Gardens’Bulletin Singapore 67(2): 361-386. 2015 361 doi: 10.3850/S2382581215000307 A contribution to the systematics ofXylopia (Annonaceae) in Southeast Asia & D.M. Johnson N.A. Murray & Department ofBotany Microbiology, Ohio Wesleyan University OH USA Delaware, 43015 [email protected] [email protected] ABSTRACT. Herbarium andfield studyofXylopia L. (Annonaceae) fortheFlora ofPeninsular Malaysia and the Flora of Thailand projects has clarified regional diversity patterns within this ecologically significant lowland rainforest genus. Two species groups represented within & Southeast Asian floras ai'e delineated, one centred onXylopiaferruginea (Hook.f Thomson) & Baill. and the other onXylopia malayana Hook.f Thomson. In XhoXylopiaferruginea group, & a new species, Xylopia erythrodactyla D.M.Johnson N.A.Murray, is distinguished from X. ferruginea, and a new combination, Xylopia sumatrana (Miq.) D.M.Johnson & N.A.Murray, is proposed, based on an earlier name for the species currently known as Xylopia stenopetala Oliv. In the Xylopia malayana group, review of the species Xylopia elliptica Maingay ex & Hook.f. Thomson resulted in the recognition oftliree additional species: Xylopia platycarpa & D.M.Johnson N.A.Murray, from southern Thailand and northwestern Peninsular Malaysia, & Xylopia ngii D.M.Johnson N.A.Murray, fi'om Peninsular Malaysia, Sumatra and Borneo, & and Xylopia heterotricha D.M.Johnson N.A.Mun*ay, from Sumatra and Borneo. The taxon & Xylopia malayana Hook.f. Thomson var. obscura Kochummen is placed in synonymy under & Xylopia elliptica sensu stricto. Xylopiafusca Maingay ex Hook.f. Thomson var. sessiliflora Kochummen & Whitmore is distinguished from Xylopiafusca, and raised to species status as Xylopia sessiliflora (Kochummen & Whitmore) D.M.Johnson & N.A.Murray. We recognise 23 Xylopia species in the Sundaic region of Southeast Asia, and provide evidence that additional collecting and taxonomic analysis in the region is needed. Keywords. Annonaceae, biogeography, Borneo, Malay Peninsula, Sumatra, Sundaland, Thailand, Xylopia Introduction The Annonaceae, a flowering plant family of2500 species, including the economically important soursop, custard apple and ylang-ylang, is widespread across the tropics. The family is most diverse, and ecologically most significant, in tropical Asia, where it is represented by c. 40 genera and 800 species. In southeastern Asia it is one of & the dominant families in lowland wet forests. Corlett Turner (1997) determined that Annonaceae ranked fourth in species-richness among flowering plant families in Singapore; Appanah et al. (1993) found that Annonaceae ranked first in species diversity among lianas ofMalaysian forests. In long-term ecological plots in Southeast 362 Gard. Bull. Singapore 67(2) 2015 Asia Annonaceae usually rank among the top ten tree families in both number of & individuals and number of species, though not in basal area (see examples in Losos Leigh, 2004). The genus Xylopia L., the only pantropical genus in the family, comprises 180-200 species oftrees and shrubs worldwide. The highest concentration of species in Southeast Asia occurs in the Sundaic region extending from the Kra Isthmus in southern Thailand to Wallace’s Line, a region of high diversity for many plant taxa. Xylopias are distinctive among the Annonaceae in their cone-shaped buds, elongate, aromatic flowers, and dehiscent fruits with seeds bearing arils or fleshy seed coats. In West Africa the