Nomenclatural changes in yams of Madagascar(Dioscorea: Dioscoreaceae)

Paul Wilkin1 & Tianjanahary Randriamboavonjy2

Summary. The accepted names and associated synonyms of 12 Dioscorea L. species present in the flora of Madagascarare lectotypified and two new synonyms established.

Key Words. lectotypification, names, species, Taxonomy.

IntroductionThe nomenclatural history of the endemic taxa of theyam genus (Dioscorea) in Madagascar began in the 19th

Century with the description of five new species byJ. G. Baker based on specimens collected by themissionary Baron (Baker 1882, 1883, 1885). Most ofthe currently recognised taxa were described in the firsthalf of the 20th Century through the work of HenriPerrier de la Bâthie and his collaborators (Jumelle &Perrier 1909, 1910; Perrier 1925, 1928, 1946; Burkill &Perrier 1950, 1951) culminating in the treatment of thegenus in the Flore de Madagascar et des Comores (Burkill &Perrier 1950). Knuth (1924) also described new taxa inhis Pflanzenreich treatment. The second half of the 20thCentury, in contrast, was an era of taxonomic stasis forMalagasy Dioscorea. The last decade has seen thepublication of several papers (Wilkin et al. 2000, 2002;Haigh et al. 2005; Weber et al. 2005; Wilkin et al. 2007,2008, 2008 publ. 2009, 2009a, b). These studies haveinvolved the lectotypification of D. trichantha Baker, D.bararum H. Perrier, D. maciba Jum. & H. Perrier, D.seriflora Jum. & H. Perrier, D. tsaratananensis H. Perrierand D. pteropoda Boivin ex H. Perrier.

In order to prepare for the production of a newtaxonomic treatment of Dioscorea (starting at specieslevel with those in Madagascar) via a scratchpad(customised taxonomic web tools, http://scratchpads.eu),the typification of all accepted and synonymous nameswas checked. While many names are already associatedwith individual type specimens, 12 others were found tohave two or more syntypes cited in the protologue. Thusthey require lectotypification as recommended underarticle 9.2 of the Vienna Code (http://ibot.sav.sk/icbn/main.htm). In all cases the lectotype selected wasthe most complete and representative specimen ofthose cited in the respective protologues, where

possible with duplicates distributed to multipleherbaria. Lectotypification of names of taxa in theD. soso Jum. & H. Perrier complex (including D.bemandry Jum. & H. Perrier and D. analalavensis Jum.& H. Perrier) will be dealt with in a separate papernow in preparation. Two new synonyms are alsopresented below.

Lectotypifications

1. Dioscorea antaly Jum. & H. Perrier (Jumelle &Perrier 1909: 485, nom. nud., 1910: 403); Knuth(1924: 88); Perrier (1925: 420, 1928: 22); Burkill &Perrier (1950: 26). Type: Madagascar, Manongarivo(Ambongo), no further data, ♂ fl. & old ♀ fr. Jan.1904, Perrier 8249 (lectotype P! (P01757345), selectedhere; isolectotype K! (K000728311)).Dioscorea antaly var. laevis Burkill & H. Perrier (1951:

136, 1950: 28). Type: Madagascar, Fianarantsoa,Midongy (Ouest), ♂ fl. & ♀ immature fr. Feb.1919, Perrier 12503 (holotype P! (P00440198) ♂fl.; isotype P! (P00440199) ♀ immature fr.).

NOTES. The specimens known to Perrier and collectedprior to 1910, when the name Dioscorea antaly wasvalidly published, were Perrier 448, 448bis, 8206, 8248,8249, 8250, 8251, 8252, Sol 505 and Pervillé 553 based onthose cited in Perrier (1928). It is therefore reasonable toassume that these specimens and their duplicates repre-sent syntypes. Perrier 8249 was selected as the lectotype inpart because the sheet at K bears Perrier’s handwritingand states ‘type des auteurs (Ann. Mus. Col. Mars. 1910)’.Knuth (1924) cited male and female specimens collectedby Perrier without collection numbers as the type(s)(‘Madagaskar: Auf Basalt und Gneiss (Herb. Perrier de

Accepted for publication February 2012.1 Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB UK. e-mail: p.wilkin@kew.org2 Royal Botanic Gardens, Kew, II J 131 B, Ambodivoanjo, Ivandry, 101 Antananarivo, Madagascar.

