A Synopsis of Girardinia (Urticaceae)

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A Synopsis of Girardinia (Urticaceae) Author(s): Ib Friis Reviewed work(s): Source: Kew Bulletin, Vol. 36, No. 1 (1981), pp. 143-157 Published by: Springer on behalf of Royal Botanic Gardens, Kew Stable URL: http://www.jstor.org/stable/4119014 . Accessed: 28/09/2012 12:36 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. . Royal Botanic Gardens, Kew and Springer are collaborating with JSTOR to digitize, preserve and extend access to Kew Bulletin. http://www.jstor.org

Transcript of A Synopsis of Girardinia (Urticaceae)

Page 1: A Synopsis of Girardinia (Urticaceae)

A Synopsis of Girardinia (Urticaceae)Author(s): Ib FriisReviewed work(s):Source: Kew Bulletin, Vol. 36, No. 1 (1981), pp. 143-157Published by: Springer on behalf of Royal Botanic Gardens, KewStable URL: http://www.jstor.org/stable/4119014 .Accessed: 28/09/2012 12:36

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp

.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact [email protected].

.

Royal Botanic Gardens, Kew and Springer are collaborating with JSTOR to digitize, preserve and extendaccess to Kew Bulletin.

http://www.jstor.org

Page 2: A Synopsis of Girardinia (Urticaceae)

A synopsis of Girardinia (Urticaceae) IB FRIIS*

Summary. The genus Girardinia has been found to comprise two species, both restricted to the montane areas in the tropics and in the warmer temperate parts of the Old World: G. bullosa is a well defined species restricted to Ethiopia and East Africa; G. diversifolia is a polymorphic species distributed from West Africa to China, Korea, Taiwan and Indonesia. G. cuspidata is found to be a Laportea, and the necessary new combination is made.

In connection with my study of African Urticaceae for the Flora of Tropical East Africa it has been necessary to study the genus Girardinia in some detail. It has long been considered difficult to name specimens of this genus with any degree of certainty, but the latest account of the genus (Letouzey 1968) suggested that only two species existed in Africa: the East African G. bullosa, and G. heterophylla, a very widespread species which occurred from West Africa through tropical Africa, Arabia and India to China and Mainland SE Asia. It appeared, however, from the synonymy given for G. heterophylla by Letouzey that the name for this species was illegitimate, and that some other epithet, probably from an Asian taxon, would have to be used instead. A study of the whole genus seemed inevitable in order to solve this problem, and material from the following herbaria was obtained on loan: A, AAU, B, BM, BR, C, CAL, COI, E, FI, G, K, L, LE, LISC, LISU, MH, MO, P, PDA, SING, TAIF, UPS, W, WU. The genus seems to be avoided by collectors because of the fiercely stinging hairs, and the total amount of material preserved in herbaria is comparatively scarce in view of the wide distribution. A total of c. 400 collections have been seen.

The study confirmed the basic findings of Letouzey, but it was also found that all hitherto described species of Girardinia, with the exception of the above mentioned G. bullosa and two species which had to be excluded from the genus, were in fact connected by intermediate forms. The only satis- factory solution to this taxonomic problem was to merge all these species into one and, in fact, species with such a wide range of distribution are well known from the Urticaceae. Laportea interrupta (L.) Chew, Laportea aestuans (L.) Chew, Boehmeria platyphylla D. Don, and Parietaria debilis G. Forster are examples of species with an equally wide distribution, and the latter three species are almost as polymorphic.

The nomenclature was found to be quite complicated, with a long range of illegitimate names, and it seems inevitable that a new combination based on the oldest available epithet, Urtica diversifolia, has to be made. More detailed notes on nomenclature and taxonomy of this species are given below.

Girardinia Gaudichaud in Freycinet, Voy. Monde Uran., Bot.: 498 (1830, '1826'); Endlicher, Gen. Pl.: 283 (1837); Benth. & Hook. f., Gen. P1. 3(1): 384 (1880); Engler in Engler & Prantl (eds.), Nat. Pflanzenf. 3(1): 107

Accepted for publication August 1980. * Institute of Systematic Botany, University of Copenhagen, 140, Gothersgade, DK- 1123 Copenhagen K, Denmark.

