Synopsis of Nekemias Raf., a segregate genus from Ampelopsis Michx. (Vitaceae) disjunct between eastern/southeastern Asia and eastern North America, with ten new combinations.

The genus Nekemias (Vitaceae) was first recognized by Rafinesque in 1838. It has been treated as a synonym of Ampelopsis Michx. Recent phylogenetic studies suggest that Ampelopsis as traditionally delimited is paraphyletic. To maintain the monophyly of each of the genera of Vitaceae, we herein segregate the Ampelopsissect.Leeaceifoliae lineage from Ampelopsis and recognize these taxa in Nekemias Raf., which has a disjunct distribution in eastern to southeastern Asia and eastern North America. Nomenclatural changes are made for nine species and one variety: Nekemiasarborea (L.) J. Wen & Boggan, Nekemiascantoniensis (Hook. & Arn.) J. Wen & Z.L. Nie, Nekemiascelebica (Suess.) J. Wen & Boggan, Nekemiaschaffanjonii (H. Lév. & Van.) J. Wen & Z.L. Nie, Nekemiasgongshanensis (C.L. Li) J. Wen & Z.L. Nie, Nekemiasgrossedentata (Hand.-Mazz.) J. Wen & Z.L. Nie, Nekemiashypoglauca (Hance) J. Wen & Z.L. Nie, Nekemiasmegalophylla (Diels & Gilg) J. Wen & Z.L. Nie, Nekemiasmegalophyllavar.jiangxiensis (W.T. Wang) J. Wen & Z.L. Nie, and Nekemiasrubifolia (Wall.) J. Wen & Z.L. Nie. A taxonomic key is provided for the genus to facilitate identification.

Introduction

Michx. () is one of the 15 recognized genera in with about 25 species (Suessenguth 1953; Chen et al. 2007; Wen 2007; Wen et al. 2013a). The genus was shown to be paraphyletic by recent phylogenetic analyses based on plastid (Soejima and Wen 2006; Ren et al. 2011; Nie et al. 2012) and nuclear GAI1 (Wen et al. 2007) analyses. The plastid data (Soejima and Wen 2006; Ren et al. 2011; Nie et al. 2012) supported that the African and the South American complex formed a clade with the simple or palmately leaved (sect. ), while the nuclear data (Wen et al. 2007) suggested that they were more closely related to the pinnately leaved (sect. , as designated by Galet 1967 in the unpublished thesis). All analyses so far have supported the monophyly of each of the two sections based on leaf morphology (Galet 1967). Apart from the differences in leaf morphology, the two sections also differ in their axillary buds, with taxa in sect. having serial accessory buds, and those in sect. having complex axillary buds as in (Bernard 1972–1973; Gerrath and Posluszny 1989; Soejima and Wen 2006; Wen et al. 2007).

Within , five or six main clades are well supported based on analysis of molecular sequence data with the -- clade as one of these major clades of (Soejima and Wen 2006; Wen et al. 2007, 2013b; Ren et al. 2011; Liu et al. 2013). Ingrouille et al. (2002), however, resolved as the basalmost branch of albeit with no support, and further argued that the presence of pinnate leaves, the thick corolla, and the floral and vegetative development in were the least-derived characters within as compared with those in the outgroup taxa from . To maintain the monophyly of each of the genera of , we herein segregate the lineage from . Rafinesque (1838) established the genus with Michx. [= (L.) Koehne] possessing pinnately compound leaves as the type species. has been rarely mentioned in subsequent taxonomic work on or treated as a synonym of (Merrill 1949; Suessenguth 1953; Wen 2007), but it represents the earliest name at the generic rank for the pinnately-compound leaved clade.

Taxonomic synopsis

Raf., Sylva Tellur. 87. 1838.

Michx., pro parte

Woody climbers.

Branchlets with prominent lenticels. Pith white, continuous through nodes. Tendrils leaf-opposed, mostly bifurcate to sometimes trifurcate, and lacking adhesive discs. Leaves alternate, petiolate, stipulate, pinnately to ternately bipinnately or sometimes tripinnately compound. Inflorescences bifurcately compound cymes, long peduncled, leaf-opposite. Flowers pedicellate, mostly bisexual; calyx saucer-like; corolla of 5 thick petals; stamens 5, opposite to petals; disc adnate to the base of the PageBreakovary; ovary 2-locular, style short, conical, stigma rounded. Fruit a berry, globose or subglobose, purple, blue or black, 1-4 seeded. Seeds obovoid.