peppery fruits ofXylopia aethiopica (Dunal) A.Rich. have long been used as a spice (Dunal, 1817; Burkill, 1985) and are sold commercially. Despite its significance, the genus has never been monographed. In preparing keys and descriptions of the genus Xylopia (Annonaceae) for the Flora ofPeninsular Malaysia and Flora ofThailandprojects we had the opportunity to study material in herbaria with important holdings for the region as well as to observe several species in the field. At the beginning ofour study 13 Xylopia taxa were known from Peninsular Malaysia, Singapore and Thailand combined (Sinclair, 1953, 1955; Kochummen et ah, 1970; Kochummen, 1972a, 1972b; Chalennglin, 2001; Gardner et ah, 2015). Analysis of morphological and preliminary molecular data (Stull et ah, 2011; Thomas et ah, 2015; Stull et ah, in prep.) has shown that the SoutheastAsian species of & Xylopia fall into two groups, one including Xylopiaferruginea (Hook.f. Thomson) & Baill. and a second including Xylopia malayana Hook.f Thomson, each with their respective allies. Our study revealed a much greater diversity of species than previously recognised in both groups. To document this diversity, and reconcile it with the previously existing taxonomy and nomenclature for the genus, the following paper A is presented. full treatment, including keys and distribution maps, is forthcoming. lUCN Conseiwation assessments using (2012) criteria are not included in this account as more data are required for these than we currently have available. Although historical EOOs and AOOs can be calculated, we are conscious that many of the collections have been made in lowland forest areas that have suffered from rapid deforestation. In these cases the Population Reduction (A) criterion would be more appropriate, as has been used for many dipterocarp species (lUCN, 2014), but again we would require more on-the-ground knowledge than we currently have. Xylopiaferruginea group The Xylopia ferruginea group is characterised by stilt roots, relatively long (5-19 mm) flower pedicels, a flat receptacle lacking a staminal cone (Fig. 2K), flat narrowly oblong stamens with a tongue-shaped apex to the anther connective (Fig. 2J), stigmas studded with small papillae (Fig. 21), and rugose seeds (Fig. 2C-D). In addition, most species oftheXylopiaferruginea group have relatively numerous (up to 20) linear and somewhat torulose monocarps. 95 Xylopia in SoutheastAsia 363 Study ofthe group resulted in demarcation ofa new species, and recognition of an earlier name for an existing species. & Xylopia erythrodactyla D.M.Johnson N.A.Murray, sp. nov. Species resembling Xylopia ferruginea in the rusty pubescence of the leaves and flowers and the long narrow monocarps, but differing consistently in the more densely pubescent and thicker leaves, thicker pedicels (1.3-2.5 mm), longer sepals (4.8-7. mm), broader outer petals (3-3.7 rmn wide at the midpoint) and narrowly oblong and weakly torulose monocarps 7.5-10.7 cm long and 0.6-1. cm wide. In contrast, in 1 mm mm Xylopiaferruginea the pedicels are 1-1.3 thick, the sepals are 3-5.5 long, the mm outerpetals are 2-2.2 wide at the midpoint, and the monocarps are linear, strongly tomlose, 6.2-11.6 cm long and 0.4-0.6 cm wide. - TYPE: Malaysia, Sarawak, Teluk Bandung, Santubong, 1st Division, 18 September 1984 (fr), Awa & Ismawi S.47080 (holotype KEP; isotypes ASU, K, L, SAR). (Fig. 1, 2) m Tree up to 30 tall, dbh up to 75 cm, bole smooth with stilt roots at