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la Bathie ♂♀ — Typus!))’. However, this does notconstitute lectotypification because it is not clear whichspecimen(s) Knuth was referring to.

The characters used to justify the recognition ofvar. laevis as a distinct taxon, specifically the absence ofdeltoid, translucent modified hairs (aspérités of Burkill& Perrier 1950) on the lower surface of the leaf andthe fruit shape, form part of the continuous variationof this widespread species. Bulbil-bearing forms areencountered occasionally in Dioscorea antaly as innumerous other members of the genus.

2. Dioscorea arcuatinervis Hochr. (Hochreutiner 1908:52); Knuth (1924: 70); Perrier (1928: 10); Burkill &Perrier (1950: 9). Type: Madagascar, Toamasina,Vatomandry, pres d'Ilaka, ♂ fl. 27 Jan. 1904, Guillot72 (holotype G, digital image! (G00018718); isotypesP! (P00442495), K! (K000098517)).Dioscorea humblotii R. Knuth (1924: 70). Type: Mada-

gascar, Toamasina, ‘Lagune Nossive au N deTamatave’ ♂ fl. 1883, Humblot 121 (holotype P!(P00442494); isotypes B, K! (K000098518)).

Dioscorea heteropoda Baker var. subdiffusa Burkill & H.Perrier (1951: 136), synon. nov. Type: Madagas-car, Toamasina, Zahamena (Reserve NaturelleNo. 3), ♂ fl. 21 March 1941, Decary 16508(holotype P! (P00442496)).

NOTES. The specimen Decary 16508 was incorrectlyidentified by Burkill & Perrier (1950, 1951) as fallingwithin the variation of Dioscorea heteropoda.

3. Dioscorea bemarivensis Jum. & H. Perrier (Jumelle& Perrier 1910: 423); Knuth (1924: 313); Burkill &Perrier (1950: 12). Type: Madagascar, Mahajanga,Haut Bemarivo (Boina), ♂ fl. Jan. 1907, Perrier 11520(lectotype P! (P00442488), designated by Perrier(1928: 8); isolectotype K! (K000098648)).Dioscorea lucida Scott-Elliot (1891: 60, nom. illegit. non

D. lucida R. Br (Brown 1810: 295); Perrier (1928:7). Type: Madagascar, Toliara, Fort Dauphin, ♂ fl.& ♀ immature fr. March 1890, Scott-Elliot 2624(holotype K! (K000098646); isotypes B, P!(P00442490)).

Dioscorea nesiotis Hemsl. (in Hemsley & Turrill 1917:288). Type: Aldabra, no further data, ♂ fl. April1907, Thomasset 241 (lectotype K! (K000098649),selected here).

Dioscorea madagascariensis R. Knuth (1924: 311). Type:Madagascar, Mahajanga, Moraitaitra, rive gauchede la Betsiboka prės Maevatanana, ♂ fl. March1899, Perrier 485bis (holotype B, digital image!(B100160981); isotypes K! (K000523632), P!(P01756327)).

Dioscorea majungensis R. Knuth (1924: 312). Madagas-car, Mahajanga, Mahajanga (Majunga), ♂ fl. &♀ immature fr. Dec. 1918, Perrier 12318 (holo-type B, digital image! (B100160980); isotypes P!(P01756370), K! (K000432448)).

NOTES. Knuth (1924) cited a specimen as followsunder Dioscorea bemarivensis ‘West-Madagaskar: Prov.Ambongo, Bemarivo (Perrier de la Bathie)’ giving nocollection number. Thus there is uncertainty regardingwhat specimen was being cited and the citation inPerrier (1928) of his own number 11520 should beregarded as lectotypification.

