril at the base or the axis, from South America nested within it. The genus consists sometimes axis becoming fleshy and foliaceous, but of approximately 12 species with a disjunct distribution still with tendrils on the peduncle (“Pterisanthes” in temperate to subtropical Asia (ca. 10 spp.) and North group) ...... 2. Ampelocissus and Central America (two spp. including one in eastern North America and one in Mexico and Guatemala). One 3. Inflorescence without a tendril at the base, tendrils species is distributed in the Philippines. normally leaf-opposite ...... 4 (1) Ampelopsis glandulosa (Wall.) Momiyama var. 4. Petals 4 ...... 5 hancei (Planch.) Momiyama 1977 J. Jap. Bot. 52(1): 30. 4. Petals 5 ...... 1. Ampelopsis (B) Ampelopsis heterophylla var. hancei Planch. 5. Inflorescences umbellate cymes, usually leaf- 1887 Monogr. Phan. 5: 457. opposed; floral disc cupular even when 4-lobed, (S) Ampelopsis brevipedunculata var. hancei (Planch.) raised above the ovary; floral disc copular and raised Rehder above the ovary; 1-seeded ...... 4. Cissus 1922 J. Arnold Arbor. 22: 177. Distribution: China (Fujian, Guangdong, Guangxi, 5. Inflorescence corymbose cymes, usually axillary Guizhou, Henan, Hunan, Jiangsu, Jiangxi, Shandong, or sometimes seemingly terminal; floral disc not Sichuan, Taiwan, Yunnan), Vietnam, Japan, Philippines cupular; fruits usually 2-4-seeded ...... 6 (the Batanes Islands, Luzon, Mindanao, Mindoro, 6. Seeds with a membrane covering the ventral infolds Palawan) ...... 5. Cayratia 2. Ampelocissus Planch. 6. Seeds without a membrane covering the ventral Ampelocissus Planch., Vigne Amér. Vitic. Eur. 8: 371. infolds ...... 6. Causonis 1884. nom. cons. Botria Lour., Fl. Cochinch.: 96. 1790. Pterisanthes Blume, Bijdr. 192. 1825. Taxonomic Enumeration of Vitaceae in the Philippines Embamma Griffith, Notul. 4: 694. 1854. Basionyms are listed as (B); and synonyms are in (S).

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Vitis L. sect. Nothocissus Miq., Ann. Mus. Lugd. Batav. 1916 Philipp. J. Sci., Bot. 11: 125 1: 73. 1863. (B) Cissus ochracea Teijsm. & Binn. Ampelocissus Planch. sect. Nothocissus (Miq.) Planch., 1864 Tijdschr. Nederl. Ind. 27: 35. in De Candolle, Monogr. Phan. 5: 369. 1887. Nothocissus (Miq.) Latiff, Feder. Mus. J. 27: 70. 1982. (6a) Ampelocissus ochracea (Teijsm. & Binn.) Merr. 1916 Philipp. J. Sci., Bot. 11: 125. Woody climbers; polygamo-monoecious; tendrils borne Distribution: Sumatra, Sulawesi, Borneo (Sabah, on the inflorescences. Leaves simple, entire or lobed, Brunei), Philippines (Basilan, Culion, Mindanao). to digitately and pedately compound with 3-9 leaflets; stipules deltate, inconspicuous, caducous. Inflorescence (6b) Ampelocissus ochracea var. trilobata Merr. leaf-opposed, a thyrse, or paniculate spikes, with the 1916 Philipp. J. Sci., Bot. 11: 125. peduncle tendril-bearing; sometimes (in the “Pterisanthes” Distribution: Philippines (Catanduanes, Luzon, group) a leaf-opposed panicle of lamella, fleshy and Mindanao, Mindoro, Polillo). foliaceous, with tendrils on the peduncle. Flowers 5-, or (7) Ampelocissus pauciflora Merr. occasionally 4-merous (e.g., in the “Nothocissus” group); 1916 Philipp. J. Sci., Bot. 11: 126. calyx copular, or saucer-shaped, entire or with 5 obscure Distribution: Philippines (Calusa, Coron, Cuyo, Golo, teeth; corolla with 5 or sometimes 4 petals, oblong to Luzon, Palawan, Panay). ovate-oblong, spreading or recurved at anthesis; stamens inserted without the disc, filaments slender; floral disc (8) Ampelocissus sinuosa (Merr.) J. Wen & Boggan, annular, erect, often vertically 5-10-furrowed; ovary more comb. nov. (Figure 4) or less embraced in the disk, styles short, conical, often Distribution: Philippines (Mindanao). 10 furrowed, stigma small in staminate flowers, discoid in (B) Pterisanthes sinuosa Merr. hermaphrodite flowers. Fruit a berry, 1-4-seeded; seeds 1907 Philipp. J. Sci., Bot. 2: 423. oblong to obovoid, abaxial surface convex, adaxial surface (9) Ampelocissus trichoclada Quisumb. & Merr. 2-furrowed, with a broad raphe, often crenately cleft at the 1928 Philipp. J. Sci. 37: 165. margin; endosperm T-shaped in cross section. 2n = 40, 80. Distribution: Philippines (Mindanao). About 100 species mostly in Africa, tropical Asia, and 3. Vitis L. Australia with only four species in the New World (Central Vitis L., Sp. Pl. 2: 230. 1753. America and the Caribbean). Nine species occur in the Philippines. Recent phylogenetic evidence suggests that the Deciduous woody climbers, polygamodioecious, usually type species of Nothocissus and species of Pterisanthes are with shred-like bark on old stems; lenticels absent or nested within Ampelocissus (Wen et al. 2007, Ren et al. 2011). inconspicuous (subgenus Vitis) or prominent (subgenus Muscadinia); pith brown, interrupted by diaphragms (1) Ampelocissus botryostachys Planch. within the nodes (subgenus Vitis) or continuous through 1887 Monogr. Phan. 5: 413. nodes (subgenus Muscadinia); tendrils 2-3-forked Distribution: Philippines (Luzon, Panay). (subgenus Vitis), or simple (subgenus Muscadinia), (2) Ampelocissus dolichobotrys Quisumb. & Merr. present opposite only two consecutive nodes or less 1928 Philipp. J. Sci. 37: 164. commonly at three to many consecutive nodes, without Distribution: Philippines (Luzon). adhesive discs. Leaves simple, often lobed, cordate to truncate at base, dentate to serrate at margin, palmately (3) Ampelocissus madulidii Latiff veined; stipules caducous. Inflorescence a panicle, present 1988 Blumea 33(2): 505. opposite only two consecutive nodes or at three to many Distribution: Philippines (Samar). consecutive nodes, sometimes with a tendril at the apex (4) Ampelocissus martini Planch. of the peduncle. Flowers 5-merous, morphologically 1884 Vigne Amer. Vitic. Eur. 8: 376. bisexual and unisexual, but functionally unisexual; Distribution: Thailand, Vietnam, Cambodia, Laos, calyx minute with 5 teeth to entire; corolla of 5 or rarely Peninsular , Borneo, Philippines (Apulit, 3-9 petals, forming a calyptra at anthesis; stamens Banton, Culion, Coron, Cuyo, Guimaras, Luzon, usually 5, erect in male and bisexual flowers, reflexed Palawan, Semirara). or occasionally absent in female flowers; floral disc of 5 glands alternating with stamens; gynoecium rudimentary (5) Ampelocissus multifoliola Merr. in male flowers; in female flowers, style conical and short, 1916 Philipp. J. Sc., Bot. 11: 127. and stigma small, capitate. Fruit a berry, pulpy, globose, Distribution: Philippines (Catanduanes, Luzon). purple to black, rarely whtish or greenish, often glaucous, 1-4-seeded; seeds obovoid to pyriform, the adaxial surface (6) Ampelocissus ochracea (Teijsm. & Binn.) Merr. with two longitudinal grooves on either side of the raphe,

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1905 Bull. Soc. Agric. Sci. Arts Sarthe 40: 36. (S) Vitis flexuosa var. parvifolia (Roxb.) Gagnep. 1911 Pl. Wilson. 1: 103. (S) Vitis flexuosa var. wallichii (DC.) F.S. Wang 2000 J. Trop. Subtrop. Bot. 8(1): 5. Distribution: , Nepal, Pakistan, China, Korea, Japan, Laos, Thailand, Vietnam, Java, Philippines (Luzon). (2) Vitis vinifera L. 1753 Sp. Pl. 202, 293. (S) Cissus vinifera (L.) Kuntze 1881 Um die Erde: 501. Distribution: Native to western Asia, cultivated in the Philippines as grape . 4. Cissus L. Cissus L., Sp. Pl. 1: 117. 1753. Spondylantha Presl, Reliq. Haenk. 2: 35. 1831. Gonoloma Raf., Sylva Tellur.: 86. 1838. Irsiola P.Browne ex Raf., Sylva Tellur.: 86. 1838. Kemoxis Raf., Sylva Tellur.: 86. 1838. Adenopetalum Turczaninow, Bull. Soc. Imp. Nat. Moscow 31: 417. 1858. Pterocissus Urban & Ekman, Arkiv Bot. Bd. 20A (5): 20, Taf. 1. 1926. Woody or herbaceous climbers or scrambling lianas, or sometimes erect shrubs, hermaphroditic to polygamo- Figure 4. Ampelocissus sinuosa (Merr.) J. Wen & Boggan (M.S. Clemens monoecious; stems terete, winged or striated, often 647, Philippines, Mindanao, Camp Keithley, Lake Lanao, Jul 1906, isotype, US, barcode 00094559). A. Overall specimen; succulent; sometimes tuberous roots present; tendrils B. Portion of enlarged inflorescence. leaf-opposite, ramified or unbranched, occasionally with adhesive discs at the tip, especially when young, each branch subtended by a bract, occasionally absent in erect species. Leaves simple, trifoliate, or palmately compound the abaxial surface with a round to elliptic chalazal with 3-5 (-7) leaflets, to pinnately and bipinnately knot, extending to ca. 1/3 of the seed length from base; compound; stipules caduceus. Inflorescence an umbellate endosperm M-shaped. 2n = 38, 40. cyme or a compound umbellate cyme, leaf-opposite. About 65 species mostly in the temperate regions of the Flowers 4-merous; calyx cupular, minute; corolla valvate northern hemisphere. Only one widespread species V. in aestivation, petal lobes adhering by interlocked flexuosa is native to the Philippines. Vitis vinifera is epidermal cells in bud, reflexed at anthesis, deltate in cultivated. shape; stamens opposite the petals; floral disc adnate to the base of the ovary, conspicuous, cupular to 4-lobed; (1) Vitis flexuosa Thunb. styles conical or cylindrical, stigma entire. Fruit a berry, 1794 Trans. Linn. Soc. London 2: 103, 332. 1-(rarely to 4) seeded, inedible; seeds ovoid, obtusely (S) Vitis wallichii DC. 3-cornered, to pyriform, with an encircling raphe, smooth 1824 Prodr. 1: 634. or faceted or pitted on either side, the testa crustaceous; (S) Vitis purani Buch.-Ham. ex D. Don endosperm ruminate, with 3 vertical lobes, M-shaped in 1825 Prod. Fl. Nep. 188. cross section; cotyledons reniform, sometimes 3, radicle (S) Vitis sylvestris Blume large. 2n = 24, 26, 28, 32, ca. 36, 40, ca. 45, 48, 50, ca. 1825 Bijdr. Fl. Nederl. Ind. 194. 85, ca. 95. (S) Vitis truncata Blume 1825 Bijdr. Fl. Nederl. Ind. 195. About 350 species widely distributed in all tropical regions (S) Vitis vitiginea var. truncata (Blume) Kuntze with a few extending into the temperate zone, with ten 1891 Revis. Gen. Pl. 1: 139. species distributed in the Philippines including one species (S) Vitis cavaleriei Lévl. commonly cultivated. The genus is the largest in Vitaceae