the base; secondary branches drooping. Bark smooth, light brown, brown tinged with red, brick-red, or orange, very finely fissured. Twigs light grey to brown, eventually dark grey, densely femiginous-pubescent/velutinous, eventually glabrate. Leaf with larger blades 12.4-26 cm long, 3.9-7 cm wide, subcoriaceous to coriaceous, strongly discolorous, oblanceolate, oblong-oblanceolate, or narrowly elliptic, base rounded to mm cuneate and short-decurrent, apex short-acuminate, the acumen 2-7 long, glabrous adaxially, densely ferruginous-pubescent, the pubescence especially pronounced along the midrib, secondary veins, and larger higher-order veins abaxially; midrib impressed adaxially, raised abaxially; secondary veins 1 1-14 per side, diverging at 50-60° from the midrib, brochidodromous, these and higher-order veins indistinct mm adaxially, strongly raised abaxially; petiole 5-12 long, deeply canaliculate (margins nearly meeting), pubescent. Inflorescences axillary or from axils of fallen mm leaves, 1-3-flowered, densely ferruginous-pubescent; peduncles 1-2 per axil, 2 mm mm long; pedicels 2 per peduncle, 7-14 long, 1.6-2.5 thick; bract attached 1, mm !/3-!/2 distance from base ofpedicel, 2.9-3 long, ovate, apex acute to obtuse; buds linear-lanceolate, sometimes somewhat falciform and slightly twisted, apex obtuse. mm mm Sepals ’/g-’A-connate, 4-7.9 long, 4-5.3 wide, coriaceous, broadly ovate to triangular, apex acute to acuminate, occasionally obtuse, pubescent along margins and at apex adaxially, ferruginous-pubescent abaxially. Petals pale yellow to white mm in vivo; outer petals curving outward at anthesis, 38-45 imn long, 4-5.3 wide at mm base, 3-3.7 wide at midpoint, linear-lanceolate, obtuse, densely grey-puberulent adaxially, densely feiTuginous-pubescent abaxially; inner petals erect at anthesis, 32- mm mm mm 40 long, c. 3.5 wide at base, c. 1.3 wide at midpoint, linear, densely grey-pubemlent on both surfaces except for glabrous concave base. Stamens up to mm 77, 2.7-3.5 long, narrowly oblong, often setose along edges of anther locules, mm apex of connective 0.4-0.6 long, oblong, densely long-papillate, filament 0.3-0. mm mm long, glabrous; staminal cone absent; outer staminodes c. 18, c. 2.7 long. 8 364 Gard. Bull. Singapore 67(2) 2015 Fig. 1. Xylopia erythrodactyla D.M.Johnson & N.A.Murray. A. Two leaves, abaxial view. B. Flowerbud and abaxial view ofmonoearp, the latter showing beginning dehiscence with single seed visible. C. Single monoearp in lateral view, resting on adaxial surface ofa leaf (Photos: N.A. Murray) mm oblanceolate, flat, apex obtuse. Carpels 16-20; ovaries 1.5-2.5 long, narrowly oblong, densely fermginous-pubeseent with hairs obseuring lower portion of stigmas; mm stigmas loosely connivent, e. 2.2 long, dark, with a few seattered hairs and mm studded with amber-eoloured papillae. Torus flat, c. 3.4 in diameter. Pedicel of mm mm mm fruit 11-20 long, 4-7 thick, pubescent; torus of fruit c. 8 high, 9-14 mm in diam., depressed-globose, sparsely pubescent to glabrate. Monocarps red with brown tomentum in vivo, up to 20 per fruit, 5.5-10.7 cm long, 0.6-1.1 cm wide and thick, linear to narrowly oblong, weakly torulose, terete in cross-section, apex rostrate, mm mm mm the beak 2.5-9 long, base contracted into a stipe 8-10 long, 3.5^ thick, mm longitudinally wrinkled, ferruginous-pubescent to glabrate; pericarp 1.7 thick. Seeds in a single row, parallel to long axis ofmonoearp, 6-12 per monoearp, 7.2-8. mm mm mm long, 5.5-5.6 wide, 4.7-5. 