Nomenclatural Changes in Apteranthes Mikan (Asclepiadoideae, Apocynaceae)

Flora Montiberica 32: 15-20 (V-2006)

NOMENCLATURAL CHANGES IN APTERANTHES MIKAN (ASCLEPIADOIDEAE, APOCYNACEAE)

Manuel B. CRESPO VILLALBA

CIBIO, Instituto de la Biodiversidad, Universidad de Alicante. Apartado 99. E-03080 Alicante. E-mail: [email protected]

RESUMEN: El género Apteranthes Mikan se considera constituido por unos ocho táxones, en su mayoría distribuidos por el norte de África. Sin em- bargo, algunos de sus componentes han sido infravalorados en las revisiones más recientes del género, pese a estar bien caracterizados morfológica y bio- geográficamente. En la presente contribución, se aportan datos que sirven para justificar la propuesta de dos nuevas combinaciones nomenclaturales: Apte- ranthes europaea subsp. negevensis (Zohari ex Feinbrun) M.B. Crespo –que incluye las poblaciones orientales de la especie–, y A. munbyana subsp. hispanica (Coincy) M.B. Crespo & Mateo –que se aplica las poblaciones del sudeste ibérico–. Para ambas se adjuntan icones de los autores originales de cada taxon.

SUMMARY: Apteranthes Mikan is currently accepted to include about eight taxa, mostly distributed throughout North Africa. However, some of the described taxa have been devaluated in recent reviews of that genus, although they are morphologically and biogeographycally well characterized. In this contribution, data are reported to justify two new nomenclatural combinations: Apteranthes europaea subsp. negevensis (Zohari ex Feinbrun) M.B. Crespo –to group the eastern populations of the species–, and A. munbyana subsp. hispanica (Coincy) M.B. Crespo & Mateo –to include the Iberian populations–. For each one, drawings from the original publications are shown.

INTRODUCTION clepioideae, Apocynaceae), which has its centre of origin in East Africa. Caralluma R. Br (sensu lato) has been In the last comprehensive review of usually accepted to include about 120 Caralluma (sensu lato), GILBERT (1990) taxa, with a wide African, Asian, South accepted 57 species arranged in several African and southwestern European dis- subgenera. Recent studies based on mor- tribution (MABBERLEY, 1993). It be- phology (PLOWES, 1995) and cladistic longs to –and in a wide sense it conforms, analyses of different molecular data (ME- following MEVE & LIEDE (2004)– sub- VE & LEIDE, 2002, 2004) have demon- tribe Stapeliinae (tribe Ceropegieae, As- strated however the convenience of split-

15 Flora Montiberica 32 (2006) ting Caralluma (s. l.) in smaller units, thern Africa, Arabia); Monolluma Plowes which accord better with the morphologi- (c. 5 taxa; northeastern Africa, Arabia, cal divergence and geographical distribu- Socotra); and Anomalluma Plowes (2 ta- tion of the complex. xa; Somalia, Omna, southern Yemen). Concerning the Mediterranean spe- Among them, Apteranthes is charac- cies, and particularly those growing in the terized by its lateral pseudo-umbellate in- Iberian Peninsula, they are currently pla- florescences, and its mostly ovate-lan- ced in Apteranthes Mikan (PLOWES, ceolate, conspicuous leaf rudiments. Taxa 1995; MEVE & LEIDE, 2002), a small included here are distributed mainly in genus highly diversified in North Africa North Africa, though some reach the Ibe- (cf. JAHANDIEZ & MAIRE, 1934: 581- rian Peninsula –A. europaea and A. mun- 583, as Caralluma). byana–, and others spread to Asia –A. Nevertheless, the taxonomic arrange- staintonii (Hara) Meve & Leide– and nor- ments of Apteranthes by PLOWES (1995) theastern India –A. tuberculata (N.E. Br.) and MEVE & LEIDE (2002) are still Meve & Leide. unsatisfactory, since they neglect of de- valuate some Mediterranean taxa. There- The complex of A. europaea fore, in the present contribution some no- Apteranthes europaea (s.l.) is widely menclatural changes are proposed to ac- distributed from southeastern Iberian Pe- commodate them to Apteranthes. ninsula to the Middle East, throughout North Africa. Every single population shows peculiarities mainly in the habit RESULTS AND DISCUSSION and the colour patterns of the flowers, which led to the description of many taxa Perhaps the most analytic rearrange- which are still in need of further taxo- ment of the Caralluma complex was pu- nomic evaluation. blished by PLOWES (1995), who accep- Despite the diversity of flower size ted 17 genera, of which 12 were described and colouring, there exist indeed some as new, on the basis of morphological constant differences separating popula- differences associated to biogeographical tions from the Middle East (Sinai, Israel distribution. and Jordan; Fig. 1), as suggested by ME- Molecular studies by MEVE & LEI- VE & HENEIDAK (2005). Among them, DE (2002, 2004) have demonstrated the the shape of leaf rudiments, the stem convenience of accepting many of Plo- colour and surface, the corolla trichomes, wes’s segregates. The arrangement of the epidermal cells and colouring patterns, former authors leaves subtribe Stapeliinae and the gynostegial features, are probably with 36 genera (MEVE & LEIDE, 2004), the most reliable characters to separate 8 of which are segregates of the Cara- both geographical groups. lluma complex: Caralluma R. Br. (c. 27 Those eastern populations have been taxa; Africa, Arabia, India); Apteranthes named in different taxonomic ranks (e.g. Mikan (c. 8 taxa; south and southwestern, Caralluma europaea var. judaica, C. ne- Mediterranean, Canary Islands, northern gevensis, or C. aaronis), and even accep- Africa, Arabia, western and central Asia); ted as independent proper taxa (cf. FEIN- Australluma Plowes (1 taxon; Namibia); BRUN, 1978). However, all of them seem Boucerosia Wight & Arn. (c. 7 taxa; sou- to belong to a unique entity which MEVE thern Asia); Caudanthera Plowes (c. 3 ta- & HENEIDAK (2005) accepted as C. eu- xa; Africa, Arabia, southwestern Asia); ropaea var. judaica. Nonetheless, diffe- Desmidorchis Ehrenb. (c. 11 taxa; nor- rences between the eastern and western