231 Philippine Journal of Science Wen et al.: Vitaceae in the Philippines… Vol. 142: Special Issue and has been shown to be polyphyletic (Rossetto et al. 1825 Bijdr. Fl. Nederl. Ind. 1: 181. 2002). But with about ten species excluded, the core Distribution: Bangladesh, India, Sikkim, Nepal, China Cissus is monophyletic (Liu et al. 2013). (Sichuan, Yunnan), Myanmar, Thailand, Vietnam, Laos, Cambodia, Malaysia, Indonesia (Java), Borneo, (1) Cissus adnata Roxb. Philippines (Luzon to Palawan). Widely cultivated and 1820 Fl. Ind. 1: 423. commonly identified as Cissus discolor. (S) Vitis adnata (Roxb.) Wall. 1834 Prodr. Fl. Ind. Orient. 1: 126. (6) Cissus luzoniensis (Merr.) C.L. Li 1996 Chinese J. Appl. Environ. Biol. 2(1): 49. Distribution: Sri Lanka, Bangladesh, India, Sikkim, Nepal, (B) Cissus repens var. luzoniensis Merrill China (Yunnan), Myanmar, Cambodia, Laos, Vietnam, 1916 Philipp. J. Sci., Bot. 9(3): 131. Thailand, Malaysia, Indonesia (Java), Borneo, Brunei, Distribution: China (Hainan, Yunnan), Philippines New , Australia (Western Australia, Northern (Luzon). Territory, Queensland), Philippines (Luzon to Mindanao). (7) Cissus nodosa Blume (2) Cissus aristata Blume 1825 Bijdr. Fl. Nederl. Ind. 182. 1825 Bijdr. Fl. Nederl. Ind. 183. (S) Vitis nodosa (Blume) Miq (S) Cissus simplex Blanco 1863 Ann. Mus. Bot. Lugd.-Bat. 1: 87. 1837 Fl. Filip. 72. Distribution: India (Andaman Islands), Malaysia (S) Cissus pyrrhodasys Miq. (peninsular), Singapore, Sumatra, Java, Borneo 1860 Fl. Ind. Bat. Suppl. 1: 517. (Brunei, Sabah), Philippines (Luzon, Mindanao, (S) Cissus assamica var. pilosissima Gagnep. Palawan). 1911 Notul. Syst. (Paris) 1: 353. (8) Cissus oblongifolia Merr. (S) Vitis pyrrhodasys (Miq.) Ridl. 1916 Philipp. J. Sci., Bot. 11: 129. 1917 J. As. Soc. Straits, 75: 23. Distribution: Philippines (Catanduanes, Luzon, (S) Vitis simplex (Blanco) Burkill Palawan, Panay). 1935 Kew Bull. 1935: 319. (9) Cissus quadrangularis L. Distribution: Bangladesh, Sri Lanka, India, Sikkim, 1767 Syst. Nat., ed. 12, 2: 124. Nicobar Island, China (Hainan, Yunnan), Myanmar, Distribution: tropical Africa to Asia, commonly Indo-China, Thailand, Malaysia, Borneo (Sabah, Brunei), cultivated; Philippines (Cebu, Luzon, Negros, Indonesia (Sumatra, Java), Papua New Guinea, Australia Siquijor). (Queensland), Philippines (Busuanga, Cuyo, Luzon, Mindanao, Panay). (10) Cissus repens Lam. 1783 Encycl. Meth. 1: 31. (3) Cissus assamica (Laws.) Craib Distribution: India, Sikkim, Bhutan, Nepal, China 1911 Kew Bull. 1911: 31. (Guangdong, Guangxi, Guizhou, Taiwan, Yunnan), (B) Vitis assamica Laws. Myanmar, Thailand, Vietnam, Cambodia, Laos, 1875 Fl. Brit. Ind. 1: 648. Malaysia (peninsular), Singapore, Indonesia (Java, (S) Cissus ambigua Elmer ex Merr., nom. nud. pro syn. Sumatra, Borneo, Sulawesi), New Guinea, Christmas 1923 Enum. Philipp. Fl. Pl. 3: 6. I., Borneo (Sabah), Australia (Queensland), Philippines Distribution: Bangladesh, India, Nepal, Bhutan, China (Luzon, Mindanao, Negros, Palawan, Panay, Ticao). (Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, 5. Cayratia Juss. Jiangxi, Sichuan, Taiwan, Xizang, Yunnan), Myanmar, Cayratia Juss. in Dict. Hist. Nat. Sci. 10: 103. 1818, nom. cons. Cambodia, Thailand, Vietnam, Malay (peninsular), Cissus sect. Cayratia (Juss.) Planch., in A. & C. DC. Borneo, Philippines (Luzon, Panay). Monogr. Phan. 5: 47. 1887. (4) Cissus hastata Miq. Columella Lour., Fl. Cochinch. 85. 1790. nom. rej. 1860 Fl. Ind. Bat. Suppl. 1: 517. Climbing shrubs and herbs; hermaphrodite; sometimes Distribution: India, Myanmar, Thailand, Vietnam, tuberous roots present; tendrils oppose to leaves, Malaysia (peninsular), Singapore, Java, Borneo usually 2-3 forked, each branch subtended by a bract. (Sabah, Brunei), New Guinea, Australia (Queensland), Leaves alternate, pedately or trifoliately compound Philippines (Luzon). with 3-7 leaflets; stipules 2, caduceus; leaflets serrate (5) Cissus javana DC. at margin. Inflorescence a dichotomous cyme, axillary, 1824 Prodr. 1: 628. or pseudoaxillary by the abortion of lateral axis, or (S) Cissus discolor Blume sometimes opposite to leaves. Flowers 4-merous; calyx

232 Philippine Journal of Science Wen et al.: Vitaceae in the Philippines… Vol. 142: Special Issue cup-shaped; corolla valvate in aestivation, cohering in bud by interlocked epidermal cells; stamens inserted on the receptacle at the base of the floral disc, opposite to petals, filaments erect, anthers introrse; disc adnate to and entirely surrounding the ovary, 4-lobed; style conical, stigma minute. Fruit a 2-4-seeded berry; seeds hemispheric pyriform to oblong, obcordate, smooth or angular, convex on the back, with 1-2 ventral cavities covered by a membrane; endosperm ruminate, transverse section U-shaped. 2n = 24, 40, 60, 80. About 35 species in tropical and subtropical Asia, Australia, and the Pacific Islands, after separating Causonis from Cayratia based on phylogenetic evidence (Lu et al. 2013; also see Galet 1967, Jackes 1987). (1) Cayratia apoensis (Elmer) Quisumb. 1944 Philipp. J. Sci. 76: 45, 201. (B) Cissus apoensis Elmer 1915 Leafl. Philipp. Bot. 8: 2880. (S) Columella apoensis (Elmer) Merr. 1923 Enum. Philipp. Fl. Pl. 3: 8. Distribution: Philippines (Mindanao). (2) Cayratia coi J. Wen & Boggan, sp. nov. (Figure 5) (S) Columella irosinensis Elmer, nom. inval. 1944 Leafl. Philipp. Bot. 10 (136): 3799. (S) Cayratia irosinensis (Elmer) Galet, nom. illeg. 1967 Recherch. Meth. Identif. & Classif. Figure 5. Cayratia coi J. Wen & Boggan. (D.A. Madulid 6784, Vitac. 2: 372. Philippines, Luzon, Mt. Isarog, Camarines Sur, 400 m, 29 Apr 1987, holotype, US, barcode 01089428). A. Overall Type: Philippines: Luzon, Mt. Isarog, Camarines Sur, 400 specimen; B. Ventral side of seed showing membrane m, 29 Apr 1987, fr, D.A. Madulid 6784 (holotype: US; covering infolds; C. Seed showing chalaza. isotype: F). Additional specimens examined: Philippines: Luzon, Province of Sorsogon, Irosin (Mt. Bulusan), in thickets along streams at low altitudes, Nov 1915, fl & Galet based on an invalid name in the thesis. We herein fr, A.D.E. Elmer 15526 (S, US, 2 sheets). describe it as a new species and name it in honor of Woody climber. Leaves pedate with 5 leaflets to trifoliate; Leonardo Co. petioles 5-8 cm long, with short hairs; leaflet blades The description by Elmer (1939) indicated that the species coriaceous, sparsely pubescent on the upper surface, is 5-foliolate. The holotype of Cayratia coi at US bears puberulent on veins and veinlets on the lower surface, pedate leaves with five leaflets. One specimen of Elmer elliptic to ovate, 9-11.5 × 4-7 cm, abruptly acute at apex, 15526 at US bears pedate leaves with 5 leaflets; and rounded to acute at base, often oblique on lateral leaflets, two sheets of Elmer 15526 at S and US, however, are serrate at margin, lateral veins 5-8 on each side; petiolules 3-foliolate. All three sheets have consistent leaf shape, 1.5-2 cm on lateral leaflets, 2.5-4.5 cm on terminal leaflets, inflorescence and floral morphology. Elmer (1939) puberulent. Inflorescence a large compound cyme, distinguished the species from Cayratia geniculata by axillary, 10-16 cm × 10-12 cm. Flowers 4-merous; calyx the number of leaflets. Furthermore, Cayratia geniculata short, 0.2-0.3 × 0.3-0.4 mm, copular; petals 1.2-1.5 × 0.6- usually has finer teeth on the margin and has denser 0.7 mm; style 1-1.5 mm long at anthesis; floral disc thick. pubescence on lower leaflet surfaces. Fruits a 2-4-seeded berry, fleshy, globose, 7-10 × 8-10.5 mm; seeds o×void, 6-6.5 × 4-4.5 mm, with a membrane (3) Cayratia geniculata (Blume) Gagnep. covering the ventral infolds, chalaza linear. 1911 Notul. Syst. (Paris) 1: 345. (B) Cissus geniculata Blume Elmer (1939) published Columella irosinensis Elmer 1825 Bijdr. Fl. Nederl. Ind. 184. without providing a Latin description. Galet (1967) (S) Cissus hirtella Blume published the combination Cayratia irosinensis (Elmer) 1825 Bijdr. Fl. Nederl. Ind. 185.