1 thick, ellipsoid, elliptic in cross section, dark brown, rugose, flattened or a little concave at micropylar end, rounded at chalazal end, raphe and antiraphe distinctly raised; aril and aril plate absent. Distribution. Occurs in Terengganu in northeastern Peninsular Malaysia, and on the northern coast of the island of Borneo in Sarawak (East Malaysia) and in Brunei. Considering its restricted habitat and the pace ofdevelopment in Sarawak, this species Xylopia in SoutheastAsia 365 & Fig. 2. Xylopia eiythrodactyla D.M.Johnson N.A.Murray. A. Habit. B. Inflorescence with flower buds, side view. C. Seed, view of micropylar end. D. Seed, side view. E. Fruit. F. Outer petal, adaxial view. G. Inner petal, adaxial view. H. Stigma apex. I. Carpel. J. Stamen. K. Close-up of flower with petals and stamens removed, to show sepals, caipels and torus. L. Schematic side view of flower at anthesis, one outer petal removed. Drawn by Kate Ball from (A) Awa S. Ismawi S.47080, ASU; (B) Sibat ak Luang S.24502, L; (C-E) Chew & Kiah SFN.40982, A; (F, G, K) Zehnder S.16803, A; (H-J) Rogstad 704, A; and (L) field sketch. 1 366 Gard. Bull. Singapore 67(2) 2015 may be more vulnerable than it appears from the collecting data. Ecology. The bulk of the Sarawak and Brunei collections have come from either lowland peat swamp forest or heath (kerangas) forest (tenninology following Saw, 2010). The collection localities in Terengganu are all from lowland dipterocarp forest, as are several in Sarawak. In general, however, the ecological range ofthis species is much narrower than that ofXylopiafermginea. Elevational range is sea level to 250 m, with one collection from Terengganu from c. 800 m. Phenology. The species shows two distinct flowering and fruiting periods throughout its distribution: flowers have been collected in late March and May-July, and then again in November-December, while fruits have been collected from February to July and also from September to December. Local name. Ako (Sarawak: Bojeng bin Sitam 9319, Zainuddin bin Bolhassan S.11941). Etymology. The species is named for the red and finger-like monocarps (Fig. IB-C), which are thicker and less torulose than those ofits congonQiXylopiafermginea. Additional specimens examined. MALAYSIA: Peninsular: Terengganu: 18th mile, Jalan Kelantan, 31 Mar 1957 (fr), Chiew & Kiah SEN40982 (A, K, L [4 sheets], M, SING [2 sheets]); Pulau Redang, Pasir Mah Kapit, 300 m, 5 Mar 1989 (fr), Saw FRl 36487 (A, K, KEP, L, SAR, SING); near Kampong Gong Nangka, Marang, 6 Jul 1 953 (st), Sinclair & Kiah bin Salleh SEN 39837 (L, SING [3 sheets] 19th mile Kuala Trengganu Besut road, 15 Nov 1954 (st), Sinclair ); & Kiah bin Salleh 40473 (L, SING); 8th mile Kuala Trengganu-Besut Road (west side), 7 Sep 1 1955 (young fr), Sinclair & Kiah bin Salleh SEN40748 (A, K, L, SING [2 sheets]); Ulu S. Loh below E face G. Mandi Angin, 2300 ft., 13 Jul 1968 (If), Whitmore FRI 12144 (K, KEP, L); Borneo: Sarawak: TelokAsam, Bako N. R, 4 Jun 1963 (W),Ashton S.17913 (A, ASU, K, KEP, L, SAR, SING); Ulu Kenyana, Mukah, 20 Oct 1963 (young fr), Ashton S. 19488 (A, K, KEP, L, SAR, SING); Kuching, Setapok F.R., 24 Jul 1957 (fl), Bojeng bin Sitam 9319 (BO, K, L, SAR, SING); G. Pueh F.R., June 1956 (st), BrunigS.6369 (SING); Selang F.R., Feb 1956 (st), Brunig S. 7225 (SING); long path from Tg. Po to Telok Kruim, Bako National Park, Kuching, 1st div., 15 m, 16 May 1980 (fr). Citing S.42286 (K, KEP, L, SAR); Ulu