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LA?O , , MAP 1. Distribution of Girardinia bullosa.

(1888); Hutch., Gen. P1. 2: 183 (1967). Type: G. leschenaultiana Decne. (lectotype, indicated in Index Nominum Genericorum) = G. diversifolia.

Tall, short-living herbaceous plants covered with long, stinging hairs, monoecious or dioecious by abortion. Leaves alternate, petiolate, elliptic to ovate in outline, but very variably divided, with various degrees of division of the leaves on the same plant, always with three major nerves from the base of the leafblade. Cystoliths dot-like. Stipules intrapetiolar, fused almost to the apex, caducous. Flowers in unisexual, axillary inflores- cences. Male inflorescence ? branched, slender, almost spike-like, with dense clusters of flowers. Female inflorescence ? branched, elongated, sometimes almost spike-like, sometimes contracted, with very dense cymules of flowers and fiercely armed with stinging hairs. Male flowers pedicellate, with 4 or 5 tepals and the same number of stamens, a rudimentary ovary always present. Female flowers apparently sessile, with 3 tepals almost completely fused into a hollow structure ? enclosing the reflexed ovary; the fourth, free tepal is ? rudimentary or lacking. Staminodia absent. Ovary

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laterally compressed, asymmetrical, reflexed, with filiform stigma. Fruit a flattened, broadly ovate or subcordate achene with uniform rugose sculptur- ing throughout. Seeds with very little endosperm, and embryo with broad cotyledons.

The genus belongs to the tribe Urereae, which is the only tribe of the Urticaceae with stinging hairs. It seems to be closely related to the genus Laportea, with which it shares the following characters: herbaceous habit, alternate leaves, intrapetiolar stipules which are fused almost to the apex, winged pedicels in female inflorescence (in Girardinia the branches of the inflorescence are also winged so that the female flowers appear to be sessile on a complex flattened structure), reflexed, laterally compressed ovary and filiform style. Girardinia differs from Laportea chiefly in the compact female inflorescences, the 3 fused tepals (and often one free tepal) in the female flower and the uniformly sculptured achenes.

KEY TO THE SPECIES OF GIRARDINIA

1. Tepals of male flower and largest tepal of female flower with a horn-like appendage on the outside; leaves broadly ovate to subcordate, usually more than 15 cm long, bullate, always with doubly serrate margin; mature stems more than 2 cm diam. ..... . 1. G. bullosa

1. Tepals without horn-like appendages, but the largest tepal of the female flower has a ? developed vertical crest on the outside; leaves elliptic to ovate in outline, entire, 3-, 5- or 7-lobed, usually less than 15 cm long, not bullate, with serrate margin (sometimes doubly serrate, but then usually only in the lower part of the leaf); mature stems less than 2 cm diam.

2. G. diversifolia

1. Girardinia bullosa (Steud.) Weddell in Ann. Sci. Nat., Bot., Ser. 4, 1: 181 (1854); Blume, Mus. Bot. Lugd.-Bat. 2: 160 (1855); Weddell, Monogr. Urt.: 168, Tab. 2, Fig. B, 5-7 (1856) & in DC., Prodr. 16 (1): 102 (1869); Engler, Hochgebirgsfl. Trop. Afr.: 193 (1892); Rendle, in Prain (ed.), Fl. Trop. Afr. 6(2): 265 (1917); Hauman, Fl. Congo Belge 1: 195 (1948); Robyns, Fl. Sperm. P.N.A. 1: 75 (1948); Cuf. in Bull. Jard. Bot. Brux. 23, Suppl.: 18 (1953); Verdcourt in Kew Bull. 21: 261 (1967); Agnew, Upl. Kenya Wild Flowers: 323 (1974). Type: Ethiopia, Tschenausa, Schimper Series II, 1409 (holotype P, Herb. Steudel; isotypes A, BM, G, L, LE, P). Urtica bullosa Steud. in Flora 33: 259 (1850); A. Rich., Tent. Fl. Abyss. 2:

262 (1851).

DISTRIBUTION. Map 1 (Ethiopia, Burundi, Kenya, Tanzania). ILLUSTRATIONS. Figs 1 & 2.