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(S) Cissus rhodocarpa Blume alternate, pedately compound or trifoliate with 3-7 1825 Bijdr. Fl. Nederl. Ind. 185. leaflets; stipules 2, caduceus; leaflets serrate at margin. (S) Cissus rubescens Blanco Inflorescence a large compound cyme, axillary, or 1837 Fl. Filip. 71. pseudoaxillary, sometimes opposite to leaves. Flowers (S) Cissus alata Blanco non Jacq. 4-merous; calyx cup-shaped; corolla valvate in aestivation; 1845 Fl. Filip. ed. 2: 51. stamens inserted on the receptacle at the base of the floral (S) Vitis geniculata (Blume) Miq. disc, opposite to petals, filaments erect, anthers introrse; 1863 Ann. Mus. Bot. Lugd.-Bat. 1: 81. disc adnate to and entirely surrounding the ovary, 4-lobed; (S) Cayratia rhodocarpa (Blume) Gagnep. style conical, stigma minute and undivided. Fruit a 1911 Notul. Syst. (Paris) 1: 346. 2-4-seeded berry; seeds hemispheric pyriform to oblong- (S) Columella geniculata (Blume) Merr. globose, smooth or somewhat angular, convex on the back, 1916 Philipp. J. Sci., Bot. 11: 132. with 1-2 ventral cavities without a membrane covering the (S) Columella geniculata var. sarcocarpa Merr. ventral infolds (cavities); endosperm ruminate, transverse 1916 Philipp. J. Sci., Bot. 11: 133. section T-shaped. 2n = 30, 40, 60, 80, 120. (S) Cayratia geniculata var. sarcocarpa (Merr.) Quisumb. 1944 Philipp. J. Sci. 76: 46. About 25 species in tropical, subtropical and temperate Distribution: Bhutan, India, Sikkim, China Asia to Australia, the Pacific Islands, and Africa. Four (Guangdong, Guangxi, Hainan, Xizang, Yunnan), species (one endemic) occur in the Philippines. The genus Laos, Vietnam, Malaysia (Sabah), Indonesia (Sumatra, is separated from Cayratia based on recent phylogenetic Java, Borneo), Brunei, Philippines (widespread). evidence (Wen et al. 2007, Lu et al. 2013). It corresponds to the Asian members of Cayratia sect. Discypharia Süess. (4) Cayratia mollissima (Wall.) Gagnep. (Süessenguth 1953, Latiff 1981). 1911 Notul. Syst. (Paris) 1: 345. (B) Vitis mollissima Wall. (1) Causonis corniculata (Benth.) J. Wen & L.M. Lu, comb. nov. 1824 Fl. Ind., ed. Carey & Wall., 2: 482. (B) Vitis corniculata Benth. (S) Cissus mollissima (Wall.) Planch. 1861 Fl. Hongk. 54. 1887 Monogr. Phan. 5: 575. (S) Cissus corniculata (Benth.) Planch. (S) Columella mollissima (Wall.) Merr. 1887 Monogr. Phan. 5: 563. 1923 Enum. Philipp. Fl. Pl. 3: 8. (S) Cayratia corniculata (Benth.) Gagnep. Distribution: India, Myanmar, Thailand, Cambodia, 1911 Notul. Syst. (Paris) 1: 347. Vietnam, Peninsular Malaysia, Sumatra, Java, Borneo, (S) Columella corniculata (Benth.) Merr. Sabah, Brunei, Philippines (Luzon, Mindanao, Panay). 1916 Philipp. J. Sci., Bot. 11: 133. Distribution: China (Fujian, Guangdong, Hainan, (5) Cayratia pedata (Lam.) Juss. ex Gagnep. Hong Kong, Taiwan), Vietnam, Malaysia, Philippines 1910 Notul. Syst. (Paris) 1: 346. (Luzon, Mindanao, Mindoro, Panay). (B) Cissus pedata Lam. 1783 Encycl. 1: 31. (2) Causonis japonica (Thunb.) Raf. (S) Cissus heptaphylla Retz. 1830 Med. Fl. 2: 122. 1788 Obs. Bot. 5: 22. (B) Vitis japonica Thunb. (S) Columella pedata (Lam.) Lour. 1784 Fl. Jap. 104. 1790 Fl. Cochinch. 1: 86. (S) Cissus japonica (Thunb.) Willd. (S) Cissus cochinchinensis Spreng. 1798 Sp. Pl. ed. 4, 1(2): 659. 1824 Syst. Veg. ed. 16, 1: 450 (S) Cissus leucocarpa Blume (S) Vitis pedata (Lam.) Wall. ex Wight 1825 Bijdr. Fl. Nederl. Ind. 189. 1833 Cat. Ind. Pl. 26. (S) Vitis tenuifolia Wight & Arn. Distribution: Sri Lanka, India, Andaman & Nicobar 1834 Prodr. Fl. Ind. Or. 1: 129. Islands, China (Guangxi, Yunnan), Bangladesh, (S) Causonia japonica (Thunb.) Raf. Myanmar, Thailand, Vietnam, Cambodia, Malaysia, 1838 Sylva Tellur. 87. Indonesia (Java), Philippines (Luzon, Mindanao). (S) Cissus tenuifolia Heyne ex Planch. 1887 Monogr. Phan. 5: 563. 6. Causonis Raf. (S) (Thunb.) Gagnep. Causonis Raf., Med. Fl. 2: 122. 1830. 1911 Notul. Syst. (Paris) 1: 349. (S) Columella tenuifolia (Heyne ex Planch.) Gagnep. Climbing herbs or shrubs; hermaphrodite; sometimes 1911 Notul. Syst. (Paris) 1: 348. tuberous roots present; tendrils oppose to leaves, usually (S) Columella tenuifolia (Heyne ex Planch.) Merr., comb. superfl. 2—3 forked, each branch subtended by a bract. Leaves 1916 Philipp. J. Sci., Bot. 11: 134.

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(S) Columella japonica (Thunb.) Merr. (S) Cissus carnosa Lam. 1918 Philipp. J. Sci., Bot. 13: 145. 1783 Encycl. 1: 31. (S) Cayratia tenuifolia (Wight & Arn.) Gagnep. (S) Cissus obtusifolia Lam. 1911 Notul. Syst. (Paris) 1: 348. 1783 Encycl. 1: 31. Distribution: India, Sikkim, Andaman & Nicobar (S) Cissus crenata Vahl Islands, China, Korea, Japan, Myanmar, Thailand, 1794 Symb. Bot. 3: 19. Laos, Cambodia, Vietnam, Peninsular Malaysia, (S) Vitis carnosa (Lam.) Wall. ex Wight & Arn. Borneo (Sabah), Indonesia (Java), New Guinea, 1834 Prodr. 1: 127. Solomon Islands, Philippines, Australia (Queensland), (S) Cissus acida Blanco Philippines (Luzon). 1837 Fl. Filip. 69. (S) Cissus psoralifolia (F. Muell.) Planch. (3) Causonis pterita (Merr.) J. Wen & L.M. Lu, comb. 1887 Monogr. Phan. 5: 567. nov. (Figure 6) (S) Cissus trifolia (L.) K.Schum. (B) Columella pterita Merr. 1889 Fl. Kais. Wilh. Land 71. 1916 Philipp. J. Sc., Bot. 11: 135. (S) Columella trifolia (L.) Merr. (S) Cayratia pterita (Merr.) Quisumb. 1916 Philipp. J. Sci., Bot. 11: 134. 1944 Philipp. J. Sci. 76: 46, 202. (S) Columella trifolia var. cinerea (Lam.) Merr. Distribution: Borneo, Philippines (Ubian Island, Sulu 1923 Enum. Philipp. Fl. Pl. 3: 10. Archipelago) (S) (L.) Domin (4) Causonis trifolia (L.) Raf. 1927 Biblioth. Bot. 89: 371. 1830 Med. Fl. 2: 122. (S) Cayratia trifolia var. cinerea (Lam.) Quisumb. 1944 Philipp. J. Sci. 76: 47. (B) Vitis trifolia L. Distribution: Africa (Gabon, Rhodesia), Madagascar, 1753 Sp. Pl. 1: 203. Pakistan, Sri Lanka, India, Assam, Nepal, Bangladesh, China (Yunnan), Myanmar, Thailand, Vietnam, Cambodia, Laos, Peninsular Malaysia, Indonesia (Sumatra, Java, Borneo), Brunei, Australia (Western Australia, Northern Territory, Queensland), Philippines (widespread). 7. Tetrastigma (Miq.) Planch. Tetrastigma (Miq.) Planch., in DC., Monogr. Phan. 5: 423. 1887. Vitis sect. Tetrastigma Miq., Ann. Mus. Lugd.-Bat. 1: 72. 1863. Climbers, mostly evergreen, polygamo-dioecious; stems striate, initially terete, often becoming flattened with age; underground root system extensive; tendrils leaf-opposite, simple or branched, branches subtended by a bract. Leaves pedate with 5-11 leaflets, or sometimes palmate with 5-7 leaflets, or trifoliate, to occasionally 1-2-foliolate or 1-foliolate; stipules caducous. Inflorescence usually axillary, an umbellate, dichotomous or corymbose cyme. Flowers 4-merous; calyx cupular, entire or slightly lobed; corolla valvate in aestivation, reflexed at anthesis, caduceus; stamens inserted on the receptacle, filaments erect; anthers in staminate flowers dorsifixed, introrse, opening by longitudinal slits, stamens in pistillate flowers reduced to staminodes; floral disc inconspicuous, adnate to and entirely surrounding the base of the ovary in pistillate flowers, almost completely embracing the reduced ovary in staminate flowers; style short, stigma large, 4-lobed in pistillate flowers, capitate and undivided in staminate Figure 6. Causonis pterita (Merr.) J. Wen & L.M. Lu (E.D. Merrill 5388, Philippines, Ubian Island, Oct 1906, isotype, US, barcode flowers. Fruit a berry, pyriform, globose to ellipsoid, 1-4 00094609). A. Overall specimen; B. Fruits. seeded; seeds ovoid, globose or pyriform, convex on the