Three specimens of Dioscorea nesiotis were cited in theprotologue: Thomasset 218 and 241 from Aldabra andDupont 118 from Assumption, which represent syntypes.Thomasset 241 alone has been placed in a type cover at K,where all three specimens are held, but it does not appearto have been designated as the lectotype until now.

The type of Dioscorea madagascariensis was indicated tobe Perrier 485 in the protologue (Knuth 1924: 311) but asPerrier 485bis by Perrier (1928) and Burkill & Perrier(1950). Both collection numbers are D. bemarivensisJum. & H. Perrier, but they come from differentlocalities in northwestern Madagascar. However, thetype specimen in B bearing Knuth’s determinationin his own handwriting is clearly Perrier 485bis, notwith-standing what is indicated in the protologue, whichmust therefore be in error and the type Perrier 485bis.The confusion in the protologue may also have resultedfrom Knuth having seen Herbier du Museum 485 andHerbier Drake 485bis, both of which were collected wellbefore his monograph was published.

4. Dioscorea decaryana H. Perrier (1946: 205); Burkill &Perrier (1950: 66). Type: Madagascar, Fianarantsoa,environs de Ambatofindrahana, ♂ fl. 21 Feb. 1938,Decary 13146 (lectotype P! (P00442487), selected here).

NOTES. The specimen comprising male floweringmaterial selected here as the lectotype bears more ofthe distinguishing characters of this species than thefemale specimen that forms the other syntype, Decary13145 (P00442486). Dioscorea decaryana is known fromjust these two specimens. The protologue cited themale plant as Decary 13145 and female as 13146 inerror. The two specimens were assigned to the correctsex when cited in Burkill & Perrier (1950).

5. Dioscorea fandra H. Perrier (1928: 27); Burkill &Perrier (1950: 58). Type: Madagascar, Toliara, PlateauMahafaly, ♀ immat. fr. June 1910, Perrier 11620 L(lectotype P! (P01736956), selected here; isolectotypeK! (K000098961)).

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NOTES. The specimens of Dioscorea fandra cited in theprotologue were Perrier 11620 A, B, C, D, E, F, I, K andL, 11620 and 11679 (the latter being a typographic errorfor 11619, because Perrier 11679 is not a specimen ofDioscorea. The lectotype selected was the most completeand representative specimen of those cited.

6. Dioscorea heteropoda Baker (1882: 270), Jumelle &Perrier (1910: 388); Knuth (1924: 70); Perrier (1925:421, 1928: 35); Burkill & Perrier (1950: 49). Type:Madagascar, ‘chiefly in Betsileo-land’, no further data,♂fl. without date, Baron 119 (lectotype K! (K000098955),selected here).

NOTES. The protologue cited two syntypes, Baron 119and 139 (K000098954). Baron 4280 was stated to be atype (‘Typus ♀’) by Knuth (1924) with Baron 139 as‘Typus ♂’. Burkill & Perrier (1950) indicated thatBaron 4280 was the type, which would have constitutedlectotypification had it been cited in the protologue.Baron 139 has just two poorly preserved male inflor-escences and thus Baron 119 was selected as thelectotype.

7. Dioscorea hexagona Baker (1882: 270); Jumelle &Perrier (1910: 388); Knuth (1924: 68); Perrier (1925: 421,1928: 28), Burkill & Perrier (1950: 38). Type: Madagas-car, ‘chiefly in Betsileo-land’, no further data, ♂ fl.without date, Baron 140 (lectotype K! (K000098655),designated by Knuth (1924)).

NOTES. The protologue cited two syntypes, Baron 95and 140. Knuth (1924) lectotypified Baron 140 byindicating that it was the type. Unfortunately Baron140 (K000098655) is a very poor specimen with just fiveleaves and one inflorescence; Baron 95 (K000098658)would have made a better choice,

8. Dioscorea hombuka H. Perrier (1928: 31); Burkill &Perrier (1950: 56, 1951: 137). Type: Madagascar, Toliara,Plateau Mahafaly, Enandria, bas bassin de Menarandra,♀ fr. June. 1910, Perrier 11616 (lectotype P! (P00442483),selected here; isolectotypes K! (K000098959), P!(P00442482)).