2. Girardinia diversifolia (Link) Friis comb. nov.

Urtica diversifolia Link, Enum. pl. hort. bot. Berol. alt. 2: 385 (1822), non Blume (1825). Type: Plant from India, cultivated in the Berlin Botanical Garden, not preserved. Neotype selected here: India, Sikkim, 5-7000 ft, Hooker s.n. (neotype K; isoneotypes B, C, CAL, L).

Urtica palmata Forssk., Fl. Aegypt.-Arab.: 159 (1775). Type: Yemen, Forsskdl s.n. (holotype C), non Girardiniapalmata Blume. I-L

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FIG. 1. Girardinia bullosa. A habit. B underside of leaf, detail. C upper surface of leaf, detail. D male bud seen from above. E part of male inflorescence. F open male flower with reflexed stamens. A-D from Chiovenda 971; E-F from Lythgoe 308.

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Urtica heterophylla Vahl, Symb. Bot. 1: 76 (1790), nom. illegit. superfl. Type: Yemen, Forsskdl s.n. (C) & Rheede-tot-Drakenstein, Hort. Ind. Malab. 2: Tab. 41 (1679), syntypes.

Urtica horrida Link, Enum. pl. hort. bot. Berol. alt. 2: 385 (1822), nom. illegit., non Humb., Bonpl. & Kunth (1817). Type: Plant from Nepal, cultivated in the Berlin Botanical Garden, not preserved.

Urtica heterophylla D. Don, Fl. Nepal Prodr.: 59 (1825), nom. illegit., non Vahl (1790). Type: Nepal, Wallich probably no. 4603 D (K).

Urtica ferocissima Sweet, Hort. Brit., 2. ed.: 363 (1827). Type: as for Urtica horrida Link.

Urtica acerifolia Zenker, P1. Ind. 1: 2 & Tab. 3-4 (1835). Type: India, Blue Mts (not traced).

Urtica linkiana Heynh., Nomencl. Bot. Hort. 1: 832 (1840), nom. illegit. superfl. Type: as for Urtica horrida Link.

Girardinia heterophylla Decne. in Jacquemont, Voy. Inde 4, Bot.: 151, Tab. 153 (1844); Weddell in Ann. Sci. Nat., Bot., S&r. 4, 1: 181 (1854); Blume, Mus. Bot. Lugd.-Bat. 2: 158 (1855); Weddell, Monogr. Urt.: 164 (1856) & in DC., Prodr. 16(1): 100 (1869); Hook. f., Fl. Brit. Ind. 5: 550 (1888) & in Trimen, Fl. Ceylon 4: 106 (1898); Strachey, Cat. P1. Kumaon, Gharwal & Tibet: 166 (1906); Collett, Fl. Simlensis: 462 (1902); Hayata in Journ. Coll. Sci. Imp. Univ. Tokyo 25(19): 196 (1908); Craib, Contr. Fl. Siam: 197 (1912); Bamber, P1. Punjab: 248 (1916); Duthier, Fl. Gangetic Plain 3(1): 125 (1915); Fyson, Fl. S. Ind. Hill Stat. 1: 544 (1932); Kanjilal & al., Fl. Assam 4: 282 (1940); Cuf. in Bull. Jard. Bot. Brux. 23, Suppl.: 19 (1953); Letouzey, Fl. Cameroun 8: 110 & Tab. 17 (1968); Wickens, Fl. Jebel Marra: 120 (1976). Based on Urtica hetero- phylla Vahl, nom. illegit.

Girardinia leschenaultiana Decne. in Jacquemont, Voy. Inde 4, Bot.: 152 (1844); Weddell in Ann. Sci. Nat., Bot., S&r. 4, 1: 181 (1854) & Monogr. Urt.: 165 (1856). Type: India, Nilgiri Hills, Leschenault 54 (holotype P).

Girardinia zeylanica Decne. in Jacquemont, Voy. Inde 4, Bot.: 152 (1844); Weddell in Ann. Sci. Nat., Bot., S6r. 4, 1: 181 (1854); Blume, Mus. Bot. Lugd.-Bat. 2: 158 (1855); Weddell, Monogr. Urt.: 166 (1856); Miq., Fl. Ind. Bat. 1, 2(2): 233 (1859); Weddell in DC., Prodr. 16(1): 101 (1869); Brandis, For. Fl. N.W. & C. India: 404 (1874); Prain, Bengal P1. 2: 721 (1903); Kanjilal & al., Fl. Assam 4: 282 (1940). Type: Sri Lanka (Ceylon), Koudji, Leschenault s.n. (holotype P).