235 Philippine Journal of Science Wen et al.: Vitaceae in the Philippines… Vol. 142: Special Issue back, perichalaza ca. 2/3 to nearly the entire length of the Distribution: Peninsular Malaysia, Indonesia (Java, dorsal surface; endosperm ruminate, T or M-shaped in Sumatra, Sulawesi), Borneo (Sabah), Philippines cross section. 2n = 22 or 44 rarely 52. (Luzon). About 95 species primarily in tropical and subtropical Asia (6) Tetrastigma diepenhorstii (Miq.) Latiff with five species in Australia (Latiff 1983, 1991, 2001a, 2001 Folia Malaysiana 2 (3): 185. b, Jackes 1989, Chen et al. 2011a, b). About 24 species (15 endemic) in the Philippines. (B) Vitis diepenhorstii Miq. 1860 Fl. Ind. Bat. Suppl. 1: 515. (1) Tetrastigma brunneum Merr. Distribution: Indonesia (Sumatra), Borneo, Philippines 1912 Philipp. J. Sc., Bot. 7: 85. (Luzon, Mindanao). Distribution: Philippines (Luzon, Mindanao) (7) Tetrastigma ellipticum Merr. (2) Tetrastigma clementis Merr. 1916 Philipp. J. Sci., Bot. 11: 138. 1916 Philipp. J. Sci., Bot. 11: 137. Distribution: Philippines (Basilan). Distribution: Philippines (Mindanao, Panay). (8) Tetrastigma everettii Merr. (3) Tetrastigma coriaceum (DC.) Gagnep 1916 Philipp. J. Sci., Bot. 11: 139. 1910 Not. Syst. (Paris) 1, 320. Distribution: Philippines (Negros). (B) Cissus coriacea DC. 1824 Prodr. 1, 632. (9) Tetrastigma glabratum (Blume) Planch. (S) Cissus leucostaphyla Dennst. 1887 Monogr. Phan. 5: 430. 1818 Schlussel Hort. Malab. 17, 19, 33. (B) Cissus glabrata Blume (S) Cissus lanceolaria Roxb. nom. nud. 1825 Bijdr. Fl. Nederl. Ind. 189. 1814 Hort. 11. Distribution: Indonesia (Java), Borneo, Philippines (S) Cissus lanceolaria Roxb. nom. illeg. (Luzon, Mindoro, Palawan, Polillo). 1820 Flora Indica 1: 430. (10) Tetrastigma harmandii Planch. Based on same type as Cissus leucostaphyla Dennst. 1887 Monogr. Phan. 5: 425. (S) Vitis lanceolaria Wallich ex Wight & Arn. 1834 Prodr. Fl. Ind. Orient. 1: 128. (S) Tetrastigma sorsogonense Elmer, nom. nud. (S) Vitis muricata Wight & Arn. nom. illeg. 1925 Leafl. Philipp. Bot. 9: 3134. 1834 Prodr. 128. Also Leafl. Philipp. Bot. 10: 3810 (1939), Based on same type as Cissus leucostaphyla Dennst. pro syn. (S) Tetrastigma lanceolarium (Roxb.) Planch., nom. illeg. (S) Tetrastigma lagunense Elmer, nom. nud. pro syn. 1887 Monogr. Phan. 5: 433. 1939 Leafl. Philipp. Bot. 10: 3810. (S) Tetrastigma leucostaphylum (Dennst.) Alston Distribution: China (Hainan), Vietnam, Cambodia, 1977 Taxon 26: 539. Laos, Philippines (Luzon, Mindoro, Panay). This species is a common host of Rafflesia and has (11) Tetrastigma laxum Merr. had a long history of nomenclatural confusion (Latiff 1916 Philipp. J. Sci., Bot. 11: 140. 2001b, Veldkamp 2009). Nevertheless, the species Distribution: Philippines (Luzon). delimitation needs reevaluation. (12) Tetrastigma littorale Merr. Distribution: India, Assam, Sikkim, Nepal, Bhutan, 1916 Philipp. J. Sci., Bot. 11: 141. Bangladesh, Myanmar, Thailand, Vietnam, Laos, Distribution: Philippines (Palawan). Cambodia, Peninsular Malaysia, Singapore, Indonesia, (13) Tetrastigma loheri Gagnep. New Guinea, Philippines (widespread). 1910 Notul. Syst. (Paris) 1: 265. (4) Tetrastigma corniculatum Merr. Distribution: Borneo, Philippines (Luzon, Mindanao, 1918 Philipp. J. Sci., Bot. 13: 26. Mindoro, Palawan). Distribution: Philippines (Leyte, Mindoro). (14) Tetrastigma magnum Merr. (5) Tetrastigma dichotomum (Blume) Planch. 1916 Philipp. J. Sci., Bot. 11: 142. 1887 Monogr. Phan. 5: 441. Distribution: Philippines (Luzon, Mindoro). (B) Cissus dichotoma Blume (15) Tetrastigma mindanaense Merr. 1825 Bijdr. Fl. Nederl. Ind. 186. 1921 Philipp. J. Sci. 17: 279. (S) Vitis dichotoma (Blume) Miq. Distribution: Philippines (Mindanao). 1863 Ann. Mus. Bot. Lugd.-Bat. 1: 77.

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(16) Tetrastigma pachyphyllum (Hemsl.) Chun 1940 Sunyatsenia 4: 235. (B) Vitis pachyphylla Hemsl. 1886 J. Linn. Soc., Bot. 23: 425. (S) Tetrastigma crassipes var. strumarum Planch. 1887 Monogr. Phan. 5: 427. (S) Tetrastigma strumarum (Planch.) Gagnep. 1910 Notul. Syst. (Paris) 1: 267. Distribution: Annam, China (Guangdong, Hainan), Laos, Vietnam, Cambodia, Philippines (Luzon). (17) Tetrastigma papillosum (Blume) Planch. 1887 Monogr. Phan. 5: 429. (B) Cissus papillosa Blume 1825 Bijdr. Fl. Nederl. Ind. 183. (S) Cissus suberosus Elmer non (Welw. ex Bak.) Planch. 1908 Leafl. Philipp. Bot. 2: 493. (S) Vitis papillosa (Blume) Backer 1911 Schoolfl. Java, 252. Distribution: Thailand, Peninsular Malaysia, Borneo, Sulawesi, New Guinea, Java, Sumatra, Philippines (Biliran, Catanduanes, Luzon, Mindanao, Negros). (18) Tetrastigma pisicarpum (Miq.) Planch. 1887 Monogr. Phan. 5: 441. (B) Vitis pisicarpa Miq. 1863 Ann. Mus. Bot. Lugd.-Bat. 1: 79. Distribution: Indonesia, Papua New Guinea, Australia (Queensland), Philippines (Luzon, Mindanao). Figure 7. Tetrastigma silvestrei Elmer ex J. Wen & Boggan (A.D.E. Elmer 14663, Philippines, Luzon, Province of Sorsogon, (19) Tetrastigma robinsonii Merr. Irosin (Mt. Bulusan), 750 ft, Oct 1915, holotype, US, barcode 1916 Philipp. J. Sci., Bot. 11: 142. 01150946). A. Overall specimen; B. Ventral side of seed; Distribution: Philippines (Luzon). C. Seed showing chalaza; D. Fruit and peduncle. (20) Tetrastigma sepulchrei Merr. 1912 Philipp. J. Sci., Bot. 7: 88. globose, pale white, fleshy, drying black, 10-12.5 mm Distribution: Philippines (Luzon). in diameter, 3-4-seeded. Seeds oval-triangular, 8-8.5 × 7-7.5 mm, ventral infolds linear and long, arching, (21) Tetrastigma silvestrei Elmer ex J. Wen & Boggan, chalaza oval. sp. nov. (Figure 7) (S) Tetrastigma silvestrei Elmer, nom. inval. Elmer (1939) published Tetrastigma silvestrei Elmer 1939 Leafl. Philipp. Bot. 10 (136): 3802. without providing a Latin description. The name was thus invalidly published. Type: Philippines: Luzon, Province of Sorsogon, Irosin (Mt. Bulusan), 750 ft, Oct 1915, fr, A.D.E. Elmer 14663 (22) Tetrastigma simplicifolia (Merr.) J. Wen & Boggan, (holotype: US). comb. nov. (Figure 8) (B) Columella simplicifolia Merr. Glabrous high woody climber. Leaves pedate, usually 1916 Philipp. J. Sci., Bot. 11: 135. 7-foliolate but occasionally 5-foliolate, glabrous; (S) Cayratia simplicifolia (Merr.) Quisumb. petioles 5-8 cm long; leaflet blades subcoriaceous, 1944 Philipp. J. Sci. 76: 46, 202. elliptic to ovate, 4-10.5 × 2.5-6 cm, abruptly acute at This species was placed in Cayratia by Quisumbing apex, rounded to broadly acute at base, often oblique on (1944). We herein transfer it to Tetrastigma, as the lateral leaflets, undulately toothed to sparsely serrate at fruits possess short, 4-lobed inconspicuous stigmas. margin, lateral veins 5-7 on each side; petiolules 1-1.5 Distribution: Philippines (Leyte). cm on lateral leaflets, 1.5-3 cm on terminal leaflets, glabrous. Infructescence on specialized lateral twigs (23) Tetrastigma stenophyllum Merr. or in leaf axils, ca. 10 cm long, peduncle stout; pedicels up to 1 cm long. Flowers not seen. Fruits compressed