NOTES. The protologue cited two collections, Perrier11616 andDecary 2577, of which the former was the mostcomplete and representative specimen. The two varie-ties of Dioscorea hombuka described by Burkill & Perrier(1950, 1951) need separate and detailed population-level taxonomic comparison, including the type varietyand related taxa, in order to uncover their patterns ofvariation before the nomenclature can be revised.

9. Dioscorea mamillata Jum. & H. Perrier (Jumelle &Perrier 1910: 422); Knuth (1924: 96); Perrier (1925: 421,1928: 37); Burkill & Perrier (1950: 16). Type: Madagascar,Mahajanga, Environs de Majunga, ♂ fl. March 1908,Perrier 8203 (lectotype P! (P00442471), selected byPerrier (1928); isolectotype P! (P00442472)).Dioscorea perpilosa H. Perrier (1946: 206), Burkill & H.

Perrier (1950: 17), synon. nov. Type: Madagascar,Mahajanga, Moraitaitra, rive droite de la Betsiboka,près Maevatanana, ♂ fl. May 1899, Perrier 846(lectotype P! (P00442467), selected here; isotypesK! (K000099037), P! (P00442464–6)).

NOTES. Perrier (1928) cited Perrier 8213 in error for 8203under Dioscorea mamillata, although the former collectionis not a Dioscorea specimen; Perrier 13910 and 13463 alongwith Campenon s.n. Jan. 1888, were also cited in Perrier(1928) and thus are also syntypes of D. mamillata.

The second syntype of Dioscorea perpilosa cited in theprotologue was Humbert 19044. However, the specimenof Perrier 846 (P00442467) in P is the most completeand has ‘type’ written on it by Perrier, although he didnot cite this specimen as a type in any publication.Knuth (1924) cited ‘Madagaskar: Mahevarano in derregion von Majunga (nach Jumelle und Perrier de laBathie)’ so it appears that he saw no specimens of thisspecies.

Dioscorea mamillata has 1 – 3 simple male inflor-escences per axil although in Perrier de la Bathie 846galling, possibly caused by insect activity, has producedmassive compound inflorescences with up to 20 partialinflorescences per axil (see Burkill & Perrier (1950)Fig. VII). This led Perrier to describe it as a distinctspecies, but it clearly represents the same entity as D.mamillata and belongs in synonymy under that name.

10. Dioscorea nako H. Perrier (1928: 30); Burkill &Perrier (1950: 46). Type: Madagascar, Toliara, Tongobory(Onilahy), ♂ fl. June 1910, Perrier 11614 (lectotype P!(P00496597) selected here; isolectotypes K! (000383683),P! (P00496596)).

NOTES. The protologue of Dioscorea nako cited thefollowing collections: Perrier 11614, 11615 and 12744,Poisson 399 2e voyage and Decary 2606, of which the firstis the most complete and representative specimen.

11. Dioscorea ovinala Baker (1882: 269); Jumelle &Perrier (1909: 486, 1910: 418); Knuth (1924: 95);Perrier (1928: 32); Burkill & Perrier (1950: 60). Type:Madagascar, ‘chiefly in Betsileo-land’, no further data,♂ fl. without date, Baron 52 (holotype K! (K000098962);isotypes B, digital image! (B1002278806), P(P00442470)!).

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Dioscorea velutina Jum. & H. Perrier (Jumelle & Perrier1910: 420), Knuth (1924: 96); Perrier (1925: 420,1928: 31). Type: Madagascar, Mahajanga, Bordsde la Mahevahingy, affluent de Mahevarano,Province d'Analalava, ♂ fl. Sept. 1908, Perrier8208 (lectotype P! (P00442469), selected here;isolectotype P! (P00442468)).