Girardinia erosa Decne. in Jacquemont, Voy. Inde 4, Bot.: 152 (1844); Weddell in Ann. Sci. Nat., Bot., S6r. 4, 1: 181 (1854) & Monogr. Urt.: 167 & Tab. 2, Fig. 8-9 (1856) & in DC., Prodr. 16(1): 102 (1869); Leandri, Fl. Madagascar 56, Urtic.: 25 & Fig. 5, 6-10 (1964). Type: Madagascar, Mt Ramassina, Bojer s.n. (holotype P not traced; isotype K).

Girardinia armata Kunth, Ind. Sem. Hort. Berol. 1846: 11 (1846) & in Ann. Sci. Nat. Bot., S&r. 3, 7: 184 (1847), nom. illegit. superfl., citing Urtica diversifolia Link, non Blume, Urtica horrida Link, non Humb., Bonpl. & Kunth, and Urtica heterophylla D. Don, non Vahl, as synonyms. Type: Plant from Nepal, cultivated in the Berlin Botanical Garden, not preserved.

Urtica adognsis Steud. in Flora 33: 259 (1850); A. Rich., Tent. Fl. Abyss. 2: 262 (1851). Type: Ethiopia, Adua, Schimper Series I, 101 (holotype P, Herb. Steudel; isotypes A, BM, G, K, LE, LISU, P, S).

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FIG. 2. Girardinia bullosa. A female inflorescence with mature fruits. B detail of A. C female flower. D young female inflorescence. All from Chiovenda 971.

Urtica condensata Steud. in Flora 33: 260 (1850); A. Rich., Tent. Fl. Abyss. 2: 263 (1851). Type: Ethiopia, Mt Selleuda near Adua, Schimper Series III, 1888 (holotype P, Herb. Steudel; isotypes BM, BR, FI, G, K, L, LE, P, S).

Girardinia hibiscifolia Miq., P1. Junghunianae 1: 31 (1851); Blume, Mus. Bot. Lugd.-Bat. 2: 161 (1855); Backer & Bakhuizen, FL. Java 1: 40 (1965). Type: Java; Mt Ged6, Junghuhn s.n., Tankuban-Prahu, Junghuhn s.n., sine loco, Junghuhn s.n. (syntypes all L).

Girardinia adoinsis (Steud.) Weddell in Ann. Sci. Nat. Bot., S&r. 4, 1: 181 (1854); Blume, Mus. Bot. Lugd.-Bat. 2: 159 (1855); Hiern, Cat. Afr. P1. Welw. 1(4): 990 (1900).

Girardinia condensata (Steud.) Weddell in Ann. Sci. Nat., Bot., Ser. 4, 1: 181 (1854); Blume, Mus. Bot. Lugd.-Bat. 2: 159 (1855); Weddell, Monogr. Urt.: 169 & Tab. 2, Fig. 1-5 (1856) & in DC., Prodr. 16(1): 103 (1869); Engler, Hochgebirgsfl. Trop. Afr.: 193 (1892) & Pflanzenw. Ost-Afr. C: 163 (1895); Diels in Engl. Bot. Jahrb. 29: 302 (1901); L6veill6, Cat. P1. Yun-Nan: 275 (1915); Rendle in Prain (ed.), Fl. Trop. Afr. 6(2): 266 (1917); Hauman in Fl. Congo Belge 1: 196 (1948); Robyns, Fl. Sperm. P.N.A. 1: 76 (1948); Andrews, Flow. P1. Sudan 2: 278 (1952); Keay, Fl. W. Trop. Afr., ed. 2, 1(2): 618 (1958).

Girardinia javanica Weddell in Ann. Sci. Nat., Bot., S6r. 4, 1: 181 (1854). Type: Java, Zollinger 2539 (holotype ? P not traced).

Girardinia vitifolia Weddell in Ann. Sci. Nat., Bot., S6r. 4, 1: 181 (1854). Type: Java, Zollinger 2212 (holotype ? P not traced; isotypes BM, G, LE).