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by the Laboratory of Analytical Biology of the National Museum of Natural History, the Smithsonian Institution. Curators of A, BO, F, K, KEP, L, LAE, LBC, PNH, SING and US are acknowledged for permitting the examination of their collections either through loan or during visits.

REFERENCES BLANCO FM. 1837. Flora de Filipinas, segun el sistema sexual de Linneo. Manila: En la Imprenta de Sto. Thomas. 887p. BLANCO FM. 1845. Flora de Filipinas, segun el sistema sexual de Linneo: segunda impression, corregida y aumentada por el mismo autor. Manila: D. Miguel Sanchez. 619p. BLANCO FM. 1877. Flora de Filipinas, Ed 3 Vol 1. Manila: Plana y C.a. 350p. CHEN I, MANCHESTER SR. 2007. Seed morphology of modern and fossil Ampelocissus (Vitaceae) and implications for phytogeography. American Journal of Botany 94(9): 1534-1553. CHEN PT, CHEN LQ, WEN J. 2011a. The first phylogenetic analysis of Tetrastigma (Miq.) Planch., the host of Rafflesiaceae. Taxon 60(2): 499-512. Figure 8. Tetrastigma simplicifolia (Merr.) J. Wen & Boggan (M. Ramos 15308, Philippines, Leyte, Dagami, Aug 1912, isotype, US, CHEN PT, WEN J, CHEN LQ. 2011b. Spatial and barcode 00094611). A. Overall specimen; B. Enlarged fruits. temporal diversification of Tetrastigma (Vitaceae). The Gardens’ Bulletin, Singapore 63(1 & 2): 307-327. 1916 Philipp. J. Sci., Bot. 11: 143 DRUMMOND A, RAMBAUT A. 2007. BEAST: Distribution: Philippines (Luzon) Bayesian evolutionary analysis by sampling trees. (24) Tetrastigma trifoliolatum Merr. BMC Evolutionary Biology 7(1): 214. 1914 Philipp. J. Sci., Bot. 9: 370 EDGAR RC. 2004. MUSCLE: multiple sequence Distribution: Philippines (Leyte). alignment with high accuracy and high throughput. Nucleic Acids Research 32(5): 1792-1797. ELMER ADE. 1915. Two hundred twenty six new species ACKNOWLEDGMENTS - II. Leaflets of Philippine Botany 8: 2719-2883. ELMER ADE. 1939. Miscellaneous new species. Leaflets We thank Benito Tan for the invitation to contribute to the of Philippine Botany 10: 3673-3810. special volume in memory of the tragic loss of Leonardo Co. This study was supported by the US National Science FELSENSTEIN J. 1985. Confidence limits on phylogenies: Foundation (grant DEB 0743474 to S.R. Manchester and an approach using the bootstrap. Evolution 39(4): 783-791. J. Wen), and the Small Grants Program of the National GALET P. 1967. Recherches sur les methods Museum of Natural History, the Smithsonian Institution. d’identification et de classification des Vitacées des We are grateful to Elizabeth Widjaja, Leng Guan Saw, temperees. II. [Doctoral Dissertation]. Montpellier, Richard Chung, Ruth Kiew, Tingshuang Yi, and Leonardo France: Montpellier University. Co for field assistance, Nora Abdul Karim of the Singapore Botanic Gardens for her kind help with sample HARTLEY TG. 2001. On the taxonomy and biogeography collection, Robin Everly for assistance with references, of Euodia and Melicope (Rutaceae). Allertonia 8(1): Larry Dorr for advice on nomenclature, and Sue Lutz 1-319. and Ingrid Lin for assistance with preparing the figures. HILLIS DM, BULL JJ. 1993. An empirical test of Laboratory work was done at and partially supported

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bootstrapping as a method for assessing confidence the Philippine flora. Philippine Journal of Science in phylogenetic analysis. Systematic Biology 42(2): (Botany) 2: 421-428. 182-192. MERRILL ED. 1912a. A flora of Manila. Manila: Bureau JACKES BR. 1987. Revision of the Australian Vitaceae, of Printing. 490p. 2. Cayratia Juss. Austrobaileya 2(4): 365-379. MERRILL ED. 1912b. Sertulum Bontocense. New or JACKES BR. 1989. Revision of the Australian Vitaceae, interesting plants collected in Bontoc Subprovince, 5. Tetrastigma (Miq.) Planchon. Austrobaileya 3 (1): Luzon, by Father Morice Vanoverbergh. Philippine 149-158. Journal of Science (Botany) 7(2): 71-107. KIRCHHEIMER F. 1939. Rhamnales I: Vitaceae. In: MERRILL ED. 1916. New or interesting Philippine Jongmans W (ed). Fossilium Catalogus Vol 2. Plantae. Vitaceae. Philippine Journal of Science (Botany) Gravenhage, Netherland: Dr. W. Junk. p. 1-174. 11(3): 125-145. LATIFF A. 1981. Studies in Malesian Vitaceae V. MERRILL ED. 1918a. New or noteworthy Philippine The genus Cayratia in the Malay Peninsula. Sains plants, XIII. Philippine Journal of Science (Botany) Malaysiana 10(2): 129-139. 13(1): 1-66. LATIFF A. 1982. Studies in Malesian Vitaceae, I-IV. MERRILL ED. 1918b. Species Blancoanae: A critical Federation Museums Journal 27: 46-93. revision of the Philippines species of plants described by Blanco and Llanos. Manila: Bureau of Science LATIFF A. 1983. Studies in Malesian Vitaceae VII. Publications No. 12. 425p. The genus Tetrastigma in the Malay Peninsula. The Gardens’ Bulletin, Singapore 36(2): 213-228. MERRILL ED. 1920. New or noteworthy Philippine plants, XVI. Philippine Journal of Science 17(3): LATIFF A. 1991. Studies in Malesian Vitaceae X. Two 239-323. new species of Tetrastigma from Borneo. Blumea 35(2): 559-564. MERRILL ED. 1923. An enumeration of Philippine flowering plants. Vol. 3. Manila: Bureau of Printing. LATIFF A. 2001a. Diversity of the Vitaceae in the Malay 628p. Archipelago. Malayan Nature Journal 55(1 & 2): 29- 42. MERRILL ED. 1926. An enumeration of Philippine flowering plants. Vol. 4. Manila: Bureau of Printing. LATIFF A. 2001b. Studies in Malesian Vitaceae XII: 515p. Taxonomic notes on Cissus, Ampelocissus, Nothocissus and Tetrastigma and other genera. Folia Malaysiana MIKI S. 1956. Seed remains of Vitaceae in Japan. Journal 2(3): 179-189. of the Institute of Polytechnics, Osaka City University, Series D, 7: 247-271. LIU XQ, ICKERT-BOND SM, CHEN LQ, WEN J. 2013. Molecular phylogeny of Cissus L. of Vitaceae NIE ZL, SUN H, CHEN ZD, MENG Y, MANCHESTER (the grape family) and evolution of its pantropical SR, WEN J. 2010. Molecular phylogeny and intercontinental disjunctions. Molecular Phylogenetics biogeographic diversification of Parthenocissus and Evolution 66(1): 43-53. (Vitaceae) disjunct between Asia and North America. American Journal of Botany 97(8): 1342-1353. LU LM, WEN J, CHEN ZD. 2012. A combined morphological and molecular phylogenetic analysis of NIE ZL, SUN H, MANCHESTER SR, MENG Y, LUKE Parthenocissus (Vitaceae) and taxonomic implications. Q, WEN J. 2012. Evolution of the intercontinental Botanical Journal of the Linnean Society 168(1): 43-63. disjunctions in six continents in the Ampelopsis clade of the grape family (Vitaceae). BMC Evolutionary LU LM, WANG W, CHEN ZD, WEN J. 2013. Phylogeny Biology 12(1): 1-17. of the non-monophyletic Cayratia Juss. (Vitaceae) and implications for character evolution and biogeography. NYLANDER JA. 2004. MrModeltest. Ver. 2. Program Molecular Phylogenetics and Evolution 68(3): 502- distributed by the author. Uppsala: Evolutionary 515. Biology Centre, Uppsala University. MERRILL ED. 1905. A review of the identifications QUISUMBING E. 1944. New or interesting Philippine of the species described in Blanco’s Flora de plants, II. Philippine Journal of Science 76(3): 37-55. Filipinas. Manila: Publication No. 27 of the Bureau QUISUMBING E, MERRILL ED. 1928. New Philippine of Government Laboratories. 132p. plants. Philippine Journal of Science 37(2): 133-212. MERRILL ED. 1907. Some genera and species new to RAMBAUT A. 2009. FigTree 1.3.1. Available from:

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http://tree.bio.ed.ac.uk/software/figtree/. grape family (Vitaceae). Systematic Botany 37(4): 941-950. RAMBAUT A, DRUMMOND AJ. 2007. Tracer. Ver. 1.4. Available from: http://beast.bio.ed.ac.uk/Tracer/. VELDKAMP JF. 2009. Notes on the names of the Tetrastigma (Vitaceae) hosts of Rafflesia REID EM, CHANDLER MEJ. 1933. The London Clay (Rafflesiaceae). Reinwardtia 13(1): 75-78. Flora. London: British Museum (Natural History). 561p. WEN J. 2007a. Vitaceae. In: Kubitzki K (ed). The families and genera of vascular plants. Vol. 9. Berlin: Springer- REN H, LU LM, SOEJIMA A, LUKE Q, ZHANG DX, Verlag. p. 467-479. CHEN ZD, WEN J. 2011. Phylogenetic analysis of the grape family (Vitaceae) based on the noncoding WEN J. 2007b. Leeaceae. In: Kubitzki K (ed). Families plastid trnC-petN, trnH-psbA, and trnL-F sequences. and Genera of Vascular Plants. Vol. 9. Berlin: Springer- Taxon 60(3): 629-637. Verlag. p. 221-225. RIDSDALE CE. 1974. A revision of the family Leeaceae. WEN J, NIE ZL, SOEJIMA A, MENG Y. 2007. Phylogeny Blumea 22(1): 57-100. of Vitaceae based on the nuclear GAI1 gene sequences. Canadian Journal of Botany 85(10): 731-745. RONQUIST F, HUELSENBECK JP. 2003. MrBayes 3: Bayesian phylogentic inference under mixed models. WEN J, Xiong ZQ, Nie ZL, Mao LK, Zhu YB, Kan Bioinformatics 19(12): 1572-1574. XZ, Ickert-Bond SM, Gerrath J, Zimmer EA, Fang XD. 2013. Transcriptome sequences resolve deep RONQUIST F, TESLENKO M, VAN DER MARK P, relationships of the grape family. PLoS ONE 8(9): AYRES DL, DARLING A, HÖHNA S, LARGET e74394. B, LIU L, SUCHARD MA, HUELSENBECK JP. 2012. MrBayes 3.2: Efficient Bayesian phylogenetic WHITFELD PR, BOTTOMLEY W. 1983. Organization inference and model choice across a large model space. and structure of chloroplast genes. Annual Review of Systematic Biology 61(3): 539-542. Plant Physiology 34(1): 279-310. ROSSETTO M, JACKES BR, SCOTT KD, HENRY RJ. 2002. Is the genus Cissus (Vitaceae) monophyletic? Evidence from plastid and nuclear ribosomal DNA. Systematic Botany 27(3): 522-533. ROSSETTO M, CRAYN DM, JACKES BR, PORTER C. 2007. An updated estimate of intergeneric phylogenetic relationships in the Australian Vitaceae. Canadian Journal of Botany 85(8): 722-730. SOEJIMA A, WEN J. 2006. Phylogenetic analysis of the grape family (Vitaceae) based on three chloroplast markers. American Journal of Botany 93(2): 278-287. STAMATAKIS A. 2006. RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22(21): 2688-2690. SÜESSENGUTH K. 1953. Vitaceae. In: Engler A, Prantl K (eds). Die Natürlichen Pflanzenfamilien, 2nd ed. Vol. 20d. Berlin: Duncker & Humbolt. p. 174-333. SWOFFORD DL. 2003. PAUP*. Phylogenetic analysis sing parsimony (*and other methods). Ver. 4.0b10. Sunderland, Massachusetts: Sinauer Associates. TIFFNEY BH, BARGHOORN ES. 1976. Fruits and seeds of the Brandon Lignite. I. Vitaceae. Review of Palaeobotany and Palynology 22(3): 169-191. TRIAS-BLASI A, PARNELL JAN, HODKINSON TR. 2012. Multi-gene region phylogenetic analysis of the

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Appendix. Vourchers (collection number, locality) and GenBank accession numbers. "–" indicates missing data. Abbreviations of herbaria are as follows: MO, the Missouri Botanical Garden Herbarium, St. Louis, Missouri; PE, Herbarium of Institute of Botany, the Chinese Academy of Sciences, Beijing; US, the United States National Herbarium, Washington, D.C. The 52 sequences starting with GenBank accession # KF were generated in this study. trnC- Taxon Voucher Locality atpB-rbcL rps16 trnH-psbA trnL-F petN2r Ampelocissus acapulcensis (Kunth) Wen 8696 (US) Mexico, JQ182472 JX476543 JF437172 JF437058 JF437058 Planch. Ampelocissus africana (Lour.) Merr. Luke & Luke 11449 (US) , Makindu JQ182448 JQ182603 − JQ182507 JQ182553 Ampelocissus cinnamomea Planch. Wen 11697 (US) Malaysia, Borneo, Sabah KF022263 KF022272 KF022281 KF022296 KF022307 Ampelocissus elegans Gagnep. Wen 11825 (US) Malaysia, Selangor KC166289 KC166380 KC166476 KC166544 KC166618 Ampelocissus elephantina Planch. Wen 9583 (US) Madagascar, Antsiranana HM585516 HM585792 − HM585659 HM585932 Ampelocissus erdwendbergii Planch. Wen 8697 (US) Mexico, Chiapas KC166290 KC166381 KC166468 KC166545 KC166619 Ampelocissus gracilis Planch. Wen 11684 (US) Singapore KF022264 KF022273 KF022282 KF022297 KF022308 Ampelocissus javalensis (Seem.) W.D. Wen 6920 (US) Costa Rica, Limon AB234911 AB234943 KC166469 KC166546 AB234984 Stevens & A. Pool Ampelocissus latifolia (Roxb.) Planch. Akfandray s.n. (US) India KF022265 KF022274 − KF022298 KF022309 Ampelocissus martini Planch. Wen 7421 (US) Thailand, HQ214193 KF022275 KF022283 − AB234985 Ampelocissus obtusata (Welw. ex Baker) , Inyonga- Luke & Luke 11590 (US) JQ182457 JQ182612 KF022284 JQ182510 JQ182556 Planch. Mpanda Ampelocissus thyrsiflora (Blume) Planch. Deden 870 (US) Indonesia, SE Sulawesi JQ182438 JQ182593 KF022285 JQ182499 JQ182546 Ampelopsis bodinieri (H. Lév. & Vaniot) Ren 55193 (US) China, Guangdong JX476427 JX476545 JF437175 JF437061 JF437284 Rehder Ampelopsis cantoniensis (Hook. & Arn.) Wen 10242 (US) Indonesia, SE Sulawesi HM585517 HM585793 JX476667 HM585660 HM5865933 K. Koch Ampelopsis chaffanjonii (H. Lév & Wen 9382 (US) China, Guangxi JQ182475 JQ182630 JF437177 JQ182528 JQ182570 Vaniot) Rehder Ampelopsis cordata Michx. Wen 7141 (US) U.S.A., Illinois (cult.) AB234916 AB234949 JF437178 JF437064 AB234997 Ampelopsis delavayana Planch. ex Wen 9377 (US) China, Guangxi JQ182476 JQ182631 JF437179 JF437065 JF437287 Franch. Ampelopsis denudata Planch. Wen 8695 (US) Mexico, Chiapas JQ182483 JQ182638 JF437180 JQ182534 JQ182577 Ampelopsis glandulosa (Wall.) Momiy. Wen 9380 (US) China, Guangxi KC166292 KC166383 JF437181 JF437067 JF437289 Ampelopsis glandulosa (Wall.) Momiy. Wen 8289 (US) Philippines, Luzon JQ182477 JQ182632 − JQ182529 JQ182571 var. hancei (Planch.) Momiy Ampelopsis glandulosa (Wall.) Momiy. Wen 8300 (US) Philippines, Luzon − − KF022286 KF022299 KF022310 var. hancei (Planch.) Momiy Ampelopsis grossedentata (Hand.-Mazz.) Wen 9336 (US) China, Hunan JQ182479 JQ182634 JF437184 JF437070 JF437292 W.T. Wang Ampelopsis humulifolia Bunge Wen 8519 (US) China, Beijing JQ182464 JQ182619 − JQ182520 JQ182563 Ampelopsis japonica (Thunb.) Makino Ren 55207 (US) China, Guangdong − − JF437185 JF437071 KF022311 Ampelopsis rubifolia (Wall.) Planch. Wen 9285 (US) China, Hunan JQ182473 JX476546 JF437186 JF437072 JF437293 Causonis japonica (Thunb.) Raf. Wen 8330 (US) Malaysia, Selangor KC166312 KC166403 JF437192 JF437078 JF437298 Causonis japonica (Thunb.) Raf. Wen 9262 (US) China, Sichuan KC166310 KC166401 JF437197 JF437083 JF437300 Causonis trifolia (L.) Raf. Wen 7488 (US) Thailand, Chiang Mai KC166323 – KC166500 KC166574 AB235007 Cayratia acris (F. Muell.) Domin Wen 12183 (US) Australia, cult. in Sydney KC166293 KC166384 KC166471 KC166548 KC166621 Indonesia, cult. in Bogor Cayratia cardiophylla Jackes Chen & Lu 160 (PE) KC428766 KC428777 KC428790 KC428809 KC428824 Botanical Gardens (F.Muell.) Domin Wen 12184 (US) Australia, cult. in Sydney KC166297 KC166388 KC166475 KC166552 KC166625 Cayratia corniculata (Benth.) Gagnep. Wen 8245 (US) Philippines, Luzon KC166300 KC166391 KC166478 KC166555 KC166628 Cayratia debilis (Baker) Suess. Carvalho 4136 (MO) Equatorial Guinea, Bioco KC166302 KC166393 JF437190 JF437076 JF437296 Cayratia geniculata Gagnep. Wen 10275 (US) Indonesia, SE Sulawesi HM585519 HM585795 KC166480 KC585662 HM585935 Cayratia gracilis (Guill. & Perr.) Suess. Luke & Luke 11685 (US) Kenya, Mombase KC166306 KC166397 KC166483 KC166559 KC166632 Cayratia imerinensis (Baker) Desc. Wen 9607 (US) Madagascar, Antsiranana KC166306 KC166397 KC166484 KC166560 KC166633 Cayratia maritima Jackes Wen 9403 (US) China, Taiwan KC166314 KC166405 JF437193 JF437079 JF437299 Cayratia mollissima Gagnep. Wen 10301 (US) Indonesia, SE Sulawesi KC166322 KC166413 KC166499 JQ182500 JQ182547 Cayratia pedata Gagnep. Wen 7428 (US) Thailand, Chiang Mai AB234919 AB234952 KC166496 KC166571 AB235004 Cayratia triternata (Baker) Desc. Wen 9643 (US) Madagascar, Antsiranana KC166324 KC166414 KC166501 KC166575 KC166644