Sungai Pasir Biawak, Lundu, Kuching, 8 Apr 1997 (if), Jamree et al. S. 76728 (K, KEP, L, SAR); Kem Pennai, Santubong, 29 Jun 1992 (fr), Othmcin et al. S.65133 (KEP, L); Sampadi F.R. (near road), 25th mile, Bau/ Lundu Road, 1st Division, 750 ft, 28 Jun 1968 (fl). Pale S.26915 (K [2 sheets], L, SAR, SING); Bako National Park,just above mangrove swamp at open area on slope at start ofJalan Lintang, 6 Dec 1981, Rogstad 703 (Afr), 704 (Afl); Nyabau Catchment area, Bintulu, 4th Division, 300 ft, 11 Jun 1966 (fl), Sibat ak Luang S.24502 (BO, KEP, L, SAR, SING); Kuching, 100’ alt., Setapok F.R., 15 Nov 1957 (fi-), Yacup 8939 (K, L, SAR, SING); Bintulu, Similajau F.R., 17 Nov 1959 (fl), Zainuddin bin Bolhassan S. 1194 (K, L, SAR, SING); Kuching, Sg. Teruntum, Sarawak Mangrove Reseiwe, 27 May 1962 (fl), Zehnder S.16803 (A, K, L, SAR, SING). BRUNEI: Andulau F.R., 18 Sep 1957 (st), Ashton BRUN 560 (L, SING); Belait District, Ulu Sungai Badas, 28 Mar 1989 (fl), Nangkat NN105 (A, AAU, K, L, SAR, SING); Belait District, Lumut Hills, 30 Mar 1968 (st), van Niel 4454 (L); Ulu Badas, Andulau Stateland, Sg. Liang, Belait (Labi Rd. 10 km), 25 Jun 1996 (fl), Ogata et al. Og-B203 (L). 367 Xylopia in SoutheastAsia Notes. Trees of this species were observed in Bako National Park in June of 2003, where the plants were frequent along the edge of dipterocarp forest on a slope bordering mangroves. At this site there were many seedlings and saplings present; m flowers or fruits were only found on individuals exceeding 15 in height. The petals offallen flowers were sweetly scented, the scent reminiscent ofGardenia (Rubiaceae). At anthesis the outer petals were widely spread while the inner petals were more or less erect (Fig. 2L). Dehiscing monocarps, gathered from the ground at this locality, were photographed (Fig. IB). The dried seeds, when soaked, become tan in colour, revealing a sarcotesta c. mm 0.2 thick, which may be somewhat incomplete at the chalazal end. The sarcotesta breaks away in chunks from the woody layer of the seed coat underneath, often with patches ofthe woody coat adhering to it. The type description ofXylopia altissima BoerL, based on a specimen collected by Teysmann on the island of Lingga (Boerlage, 1 899), suggests the new species in its emphasis on the dense tomentum of the abaxial leaf surface, but Boerlage (p. 203) also emphasised the strongly reticulate leaves (“venis tenuibus in nervos tiansversis et reticulatis pertensa”) with acuminate apices. These features are frequent in Sumatran specimens ofXylopiaferniginea s.s., which in general have larger and broader leaves than those ofPeninsular Malaysian and Bornean plants, as well as a tendency toward a subcordate rather than truncate leafbase. These leaffeatures do not occur inXylopia erythrodactyla. M Two sterile specimens found in the herbarium at bear leaves resembling those of Xylopia e/ythrodactyla. The specimens were collected from a plant grown in the Bogor Botanic Garden. The provenance of the plants is not certain, but “Borneo” is written in pencil on one ofthe sheets. The actual labels give the following information: Hort. Bogor. IV B 1 8, 2 May 1 895, Spelta [?] s.n. (M [2 sheets], as “Sapotacea Sambas V. d. Florsf’ on one sheet, “Columnifera?” on the other). Duplicates ofthese specimens were not seen at L or BO; the catalogue by Dakkus (1957) did not show any listings under either the names or the number. Xylopia sumatrana (Miq.) D.M.Jolmson & N.A.MuiTay, comb. nov. - Unona sumatrana Miq., FI. Ned. Ind., Eerste bijv. 3: 377 (1861). -Xylopia malayana Hook.f. & Thomson var. macrocarpa BoerL, Icon. Bogor. 