NOTES. Perrier (1928) cited the specimens Perrier 8208,Decary 64 and Decary 68 as syntypes of Dioscorea velutina.The latter two specimens give their collecting localityas Alottra (presumably Lac Alaotra in ToamasinaProvince, which is highly unlikely) while Perrier(1928) states that they were from Maromandia. ThusPerrier 8208 was selected in part because it was the onlysyntype of unequivocal provenance. Knuth (1924)cited ‘Madagaskar: Analalave, in Gebüschen an denUfern deMahevahinja, einemNebenfluss des Maivarano(nach Jumelle und Perrier de la Bathie’ so it appears thathe saw no specimens of this species.

12. Dioscorea proteiformis H. Perrier (1946: 199);Burkill & Perrier (1950: 17). Type: Madagascar, Fianar-antsoa, Mananjary, ♀ fl, March – April 1909, Geay 7036(lectotype P! (P01757329), selected here).

NOTES. The protologue cited Geay 7266, 7266bis, 7706,7925, 7926, 7471, 7037 and du Petit Thouars s.n. assyntype collections in addition to the specimen selectedas a lectotype.

13. Dioscorea sansibarensis Pax (1892: 146); Knuth(1924: 87); Perrier (1946: 200); Burkill & Perrier(1950: 4). Type: Tanzania, Bagamoyo (Bagamojo), ♂fl. May 1874, Hildebrandt 1284 (holotype B†).Dioscorea toxicaria Bojer (1837: 352), nom. nud.Dioscorea macroura Harms (1896: 266). Type: Yaounde,

♂ fl. Sept. 1894, Zenker & Staudt 414 (lectotypeK!, selected here; isolectotypes B†, BM!).

Dioscorea welwitschii Rendle (1899: 39). Type: Angola,Golungo Alto, Sobato, (Quilombo-Quiacatubia),♂ fl. Feb. 1855, Welwitsch 4041 (holotype BM!;isotype K!, sterile).

Dioscorea macabiha Jum. & H. Perrier, (Jumelle &Perrier 1909: 485, nom. nud., 1910: 407); Knuth(1924: 88); Perrier (1925: 419, 1928: 13). Type:Madagascar, Mahajanga, Autour de la ville deMaravoay (Boina), sterile without date, Perrier8247 (lectotype P!, selected here).

NOTES. Bojer (1837) states that Dioscorea toxicaria wascultivated ‘a Rivière Noire et a Mon Plaisir’ inMauritius and flowered in April and June. We suspectthat no specimens were made at that time. However,

Bojer had a drawing made of the leaves and bulbilswhich Burkill (1939) was able to identify asD. sansibarensis.

Dioscorea macroura was lectotypified using Zenker &Staudt 414 because the syntype Zenker 620 could not belocated in Berlin and was presumed lost.

Three specimens of Dioscorea macabiha were cited inPerrier (1928) and collected prior to 1910 and couldtherefore be syntypes: Perrier 8247 and Boivin 2013 and1685. Boivin 1685 was not located in P. Both Perrier8247 and Boivin 2013 are sterile, but the locality of theformer (environs de Maravoay) was mentioned byJumelle & Perrier (1910), suggesting that Perrier 8247was both in existence at that point and part of theirconcept of D. macabiha. Knuth (1924) cited ‘Madagas-kar: Ambongo, Boina, bei Maravoay auf Kalkboden(Herb. Perrier de la Bathie)’ which is also likely tohave been a reference to Perrier 8247. The vegetativemorphology of D. sansibarensis is highly distinctive,allowing sterile specimens to be definitively identifiedas belonging to this species.

AcknowledgmentsThe COLPARSYST programme provided financialsupport for a visit by PW to P in October 2003, andthe SYNTHESYS programme a further visit in October2006; he would like to thank Leandro Leoz and all thestaff of the herbarium for their help, especially the lateJean-Noël Labat and Thierry Deroin. Thanks are alsodue to Corine Koyamba of the Service du PEB,Bibliothèque centrale, P, Annette & Marcel Hladik,Vololoniaina Jeannoda, Mamy Tiana Rajaonah andtwo anonymous reviewers for the many improvementsthey made to this paper.

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