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Cayratia wrayi (King) Gagnep. Deng 3983 (US) Myanmar KC166325 KC166415 KC166503 KC166576 KC166645 Cissus adnata Roxb. Wen 10268 (US) Indonesia, SE Sulawesi JX476429 JX476547 JX476673 JX476772 JX476858 Cissus albiporcata Masinde & L.E. Luke & Luke 11456 (US) Kenya, Chyulu Plains JX476430 JX476548 JF437201 JF437087 JF437304 Newton Cissus antarctica Vent. Wen 6684 (US) USA, California (cult.) JX476433 JX476551 JX476677 JX476775 JX476860 Kenya, Tsavo West Na- Cissus aphyllantha Gilg Luke 11674 (US) JX476435 JX476553 JX476679 JX476777 JX476862 tional Park Belgium, National Bo- Cissus aralioides Planch. Aplin 19870062 (US) JX476438 JX476556 JF437202 JF437088 JF437305 tanical Garden (cult.) Cissus assamica (M.A. Lawson) Craib Wen 10273 (US) Indonesia, SE Sulawesi JX476439 JX476557 JX476681 JX476780 JX476865 Kenya, Nr KWS Rhino Cissus cornifolia (Baker) Planch. Luke & Luke 11452 (US) JX476449 JX476567 JF437205 JF437091 JF437308 Camp. Cissus diffusa (Miq.) Amshoff Wen 10756 (US) Indonesia, Papua JX476451 JX476569 JX476689 JX476787 JX476871 Jongkind et al. 1813 Ghana, Atewa Range For- Cissus diffusiflora (Baker) Planch. JX476452 JX476570 − JX476788 JX476872 (MO) est Reserve Belgium, National Bo- Cissus discolor Blume 20061111 (US) JX476454 JX476572 JF437206 JF437092 JF437309 tanical Garden (cult.) Cissus erosa Rich. Wen 8574 (US) Peru, Arequipa JX476457 JX476575 JF437207 JF437093 JF437310 Cissus faucicola Wild & R.B.Drumm. Kayombo 2344 (MO) Tanzania, Mbeya JX476458 JX476576 JX476694 JX476792 JX476874 Cissus gongylodes (Burch. ex Baker) Nee & Wen 53792 (US) Bolivia, Santa Cruz JX476460 JX476578 JX476696 JX476794 JX476876 Planch. Cissus granulosa Ruiz & Pav. Wen 8573 (US) Peru, Oxapampa JX476462 JX476580 JX476698 JX476796 JX476878 Cissus hastata Miq. Wen 7509 (US) Singapore JX476465 AB234955 JX476701 JX476799 AB235012 Cissus hypoglauca A. Gray Wen 12185 (US) Australia, Wollongong JX476466 JX476583 JX476702 JX476800 JX476881 Cissus integrifolia (Baker) Planch. Luke & Luke 11475 (US) Kenya, Shimba Hills JX476467 JX476584 JX476703 JX476801 JX476882 Cissus javana DC. Shui et al. 81970 (US) China, Yunnan JX476468 JX476585 JX476704 JX476802 JX476883 Cissus lanea Desc. Wen 9536 (US) Madagascar, Antsiranana JX476469 JX476586 JX476705 JX476803 JX476884 Cissus leucophleus (Scott-Elliot) Suess. Wen 9676 (US) Madagascar, Toliara JX476470 JX476587 JX476706 JX476804 JX476885 Cissus madecassa Desc. Wen 9597 (US) Madagascar, Antsiranana JX476471 JX476588 JX476707 JX476805 JX476886 Panama, Barro Colorado Cissus microcarpa Vahl. Wen 11954 (US) JX476472 JX476589 JX476708 JX476806 JX476887 Island Cissus microdonta Vahl. Wen 9635 (US) Madagascar, Antsiranana JX476474 JX476591 JX476710 − JX476889 Cissus nodosa Blume Wen 10713 (US) Indonesia, Papua JX476475 JX476592 JX476711 HM585671 HM585945 Cissus phymatocarpa Masinde & Luke & Luke 11474 (US) Kenya, Diani Forest JX476479 JX476596 JF437209 JF437095 JF437311 L.E.Newton Cissus polita Desc. Wen 9577 (US) Madagascar, Antsiranana JX476482 JX476599 JX476717 JX476813 JX476895 Tanzania, Udzungwa Cissus producta Afzel. Luke et al. 11528 (US) JX476483 JX476600 JF437210 JF437096 JF437312 Mountain Thailand, Chiang Mai Cissus quadrangularis L. Wen 7368 (US) JX476486 JX476603 JF437211 JF437097 JF437313 (cult.) Cissus quarrei Dewit Luke 11581 (US) Tanzania, Tabora-Inyonga JX476488 JX476605 JX476722 − JX476899 Cissus repanda Vahl Wen 7396 (US) Thailand, Chiang Mai JX476489 JX476606 JX476723 JX476818 AB235015 Cissus repens Lam. Ren 55094 China, Yunnan JX476491 JX476608 JX476725 JX476820 JX476901 Cissus rhombifolia Vahl Wen 6992 (US) Costa Rica, Puntarenas JX476495 JX476612 JX476729 JX476824 JX476905 Cissus rotundifolia (Forssk.) Vahl. Luke & Luke 11478 (US) Kenya, Taru JX476499 JX476615 JF437212 JF437098 JF437314 Cissus sagittifera Desc. Wen 9605 (US) Madagascar, Antsiranana JX476501 JX476617 JX476733 JX476827 JX476908 Cissus sciaphila Gilg. Luke & Luke 11477 (US) Kenya, Shimba Hills JX476503 JX476619 JF437214 JF437100 JF437316 Cissus simsiana Roem. & Schult. Nee & Wen 53805 (US) Bolivia, Santa Cruz JX476504 JX476620 JX476734 JX476828 JX476910 Cissus striata Ruiz & Pav. Wen 7424 (US) Chile, Valdivia AB234921 AB234958 JX476747 JX476841 AB235017 Cissus subtetragona Planch. Ren 55110 (US) China, Yunnan JX476519 JX476635 JF437216 JF437102 − Cissus trianae Planch. Wen 8585 (US) Peru, Oxapampa JX476521 JX476637 JX476752 JX476844 JX476925 Cissus trifoliata (L.) L. Wen 7287 (US) USA, Texas JX476523 AB234956 JF437208 JF437094 AB235014 Tanzania, Udzungwa Cissus trothae Gilg & M. Brandt Luke et al. 11537 (US) JX476525 JX476640 JF437217 JF437103 JF437318 Mountain Cissus verticillata (L.) Nicolson & C.E. Wen 8734 (US) Mexico, Chiaps JX476534 JX476648 JF437215 JF437101 JF437317 Jarvis Clematicissus opaca (F. Muell.) Jackes Sun 11005 (US) Australia, Brisbane JX476538 JX476652 JX476767 JX476857 JX476935 & Rossetto Cyphostemma cyphopetalum (Fresen.) Kenya, Nr KWS Rhino Luke & Luke 11451 (US) KC166331 KC166421 JF437221 JF437108 JF437323 Wild & R.B. Drumm. Camp.