1(2): 123 (1 899). -TYPE: Indonesia, Sumatra occid. in prov. Priaman, Diepenhorst s.n. (holotype U). Xylopia stenopetala Oliv., Hook. Icon. PL 21: 1563 (1887). - TYPE: Malaysia, t. Penang, Government Hill, 600 ft, June 1886, Curtis 857 (lectotype K, designated by Turner (2011); isolectotype SING). Distribution. Peninsular Malaysia, Sumatra, Borneo, Philippines (Mindanao). Additionalspecimensexamined(representativespecimens). MALAYSIA: Peninsular: Pahang: Frasers Hill at the lower gate, 3000’ alt, 29 May 1968 (fl, fr), Ng FRl 6172 (A, K, SING); G. 368 Gard. Bull. Singapore 67(2) 2015 Benom Game Reserve, Ulu Ki'au, 1800’ alt, 22 Apr 1967 (fl), YusoffKEP 99124 (A, K, SING); Penang: Tunnel road, Penang Hill, 2200 ft, 28 May 1938 (fr), Henderson SFN 21425 (SING [2 sheets]); Selangor: Kepong, F.R. Inst., 50th mile. Gap Road, 15 April 1990 (fl), Kochummen FRl 29090 (A, K, SAR); Ulu Gombak F.R., 1800 ft alt, 18 Jan 1966 (fr), Kochummen FRl 80497 (A, K, SING); Borneo: Sabah: Lamag District, close to exit stream ofGunong Lotung, lake Inarat, 1200 ft, 21 May 1976 (ft), Cockburn SAN 83321 (K, L, SAR, SING); Tawau Dist., Mile 26, Apas Rd, 120 ft, 25 Jun 1959 (If), Meijer SAN 19321 (K, L, SING); Sarawak: Pk in Kuching, Dec 1892 (fl), Haviland =10[?] (BM, L, MO, SING); Perkulen Ampat, b.p.m.d. (fl, fir), Haviland 142/10 [A.13.9] (SAR, SING); Bkt. Lobang, Punung Lusong, Sg. Linau, Belaga, 7th division, 470 m, 14 Jun 1 979 (fr), Lee S.39802 (ASU, K, L, SAR); path from Kpg. Seropak to Bungoh Range, Bau, secondary forest on hillside, c. 1000 ft, 29 Nov 1969 (fl), Paie & Mamit S.29584 (A, K, L, SAR, SING), INDONESIA: Borneo: East Kalimantan: Meratus, 5230 area PT. ITCI, Kenangan Balikpapan, 70 m, 27 Sep 1992 (fr), AmhriansyahA. A:611 (A, K, L); PT. ITCI, road Kenangan to G. Meratus, than to Basecamp Birawa, km 52, 500 m, 28 Mar 1995 (fl), Kessler et al. PK.949 (A, K); E Kutei, Sangkulirang subdivision, Sg. Susuk region, 10 m, 26 Jun 1951 (fl), Kostermans 5456 (A, BM, K, L [2 sheets], SING); West Kalimantan: G. Bentuang area, 5-10 km N ofMasa village, 150 km NE ofPontianak, steep ridge above Semawang River, 00°52’N, 110°26’E, 50 m, 23 Jun 1989 (fl, fr), Burley et al 2823 (A [2 sheets], F, K, L, MO, NY [2 sheets], SING); Sumatra: Aceh: Gunung Leuser Nature Resei^ve, Gunung Bandahara c. 6 km NE of Kampung Seldok (Alas Valley), c. 25 km N of Kutatjane, c. 800-1000 m alt, 20 Mar 1975 (fl, fir), de Wilde & de Wilde-DuyJjes 15599 (K [2 sheets], MO, US); North Sumatra: East Coast, vicinity ofLoemban Ria, Asahan, 5 Feb-12 Apr 1934 (st), Rahmat si Boeea 8096 W (A, MICH, US); East Coast, Asahan NE ofTamuan Delok and of Salabat, 500 m, 15 Jun-9 Jul 1936 (fl), Rahmatsi Boeea 9269 (A [2 sheets], K, L, MICH, NY, US, W). PHILIPPINES: Mindanao: Surigao Sur: Manobo District, PICOP Bislig, Apr 1976 (fl), Rojo 325 (MO). Notes. The type specimen of Unona sumatrana Miq. from the Utrecht herbarium (now housed at L) includes two separate branches, one with a fruit attached, and the other with two small leaves. The two leaves are slightly obovate-oblong, one 6.5 cm long and 3 cm wide and the other 3.9 cm long and 2.3 cm wide. The leaves are retuse at the apex and broadly cuneate and decurrent at the base. The fruit shows the distinctive characteristics of Xylopia stenopetala: there are two monocarps and a portion of a