Girardinia vahlii Blume, Mus. Bot. Lugd.-Bat. 2: 158 (1855), nom. illegit. superfl. Based on Urtica palmata Forssk. and U. heterophylla Vahl; type as for Urticapalmata Forssk.

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Girardinia palmata Blume, Mus. Bot. Lugd.-Bat. 2: 158 (1855), non Urtica palmata Forssk., nom. illegit. superfl.; Weddell in DC., Prodr. 16(1): 101 (1869); Diels in Engl. Bot. Jahrb. 29: 302 (1901); Leveill, Cat. P1. Yun- Nan: 275 (1915); Handel-Mazzetti, Symb. Sinicae 7(1): 119 (1929); Backer & Backhuizen, Fl. Java 1: 39 (1965); Hara, Fl. East Himalaya: 59 (1966); Hamzah, Toha & van Steenis, Mt. Fl. Java: Tab. 55, 4 (1972). Type as for Girardinia leschenaultiana Decne.

Girardiniafurialis Blume, Mus. Bot. Lugd. -Bat. 2: 159 (1855). Type: Mada- gascar, Bojer I 142 (holotype L; no specimen traced at P).

Girardinia condensata (Steud.) Weddell var. nana Blume, Mus. Bot. Lugd.-Bat. 2: 160 (1855). Type: Asia (? where), Hiigel 4712 (holotype ?L not traced).

Girardinia zeylanica Decne. var. vitifolia (Weddell) Weddell, Monogr. Urt.: 166 (1856) & in DC., Prodr. 16(1): 102 (1869); Miq., Fl. Ind. Bat. 1, 2(2): 233 (1859). Type as for Girardinia vitifolia Weddell.

Girardinia vitifolia Franch. in Nouv. Arch. Mus. Hist. Nat., Str. 2, 10: 80

(1887), nom. illegit., non Weddell; Liveillk, Cat. P1. Yun-Nan: 275 (1915). Type: China, Eastern Tibet, Moupin, David s.n. (holotype P).

Girardinia heterophylla Decne. var. zeylanica (Decne.) Hook. f., Fl. Brit. India 5: 551 (1888); Duthier, Fl. Gangetic Plain 3(1): 125 (1915). Type as for Girardinia zeylanica Decne.

Girardinia heterophylla Decne. var. palmata Hook. f., Fl. Brit. India 5: 551

(1888) & in Trimen, Fl. Ceylon 4: 106 (1898). Based on Girardinia palmata Blume, nom. illegit.; type as for Girardinia leschenaultiana Decne.

Girardinia condensata (Steud.) Weddell var. adoinsis (Steud.) De Wildem. in Ann. Mus. Congo, S&r. 4, Bot. 1(3): 173 (1903) & Contrib. etud. Fl.

Katanga: 50 (1921) & P1. Bequaert. 1(2): 197 (1922); Peter in Fedde

Repert. Sp. Nov., Beih. 40, 2(1): 120 (1932). Type as for Urtica adoinsis Steud.

Girardinia formosana Yamemoto, Suppl. Icon. P1. Formosa. 1: 3 (1925); Sasakin in Rep. Res. Inst. Formosa 9: 180 (1930). Type: Taiwan (For- mosa); Tappansha, Kawakami & Mori s.n., Mt Arisan, Nimandira s.n.

Jan. 1915, Kiraikei & Nokozan s.n. Aug. 1919 (all syntypes, none traced). Girardinia longispica Handel-Mazzetti in Sinensia 2: 133 (1932). Type: China,

Kweichow, Tsiang 4226 (holotype ? IBSC not traced). Girardinia chingianae Chien in Contr. Biol. Lab. Chin. Assoc. Advanc. Sci.,

Sect. Bot. Ser. 9: 259 & Fig. 25 (1934). Type: China, Yutsien, Ching 4698 & Chang 121 (both syntypes, neither traced).

Girardinia erosa Decne. var. occidentalis Leandri in Ann. Mus. Col. Marseille 7: 31 (1951) & Fl. Madagascar 56, Urtic.: 26 (1965), nom. non rite publ. Type: Madagascar, Analfanja, Humbert 14300 (holotype ?P not traced).