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Cyphostemma duparquetii (Planch.) Tanzania, Udzungwa Luke & et. 11534 (US) KC166332 JX476654 JF437222 JF437109 JF437324 Desc. Mountain Cyphostemma dysocarpum (Gilg & M. Luke & Luke 11457 (US) Kenya, Chyulu Plains KC166334 KC166423 JF437223 JF437110 JF437325 Brandt) Desc. Cyphostemma echinocarpa Desc. Wen 9488 (US) Madagascar, Fianarantsoa KC166335 KC166424 KC166509 KC166583 KC166651 Cyphostemma heterotrichum (Gilg & Lovett 4027 (MO) Tanzania, Mbeya KC166338 KC166427 JF437224 JF437111 JF437326 R.E. Fr.) Desc. ex Wild & R.B. Drumm. Cyphostemma kibweziense Verdc. Luke & Luke 11481 (US) Kenya, Mbinzau KC166346 KC166435 JF437229 JF437116 JF437330 Cyphostemma kilimandscharicum (Gilg) Luke & Luke 11469 (US) Kenya, Chyulu Hills KC166348 KC166437 JF437225 JF437112 JF437327 Wild & R.B. Drumm. Cyphostemma kirkianum (Planch.) Wild Luke & Luke 11473 (US) Kenya, Diani Forest KC166349 KC166438 JF437226 JF437113 JF437328 & R.B. Drumm. Cyphostemma macrocarpa Desc. Wen 9589 (US) Madagascar, Antsiranana KC166350 KC166439 KC166518 KC166595 KC166663 Cyphostemma maranguense (Gilg) Desc. Luke & Luke 11468 (US) Kenya, Chyulu Hills KC166352 JX476656 JF437227 JF437114 JF437329 Cyphostemma microdipterum (Baker et Wen 9687 (US) Madagascar, Toamasina KC166354 KC166442 KC166522 KC166598 KC166665 al.) Desc. U.S.A., Missouri Botani- Cyphostemma montagnacii Desc. Wen 6672 (US) AB234923 AB234961 JF437228 JF437115 AB235027 cal Garden (cult.) Cyphostemma sp. Wen 7405 (US) Thailand, Mae Hong Son KC166369 KC166457 – KC166609 KC166676 Cyphostemma thomasii (Gilg & M. Luke & Luke 11448 (US) Kenya, Makindu KC166365 KC166453 JF437230 JF437117 JF437331 Brandt) Desc. Cyphostemma vogelli (Hook.) Desc. Carvalho 4127 (MO) Equatorial Guinea KC166367 KC166455 JF437221 JF437118 JF437332 Cyphostemma zimmermannii Verdc. Luke & Luke 11476 (US) Kenya, Shimba Hills KC166368 KC166456 JF437222 JF437119 JF437333 Leea aculeata Blume Wen 8248 (US) Philippines, Laguna − − KF022287 KF022300 AB235088 Leea acuminatissima Merr. Wen 8242 (US) Philippines, Laguna − − KF022288 KF022301 AB235096 USA, Maryland, cult. Leea guineensis G. Don Wen 2010-091 (US) in Smithsonian Botany KC166370 KC166458 KC166533 KC166610 KC166677 Greenhouse Leea guineensis G. Don Wen 8250 (US) Philippines, Laguna − − JF437234 JF437121 − Leea indica (Burm. f.) Merr. Wen 8341 (US) Malaysia, Selangor JX476542 JX476658 JF437236 JF437123 JF437334 Leea macrophylla Roxb. ex Hornem. & Ren 55105 (US) China, Yunnan − JX476659 JF437237 JF437124 JF437335 Roxb. Leea rubra Blume MAC 05-716 (US) Thailand, − − JF437238 JF437125 − Singapore, Singapore Nothocissus spicifera (Griff.) Latiff Wen 11675 (US) KC166371 KC166459 KC166534 KC166611 KC166678 Botanic Gardens (cult.) Parthenocissus chinensis C. L. Li Nie& Meng 455 (US) China, Sichuan HM223373 HM223320 JF437240 JF437127 HM223263 Parthenocissus dalzielii Gagnep. Wen 9372 (US) China, Hunan − − JF437243 JF437130 JF437338 Parthenocissus henryana (Hemsl.) Nie & Meng 359 (US) China, Guizhou HM223383 HM223329 JF437244 JF437131 HM223272 Graebn. ex Diels & Gilg Parthenocissus laetevirens Rehder Wen 9379 (US) China, Guangxi HM223378 HM223323 JF437245 JF437132 JF437339 Parthenocissus quinquefolia (L.) Planch. Wen 8662 (US) USA, Virginia HM223380 HM223325 KF022289 KF022302 HM223269 Parthenocissus suberosa Hand.-Mazz. Nie & Meng 358 (US) China, Guizhou HM223384 HM223330 JF437247 JF437134 HM223273 Parthenocissus tricuspidata Planch. Nie & Meng 355 (US) China, Guizhou HM223385 HM223331 JF437248 JF437135 HM223274 Parthenocissus vitacea (Knerr) Hitchc. Wen 7234 (US) USA, Texas KC166372 KC166460 JF437249 JF437136 JF437340 Pterisanthes polita M.A. Lawson Wen 11831 (US) Malysia, Selangor KC166374 KC166462 KC166536 KC166613 KC166680 Pterisanthes stonei Latiff Wen 8346 (US) Malaysia, Selangor KF022266 JX476662 KF022290 JF437137 AB235046 Rhoicissus tomentosa (Lam.) Wild & Belgium, National Bo- 19656252 (US) JQ182454 JQ182609 JF437251 JF437139 JF437342 R.B. Drumm. tanical Garden (cult.) Rhoicissus tridentata (L.f.) Wild & R.B. Luke & Luke 11453 (US) Kenya, Chyulu Hills JQ182443 JQ182598 JF437250 JF437138 JF437341 Drumm. Tetrastigma bioritsense (Hayata) Hsu & Wen 9451 (US) China, Taiwan HM585548 HM585824 JF437252 JF437140 HM585964 Kuoh Tetrastigma brunneum Merr. Wen 8240 (US) Philippines, Luzon HM585549 HM585825 − HM585689 HM585965 Tetrastigma campylocarpum Planch. Wen 10517 (US) China, Yunnan HM585550 HM585826 – HM585690 HM585966 Tetrastigma ceratopetalum C.Y. Wu Shui 81836 (US) China, Yunnan HM585557 HM585833 KC166538 HM585697 HM585973 Tetrastigma curtisii (Ridl.) Suess. Wen 10277 (US) Indonesia, SE Sulawesi HM585563 HM585839 – HM585703 HM585979 Tetrastigma delavayi Gagnep. Wen 7443 (US) Thailand, Chiang Mai AB234932 AB234968 KC166539 HM585705 AB235050 Tetrastigma diepenhorstii (Miq.) Latiff Wen 8261 (US) Philippines, Luzon HM585567 HM585843 − HM585707 HM585983 Tetrastigma ellipticum Merr. Wen 8260 (US) Philippines, Luzon HM585569 HM585845 − HM585709 HM585985

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Tetrastigma erubescens Planch. Ren 55116 (US) China, Yunnan − − JF437253 JF437141 JF437343 N.P. Pui s.n. (GenBank Tetrastigma garrettii Gagnep. Thailand, Chiang Mai − − JF437254 JF437142 JF437344 sequences) Tetrastigma glabratum Planch. Wen 10666 (US) Indonesia, West Java HM585579 HM585855 − − HM585995 Tetrastigma hemsleyanum Diels & Gilg Nie & Meng 451 (US) China, Sichuan HM585584 HM585860 JF437255 JF437143 HM586000 Tetrastigma jinghongense C.L. Li Wen 8471 (US) China, Yunnan HM585590 HM585866 JF437256 JF437144 HM586006 Tetrastigma lanyuense C.E. Chang Wen 9404 (US) China, Taiwan HM585593 HM585869 JF437257 JF437145 HM586009 Tetrastigma laxum Merr. Wen 8314 (US) Philippines, Luzon HM585602 HM585877 − HM585740 HM586018 Tetrastigma loheri Gagnep. Wen 10202 (US) Indonesia, SE Sulawesi HM585605 HM585880 − HM585743 HM586021 Tetrastigma obtectum (Wall.) Planch. Nie & Meng 454 (US) China, Sichuan HM585614 HM585888 JF437266 JF437154 JF437349 Tetrastigma pachyphyllum (Hemsl.) Chun Wen 8319 (US) Philippines, Luzon HM585616 HM585891 JF437259 JF437147 HM586032 Tetrastigma papillosum Planch. Wen 8401 (US) Malaysia, Pahang HM585617 HM585892 − HM585754 HM586033 Tetrastigma planicaule Gagnep. Wen 10904 (US) Vietnam, Ninh Binh HM585622 HM585897 KC166540 HM585759 HM586037 Tetrastigma pyriforme Gagnep. Wen 8370 (US) Malaysia, Pahang HM585624 HM585898 JF437268 JF437156 JF437351 Tetrastigma rumicispermum Planch. Tibet 2003 (US) China, Tibet HM585626 HM585900 – HM585763 HM586041 Tetrastigma serrulatum Planch. Nie & Meng 445 (US) China, Yunnan HM585627 HM585901 JF437261 JF437149 HM586042 Thailand, Bahng Mah Pah Tetrastigma siamense Gagnep. 03-439 (US) – – JF437262 JF437150 JF437347 District Tetrastigma sp. Wen 8256 (US) Philippines, Luzon HM585638 HM585911 − HM585775 HM586053 Tetrastigma sp. Wen 11412 (US) China, Guangdong KC166376 KC166463 KC166541 KC166615 KC166681 Tetrastigma triphyllum (Gagnep.) W.T. Nie & Meng 342 (US) China, Yunnan HM585646 HM585919 JF437263 JF437151 HM586061 Wang Tetrastigma xishuangbannaense C. L. Li Ren 55108 (US) China, Yunnan − − JF437265 JF437153 − Vitis aestivalis Michx. Wen 10428 (US) USA, Delaware KF022267 KF022276 KF022291 KF022303 KF022312 Vitis betulifolia Diels & Gilg Wen 8217 (US) China, Chongqing AB234937 KF022277 KF022292 KF022304 AB235075 Vitis chunganensis Hu Wen 11406 (US) China, Guangdong KF022268 KF022278 KF022293 KF022305 KF022313 Vitis flexuosaThunb. Wen 10647 (US) China, Yunnan HM585656 HM585929 KF022294 HM585789 HM586071 Vitis heyneana Roem. & Schult Wen 9378 (US) China, Guangxi KF022269 JX476666 JF437273 JF437161 JF437354 Vitis labrusca L. Wen 8652 (US) USA, Virginia JX507360 JX507361 JX507362 JX507363 JX507364 Vitis lanata Roxb. Wen 9197 (US) China, Tibet KF022270 KF022279 JF437275 JF437163 JF437356 Vitis mengziensis C. L. Li Nie & Meng 415 (US) China, Yunnan HM223387 HM223333 JF437270 JF437158 HM223276 Vitis popenoei J.L. Fennell Wen 8724 (US) Mexico, Chiapas HM585657 HM585930 JF437276 JF437164 HM586072 Vitis riparia Michx. Wen 8658 (US) USA, Virginia JQ182453 JQ182608 JF437277 JF437165 JF437357 Vitis rotundifolia Michx. Wen 11087 (US) USA, Arkansas KC166379 KC166466 KC166542 JF437166 JF437358 Vitis thunbergii Sieb. & Zucc. Wen 9446 (US) China, Taiwan − AB234976 JF437278 JF437167 AB235082 Vitis tilifolia Humb. & Bonpl. Wen 11894 (US) Panama KF022271 KF022280 KF022295 KF022306 KF022314 Yua austro-orientalis (F.P. Metcalf) C.L. S. Ickert-Bond 1313 (US) China, Guangdong − AB234977 KC166543 JF437170 AB235085 Li Yua thomsoni (M.A. Lawson) C.L. Li Nie & Meng 469 (US) China, Sichuan HM223389 HM223335 − JF437171 HM223277

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