third on the specimen, none ofthem fully mature. The intact monocarps are naiTowly oblong, one 6.8 cm long and 0.8 cm wide and slightly torulose, sparsely pubescent, with about 7 seeds arranged in a single row. The monocarps are acute at the apex and mm mm taper to a stipe c. 8 long. The monocaips are borne on a pedicel 10 long and mm mm mm 4.3 thick, and a torus 1 1 in diameter, 8 high, and depressed-globose in shape. Although the name Unona sumatrana was reduced to a variety of Xylopia A malayana by Boerlage (1899), and placed in synonymy under malayana by Turner (2011), the fmit on the type specimen of Unona sumatrana clearly distinguishes it fromA. malayana: the narrow monocarps have up to 7 seeds (seeds usually 3 or fewer A per monocaip in malayana), and, at 6.8 cm in length, are already longer than those A found in malayana (which never exceed 4 cm in length). 369 Xylopia in SoutheastAsia The type ofthe name Unona sumatrana from 1861 was thus found to represent the same species as Xylopia stenopetala from 1887 and the earlier name must take precedence. This name is not to be confused with Xylopicrum siimatranum (Miq.) Kuntze, Revis. Gen. PI. 8, 1891, which is based on Parartabotrys sumatranus Miq., now considered a taxonomic synonym ofXylopia malayana (Sinclair, 1955; Turner, 2011 ). Xylopia sumatrana has been collected in Peninsular Malaysia in the states of Pahang, Penang and Selangor, and on the islands of Sumatra, Borneo, and Mindanao (Philippines), where it is found at elevations of 50-1000 m, the widest elevational range ofany Asian species ofXylopia. Xylopia malayana group Members of Xylopia malayana group lack stilt roots (with the exception ofXylopia mm sessiliflora treated below), the flower pedicels are 5 long or less, the receptacle bears a distinct but low and irregularly laciniate staminal cone fonned from the connate bases ofthe filaments, the stamens are clavate with a transversely flattened apex to the anther connective, the stigmas lack papillae, and the seeds are smooth. Within this group the monocarps tend to be relatively few in number (ten or fewer, but sqqXylopia heterotricha below) and relatively broad and oblong. Species of the Xylopia malayana group are distinctive and well circumscribed We with the exception ofX. elliptica and X. malayana. found that the primary source oftaxonomic difficulty with these two species stemmed from the inclusion ofmultiple distinct taxa under the name Xylopia elliptica. The simplest route to clarification is to retrace the taxonomic history ofthat species. & Hooker Thomson (1872: 86) originally Xylopia elliptica on a single Maingay collection from Malacca. The protologue, reproduced verbatim below, shows that the diagnosis emphasised the glabrous branches, the small elliptic, obtuse, membranous, glabrous leaves and the solitary flowers: 14. X. elliptica, Maingay niss,\ branches glabrous, leaves small elliptic obtuse mem- branous glabrous, tip rounded, neives faint reticulate, flowers small solitary erect pubescent, sepals subacute united to the middle, ovaries 1-3. Malacca, Maingay. A lofty tree; trunk thick; branches glabrous, almost black; branchlets pubescent. Leaves 1 1/2-2 by 1 %-l 14 in., base obtuse or acute, pale on both surfaces, browner beneath; petiole 14 -% in., puberulous. Flowers 14 in., slender; peduncle halfas long or shorter, and calyx rusty-pu- bescent; bratcs [sic] median, minute. Petals pale brown-tomentose; outer linear-subulate, from a rather