Girardinia heterophylla Decne. subsp. adoinsis (Steud.) Cuf. in Bull. Jard. Bot. Nat. Belg. 39, suppl.: XX (1969). Type as for Urtica adoinsis Steud.

NOMENCLATURE. Vahl (1790) altered the name of ForsskAl's taxon and

amplified the protologue to include a plate in Rheede-tot-Drakenstein (1679), showing a Girardinia from India. Vahl thus created an illegitimate, superfluous name according to art. 63 of the International Code. Link

(1822) described two new species of Urtica, U. diversifolia and U. horrida. The latter is illegitimate because it is a later homonym. Urtica diversifolia Blume is a later homonym for the former.

Gaudichaud (1830) established the genus Girardinia without indicating

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FIG. 3. Girardinia diversifolia. A habit. B upper surface of leaf, detail. C underside of leaf, detail. D female flower. E open male flower, from above. F open male flower, side view. All from Chiovenda 1563.

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any specific epithets to be used in combination with his new generic name. His statement 'Species hujus generis sunt: Urtica palmata Lesch. (Herb. Mus.), &c.' refers to a specimen collected in India by Leschenault and preserved at P. On the label of this specimen is written 'No. 54. Urtica palmata, nom indigene: tourrouss6. Mts. Nillijgerry, Leschenault. Plante herbac6e plus vermineuse que l'ortie d'Europe.' There is thus no reference, directly or indirectly, to ForsskAl's Urtica palmata. The mere identity of the two names does not allow us to assume a connection; there are other examples in Girardinia where identical epithets have been given absolutely independently to two species, even by botanists associated with the same institution (G. vitifolia)! Gaudichaud must therefore be considered to have published a nomen nudum 'Urtica palmata Leschenault ex Gaudichaud'. Even if there had been a reference to Forsskil's species it would not have been possible to ascribe a combination with the epithet palmata to Gaudichaud because of art. 33 of the Code, under which examples from Bentham & Hooker's Genera Plantarum very similar to this are cited as examples of combinations not validly published. Nor is it possible to regard Gaudichaud as publishing a new species validated by a descriptio generico-specifica since this, according to art. 42, requires that the author regards his new genus as monospecific, and Gaudichaud clearly states that the genus has more than one species. It is therefore necessary to accept that the first validly published specific names in Girardinia were those published by Decaisne (1844), a view which has also been adopted by the compiler of the Index Nominum Genericorum.

Decaisne adopted the name Girardinia heterophylla for ForsskMl's taxon, indicating his intention by an indirect reference to Vahl's Urtica hetero- phylla, and provided it with a Latin description; but as Urtica heterophylla Vahl is illegitimate the combination Girardinia heterophylla has, according to art. 72, to be ascribed solely to Decaisne. The Leschenault specimen quoted by Gaudichaud was by Decaisne referred to a distinct species called Girardi- nia leschenaultiana.

Blume (1855) published a synopsis of Girardinia as it was known then. He reinstated the name 'Girardinia palmata Gaud'. and provided it with a validating Latin description, citing Girardinia leschenaultiana Decne. as a synonym. As the combination G. palmata had not previously been validly published the binomial must be ascribed to Blume, and as it is based on the type of the already validly published G. leschenaultiana it is nomenclaturally superfluous. It prevents the reinstatement of the epithet palmata of ForsskMl in a legitimate combination with Girardinia. In fact, Blume specifically excluded the Forsskal-specimen from his circumscription of G. palmata, stating that he proposed the name Girardinia vahlii for ForsskMl's taxon. Under the present rules of botanical nomenclature it is therefore necessary to find the next validly published specific epithet, which is Urtica diversifolia Link, and propose the combination G. diversifolia.

TAXONOMY. The latest revision of the genus as a whole was made by Weddell (1856 & 1869) who accepted the following taxa: G. heterophylla Decne. (northern India to Burma, Yemen), G. palmata Blume (southern India), G. zeylanica Decne. (southern India, Ceylon, Java), G. erosa Decne. (Madagascar), G. condensata (Steud.) Weddell (Ethiopia, northern India), G. bullosa (Steud.) Weddell (Ethiopia) and G. cuspidata Weddell (China). Hooker (1888) united G. palmata and G. zeylanica with G. heterophylla under

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the latter name, suggesting an infraspecific classification to express the variation. The more recently described species have all by their authorities been considered to be near G. heterophylla (or G. palmata).