Type species.

(Michx.) Raf. [= (L.) J. Wen & Boggan].

Nine species with eight occurring in warm temperate to tropical areas of eastern and southeastern Asia (Suessenguth 1940; Galet 1967; Chen et al. 2007), and one species distributed in eastern North America extending to the Caribbean (Brizicky 1965). This intercontinental disjunct distribution between eastern Asia and eastern North America represent a classical biogeographic pattern of the Northern Hemisphere (Wen 1999; Wen et al. 2010).

Below we provide a taxonomic synopsis for the genus.

(L.) J. Wen & Boggan comb. nov.

urn:lsid:ipni.org:names:77142315-1

Figure 1A–B

Figure 1.

Images of representative species of Raf. A–B (L.) J. Wen & Boggan, voucher specimen: J. Wen 12005 (US), collected from Montgomery Co., Texas, USA C–D (Hook. & Arn.) J. Wen & Z.L. Nie, voucher specimen: J. Wen 10613 (US), collected from Xichou Xian, Yunnan province, China E–F (Suess.) J. Wen & Boggan, voucher specimen: J. Wen 10242 (US), collected from SE Sulawesi, Indonesia.

L., Sp. Pl. 1: 203. 1753. Basionym

Michx., Fl. Bor.-Amer. 1: 160. 1803, nom. illeg.

Pers., Syn. Pl. 1: 143. 1805, nom. illeg.

(Michx.) Nutt., Gen. N. Amer. Pl. 1: 144. 1818, nom. illeg.

(Michx.) Raf., Sylva Tellur. 87. 1838, nom. illeg.

(Michx.) Torrey & A. Gray, Fl. N. Amer. 1: 243. 1838, nom. illeg.

(L.) Des Moulins in Durand, Actes Soc. Linn. Bordeaux 24: 156. 1862.

(L.) Koehne, Deutsch. Dendrol. 400. 1893.

(L.) Rusby, Mem. Torrey Bot. Club 5: 221. 1894, comb. superfl.

Distribution.

USA (Alabama, Arkansas, Florida, Georgia, Illinois, Indiana, Kentucky, Louisiana, Mississippi, Missouri, New Mexico, North Carolina, Ohio, Oklahoma, South Carolina, Tennessee, Texas, Virginia and West Virginia) and the Caribbean.

(Hook. & Arn.) J. Wen & Z.L. Nie comb. nov.

urn:lsid:ipni.org:names:77142316-1

Figure 1C–D

Hook. & Arn., Bot. Beech. Voy.: 175. 1833. Basionym

Walp., Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 19 (Suppl. 1): 314. 1843, nom. illeg., non DC. 1824.

(Hook. & Arn.) K. Koch, Hort. Dendrol. 48: 11. 1853.

Maxim., Mel. Biol. Acad. Sci. St. Petersb. 9: 148. 1873.

(Hook. & Arn.) Seem., Bot. Voy. Herald: 370. 1875.

(Hook. & Arn.) Planch., Monogr. Phan. 5: 460. 1887, comb. superfl.

Planch., Monogr. Phan. 5: 460. 1887.

(Maxim.) Planch. [epithet published in error as “

Hort. Mazel. ex Planch., Monogr. Phan. 5: 461. 1887, nom. nud. pro syn.

H. Lév. & Vaniot, Bull. Soc. Agric. Sci. Arts Sarthe 40: 41. 1905.

H. Lév., Repert. Spec. Nov. Regni Veg. 8: 58. 1910.

Gagnep., Bull. Soc. Bot. France 92: 166. 1946.

(Maxim.) F.Y. Lu [published in error as “

China (Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hong Kong, Hubei, Hunan, Taiwan, Xizang, Yunnan and Zhejiang), India, Japan (including the Ryukyu Islands), Vietnam, Laos, Malaysia (peninsular), and Indonesia (Java).

(Suess.) J. Wen & Boggan comb. nov.

urn:lsid:ipni.org:names:77142317-1

Figure 1E–F

Suess., Repert. Spec. Nov. Regni Veg. 49: 14. 1940. Basionym

Indonesia (Sulawesi).

(H. Lév. & Van.) J. Wen & Z.L. Nie comb. nov.

urn:lsid:ipni.org:names:77142323-1

H. Lév. & Van., Bull. Soc. Agric. Sci. Arts Sarthe 40: 37. 1905. Basionym

H. Lév., Repert. Spec. Nov. Regni Veg. 8: 58. 1910.

H. Lév., Repert. Spec. Nov. Regni Veg. 9: 457. 1911.

E.H. Wilson, J. Roy. Hort. Soc. 42: 37. 1916, nom. nud.