broad base, concave; inner trigonous, base excavated. Stamens minute. Ovaries sunk in the deeply urceolate torus, hidden amongst long white hairs; ovules 4-6. The circumscription ofthe species was enlarged by King (1 892), who identified two additional specimens, Wray 3194 from Perak and Curtis 2482 from Penang, as belonging to this species. The latter specimen in particular departed from Hooker and 370 Gard. Bull. Singapore 67(2) 2015 Thomson’s protologue, however, in having distinctly pubescent leaves and multiple flowers per inflorescence home on longer pedicels. Ridley (1922) subsequently maintainedKing’s concept ofthe species mtact. Sinclair(1955) identifiedfour additional specimens as Xylopia elliptica: Awang 42444 from Kedah, Wray 3562 from Perak, 20309, without collector’s name, from Pahang, and Yeob 5037 from Selangor. Sinclair stressed the significance of the pubescence of twigs and leaves as a characteristic important in distinguishing this species from the otherwise similar Xylopia malayana, and apparently on this basis identified a number ofspecimens ofXylopia Ifom Sarawak and Sabah in various herbarium collections as belonging to X. elliptica. It has become clear from re-examination of these specimens, plus study of a wider range of material than was available to these authors, Xylopia elliptica in the sense ofSinclair is a mixture offour very distinctive species. The concept adopted here is one ofXylopia elliptica in a restricted sense, and the recognition ofthree previously undescribed species to accommodate the variants. & - Xylopia elliptica Maingay ex Hook.f Thomson, FI. Brit. India 1: 86 (1872). & Xylopicrum ellipticum (Maingay ex Hook.f. Thomson) Kuntze, Revis. Gen. PI. 1:8 (1891). - TYPE: Malaysia, Malacca, 9 May 1867, Maingay 2376 [Kew Distribution K no. 82] (lectotype [2 sheets, barcode nos. K000574709, K000574712], designated by Turner (201 second-step lectotype, designated here: K000574709; isotype CAL). 1); Xylopia malayana var. obscura Kochummen, Gard. Bull. Singapore 26(1): 49 (1972). - TYPE: Malaysia, Terengganu, Gunong Padang Expedition, Ulu Brang, camp 1 nr. K. Lallang, 1000’ alt, 15 September 1969 (fl, fr), Whitmore 12594 (holotype KEP; isotypes A, K, L, SING). Distribution. Peninsular Malaysia. Although it has been collected from four states in Peninsular Malaysia, Xylopia elliptica is known from only eight collections, all from m forests below 300 in elevation. Additional specimens examined. MALAYSIA: Peninsular: Johor: Compt 10, Rengam F.R., 14 Nov 1966 (fl), Kochummen FRI 2188 (A, K, KEP, L, SING); Compt 34, Gunong Arang F.R., 13 Sep 1969 (fr), Kochummen FRI2761 (K, KEP, L); Keluang, 20 Nov 1990 (fl, fl-), Teo & Remy KL 3968 (KLU); Hutan Simpan Endau, 23 Oct 1997 (fl), Teo & Tetu KL 4951 (KEP); Pahang: Chini Forest Reserve, 11 Dec 2008 (fl), Khairil bin Mahmoud et al, s.n. (UKMB); KEP without definite locality [Raub, according to Sinclair (1955)], collector unknown, 20309 (KEP). Notes. Xylopia elliptica in the sense of its type applies to relatively small-leaved plants with glabrous to sparsely pubescent leaves, inflorescences of a single flower, and relatively short petals. This same circumscription also includes the taxon Xylopia malayana var. obscura.Xylopia elliptica in this restrictedsense is endemic to Peninsular Malaysia, where it is infrequent in lowland forests up to an elevation of300 meters in Johor, Melaka, Pahang, and Terengganu. The report ofXylopia elliptica for southern