The present study indicates that all these taxa, with the exception of G. bullosa and G. cuspidata, are so closely related that it is impossible to keep them as distinct species in spite of a considerable variation in the material. The most striking variation was found in the leaf morphology, as illustrated in Fig. 4. Very similar leaf shapes may be found in specimens from widely separated localities. Leaves with very deep incisions are observed on specimens from Madagascar, Ceylon and southern India (no. 12, 13, 15 & 22). The 'vitifolia' type of leaf has been seen on specimens from China and Java (no. 19 & 21), and a type with regularly serrate leaves can be found on specimens from the Himalayas and West Africa (no. 7, 8, 10 & 16). Other recognizable forms can be found on specimens from China, India and Java (no. 1, 3 & 5), and on specimens from Africa, Yemen and India (no. 4, 6, 12 & 20, specimens from the Yemen not illustrated). But the overall variation within each local area, and sometimes indeed on the same plant, tends to blur any clear pattern.

There is also a considerable but continuous variation in the shape and size of the female inflorescence. Extremely long and virtually unbranched types are seen on specimens from India and Indonesia, whereas extremely condensed types are found on specimens from Africa ('G. condensata'), India and Ceylon. Intermediate types occur with the extreme ones throughout the range, and characters from the inflorescence seem in no way to be correlated with leaf characters. The same applies to the noticeable variation in the size and sculpturing of the achenes.

It seems that it is necessary either to accept a very high number of more or less arbitrarily defined taxa with almost inconceivable patterns of dis- continuous distributions, or to recognize one polymorphic species with no formal infraspecific taxa. It is my impression that the latter solution is by far the most appropriate.

DISTRIBUTION. Map 2. (Guin&e, C6te d'Ivoire, Nigeria, Cameroun, Zaire, Sudan, Ethiopia, Kenya, Uganda, Burundi, Tanzania, Angola, Zambia, Malax^i, Zimbabwe, Mozambique, South African Republic (Transvaal), Madagascar, Yemen, India (Himalaya, western and southern India), Nepal, Sikkim, Sri Lanka, Burma, Thailand, Laos, China (Tibet, Yunnan, eastern and northeastern China), North Korea, Taiwan, Indonesia (Java, Sumatra, Bali, Celebes, Timor)).

ILLUSTRATIONS. Figs. 3 & 4.

SPECIES EXCLUDED FROM GIRARDINIA

1. Girardinia cuspidata Weddell was described from specimens collected in China by Pierre David. Two sheets of this collection have been seen from P, both answering to Weddell's quite detailed description and both annotated as G. cuspidata by Weddell. A drawing of the female flower, apparently also by Weddell, was attached to one of the sheets. This drawing shows only two perianth segments, a composed larger segment and a smaller free one, as in Girardinia, but closer examination of the specimens shows that the female

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154 KEW BULLETIN VOL. 36(1)

FIG. 4. Girardinia diversifolia. Variation in leaf morphology. 1 Bretschneider 1957 (China, mts W of Peking). 2 Dalhousie s.n. (India, Simla). 3 Madras Herbarium 49509 (India, Palni Hills, Kodaikanal Ghat). 4 Tweedie 2240 (E Africa, N side of Mt Elgon on Kenya/Uganda border). 5Junghuhn s.n. (Java). 6 Pobiguin 2278 (WAfrica, Guinee, Pita). 7 Hooker s.n. (Sikkim). 8 Hooker s.n. (Himalaya). 9 Garrett 600 (Thailand, Doi Angka). 10 Anderson s.n. (N India, Mussoorie).

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A SYNOPSIS OF GIRARDINIA (URTICACEAE) 155

FIG. 4, continued: 11 Simpson 9211 (Sri Lanka, Hanguranketa to Hewaheta). 12 Leschenault 54 (S India, Nilgiri Hills). 13 Leschenault s.n. (Sri Lanka, Koudij). 14 Chevalier 18569 (W Africa, Guinee, Fouta djalon). 15 Thwaites C. P. 3520 (Sri Lanka sine loc.). 16 Jacques-Fdlix 8994/b (Cameroun, Tchabal Mbabo). 17 Gamble 12616 (S India, Nilgiri). 18 Henry 9065 (China, Yunnan, Mengzi). 19 David s.n. (China, Moupin). 20 Mooney 1467 (India, Bastar State, Bailadila Hill). 21 Backer 9705 (Java, Juang Plateau). 22 Perrier de la Bdthie 13063 (Madagascar, Betafo).