(E.H. Wilson) Bean, Trees & Shrubs Brit. Isles 2: 673. 1921.

(H. Lév. & Van.) Rehder, J. Arnold Arbor. 15: 25. 1934.

China (Anhui, Chongqing, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan and Yunnan).

(C.L. Li) J. Wen & Z.L. Nie comb. nov.

urn:lsid:ipni.org:names:77142318-1

C.L. Li, Chinese J. Appl. Environ. Biol. 2(1): 48. 1996. Basionym

China (Yunnan).

(Hand.-Mazz.) J. Wen & Z.L. Nie comb. nov.

urn:lsid:ipni.org:names:77142325-1

Hand.-Mazz., Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Cl., Abt. 1, 59: 105. 1877. Basionym

(Hand.-Mazz.) W.T. Wang, Acta Phytotax. Sin. 17(3): 79. 1979.

China (Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi and Yunnan).

(Hance) J. Wen & Z.L. Nie comb. nov.

urn:lsid:ipni.org:names:77142319-1

Hance, Ann. Bot. Syst. 2: 724. 1852. Basionym

(Hance) C.L. Li, Chinese J. Appl. Environ. Biol. 2(1): 48. 1996.

China (Fujian, Guangdong, Hong Kong and Jiangxi).

(Diels & Gilg) J. Wen & Z.L. Nie comb. nov.

urn:lsid:ipni.org:names:77142320-1

Diels & Gilg, Bot. Jahrb. Syst. 29: 466. 1900. Basionym

(Diels & Gilg) Börner, Abh. Nat. Ver. Bremen 21: 280. 1913.

China (Chongqing, Gansu, Guizhou, Hubei, Shaanxi, Sichuan and Yunnan).

(W.T. Wang) J. Wen & Z.L. Nie comb. nov.

urn:lsid:ipni.org:names:77142321-1

W.T. Wang, Bull. Bot. Res. North-East. Forest. Inst. 1(1–2): 170. 1981. Basionym

(W.T. Wang) C.L. Li, Chinese J. Appl. Environ. Biol. 2(1): 48. 1996.

China (Jiangxi).

(Wall.) J. Wen & Z.L. Nie comb. nov.

urn:lsid:ipni.org:names:77142322-1

Wall., Fl. Ind., ed. Carey & Wall., 2: 480. 1824. Basionym

(Wall.) Planch., Monogr. Phan. 5: 463. 1887.

W.T. Wang, Acta Phytotax. Sin. 17(3): 79, 90. 1979.

China (Guangxi, Guizhou, Hunan, Jiangxi and Sichuan), and India.

1	Leaves abaxially strongly glaucous	2
1	Leaves green on both surfaces	3
2	Lower leaves pinnately compound, leaflet blades 7–15 × 3–7 cm	Nekemias chaffanjonii
2	Lower leaves bipinnately compound, leaflet blades 2.5–6 × 1–3.5 cm	Nekemias hypoglauca
3	Leaves pinnately compound	4
3	Leaves bipinnately, ternately bipinnately to tripinnately compound	5
4	Leaflets 3.5–14 × 2–6.5 cm, margin 5–15-toothed, abaxially densely ferruginous pilose; berries 8–15 mm in diameter	Nekemias rubifolia
4	Leaflets 3–6 × 0.5–3 cm, margin entire or with 1 to several inconspicuous teeth, midvein abaxially sparsely pilose; berries 5–7 mm in diameter	Nekemias gongshanensis
5	Tendril trifurcate; leaflets 4–12 × 2–6 cm	Nekemias megalophylla
5	Tendril bifurcate; leaflets 1–5 × 0.5–2.5 cm	6
6	Leaflet margin with 2–4 large coarse teeth; from North America or the Caribbean	Nekemias arborea
6	Leaflet margin serrate with 5–15 teeth on each side; from Asia	7
7	Leaflet margin coarsely serrate, central leaflet ovate-elliptical	Nekemias grossedentata
7	Leaflet margin ± undulate, central leaflet obovate or ovate	8
8	Leaves and inflorescences pilose to glabrescent	Nekemias cantoniensis
8	Leaves and inflorescences pubescent to densely so	Nekemias celebica