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156 KEW BULLETIN VOL. 36(1) flowers have in fact four tepals, as the female flowers of Laportea, to which genus G. cuspidata must consequently be transferred. A comparison of the specimens with Chew's monograph of Laportea (Chew 1969) shows that G. cuspidata is identical with Laportea macrostachya (Maxim.) Ohwi, and indeed antedates the basionym of the latter name. The following new combination has therefore to be made:

Laportea cuspidata ( Weddell) Friis comb. nov.

Girardinia cuspidata Weddell in DC., Prodr. 16(1): 103 (1869); Hance in Journ. Linn. Soc. Bot. 13: 86 (1873); Forbes & Hemsley in Journ. Linn. Soc. Bot. 26: 475 (1899); Komarov, Fl. USSR 5: 395 (1936). Type: China, near Peking, David 2278 (two isotypes, P).

Sceptrocnide macrostachya Maxim. in Bull. Acad. Petersb. 22: 238 (1877). Type: Japan, Maximowicz s.n. (isotypes BM, K).

Laportea macrostachya (Maxim.) Ohwi in Journ. Jap. Bot. 12: 331 (1936) & Fl. Japan (English ed.): 387 (1965); Chew in Garden's Bull. Singapore 25: 142 (1969). For further synonymy, see Chew 1.c. The species is distributed in Burma,

China and Japan.

2. Girardinia marginata Engler, Bot. Jahrb. 33: 123 (1902). The type seems no longer extant, but Engler states in the diagnosis that the female flowers have lateral tepals (as in Laportea) to which genus Rendle (1917) and all subsequent authors have referred it as a synonym of Laportea alatipes Hook. f.

ACKNOWLEDGEMENTS

I am grateful to a group of research students at the Institute of Systematic Botany who, as part of their training, took part in the work and helped scoring of characters, scanning of literature and tracing of localities. The students were Messrs J. S. Andersen, N. Jacobsen, T. O. Pedersen, F. N. Rasmussen and K. B. Vollesen. I am further indebted to Dr B. Hansen, Botanical Museum, Copenhagen, for discussions of the taxonomic problems in G. diversifolia and for help with Asian literature and localities, and to Mr R. D. Meikle, Kew, for a discussion of the nomenclatural intricacies of the same species. The drawings were done by the artist Mr B. Johnsen. Finally I am obliged to the many herbarium curators who generously have put the material of Girardinia in their care at my disposal.

REFERENCES

Blume, C. L. (1855). Museum Botanicum Lugduno-Batavum, 2, Fasc. 10-11 (p. 485-528). Leiden.

Chew, W.-L. (1969). A monograph of the genus Laportea (Urticaceae). Garden's Bull. Singapore 25: 111-178.

Decaisne, J. (1844). Botanique, p. 57-183, in V. Jaquemont, Voyage dans l'Inde 4. Paris.

Gaudichaud, C. (1830, '1826'). Botanique, part 12. In L. de Freycinet, Voyage autour du monde.. . excut6 sur les corvettes de S.M. l'Uranie et la Physicienne. Paris.

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Hooker, J. D. (1888). Flora of British India 5, Part 15. London. Letouzey, R. (1968). Ulmactes, Urticactes. Flore du Cameroun 8. Paris. Link, H. F. (1822). Enumeratio plantarum horti regii botanici Berolinensis

altera 2. Berlin. Rendle, A. B. (1917). Urticaceae. In D. Prain (ed.), Flora of Tropical

Africa 6(2). London. Rheede-tot-Drakenstein, H. A. (1679). Hortus Indicus Malebaricus 2.

Amsterdam. Vahl, M. (1790). Symbolae botanicae 1. Kobenhavn. Weddell, H. A. (1856). Monographie de la famille des Urticac6es. Paris.

- (1869). Urticacees. In A. De Candolle, Prodromus systematis naturalis regni vegetabilis 16(1). Paris.