16 M.B. CRESPO: Nomenclatural changes in Apteranthes (Asclepiadoideae-Apocynaceae) taxa are sufficient to regard them at the NOMENCLATURAL CHANGES subspecies rank, as proposed here. According to the discussion above, the The complex of A. munbyana following combinations are proposed for Apteranthes munbyana is endemic to some Mediterranean taxa of Apteranthes the southwestern Mediterranean areas, na- (a more detailed synonymy is reported in mely to Morocco, Algeria, and Spain. It MEVE & LEIDE, 2004): was described in the genus Boucerosia from plants growing in Oran (Algeria). 1. Apteranthes europaea (Guss.) Plowes, The Iberian populations were segre- Haseltonia 3:59 (1995) ≡ Stapelia europaea Guss., Fl. Sic. Prod. gated as B. munbyana var. hispanica by Suppl. 1: 65 (1832) [basion.] ≡ Boucerosia DE COINCY (1898), on the basis of europaea (Guss.) Caruel in Parl., Fl. Ital. plants previously identified as Apteran- 6: 725 (1886) ≡ Caralluma europaea thes gussoneana (DE COINCY, 1893: 24, (Guss.) N.E. Br., Gard. Chron. 12: 369 tab. X, fig. A). They differed from the (1892) typical Algerian plants by its leaf rudi- subsp. europaea ments smaller, sessile and obtuse to sub- acute; its stems thinner and obtusely an- = A. gussoneana Mikan, Nov. Act. Nat. gled; its corona pieces shorter and shortly Cur. 17: 594 (1835) ≡ Boucerosia gussoneana (Mikan) Hook. f., Bot. connate; and its fruits straight, only Mag.: 100, t. 6137 (1874) hooked at the apex. Finally, this author (DE COINCY, 1899) studied with more = Boucerosia maroccana Hook. f., Bot. Mag.: 100, t. 6137 (1874) ≡ Ca- detail the Spanish plants and suggested ralluma maroccana (Hook. f.) N.E. they should constitute a proper species he Br., Gard. Chron. 12: 370 (1892) drew with detail later (DE COINCY, = Caralluma simonis Hort. ex Berger, 1901: Tab. 7; Fig. 2). Monatsschr. Kakt. 14: 6 (1904) ≡ Although several authors (BRUYNS, Boucerosia simonis (Hort. ex Ber- 1987; GILBERT, 1990; MEVE & LEI- ger) A.C. White & C. Sloane, Stap.: DE, 2004) have neglected or devaluated 191 (1933) the Iberian taxon, M.B. Crespo and G. = Caralluma europaea var. confusa Mateo (in JUAN & al., 1995), accepted it Font Quer, Mem. Mus. Ci. Nat. Bar- at the subspecies rank on the basis of celona, Ser. Bot. 1(2): 12 (1924) morphological and biogeographical dif- subsp. negevensis (Zohary ex Fein- ferences. This treatment has been fol- brun) M.B. Crespo comb. & stat. nov. lowed by other authors (cf. SÁNCHEZ GÓMEZ & al., 1997; LAGUNA & al., ≡ Caralluma negevensis Zohary ex Feinbrun, Fl. Palaestina 3: 449, Ap- 1998; BOLÒS & al., 2005), and the taxon pendix (1978) [basion.] ≡ A. nege- has been included in the Spanish “red vensis (Zohary ex Feinbrun) Plowes, book” as Vulnerable, as well as C. euro- Haseltonia 3: 61 (1995) paea (BAÑARES & al., 2003). = Caralluma europaea var. judaica Differences between the North Afri- Zohary, Pal. J. Bot. ser. 2: 173 can and the Spanish plants are significant (1941) ≡ A. europaea var. judaica enough to consider recognition of De Co- (Zohary) Plowes, Haseltonia 3: 61 incy’s taxon as a proper subspecies, tho- (1995) ugh in the genus Apteranthes as shown = Boucerosia aaronis Hart, Trans. Roy. below. Irish Acad. 28: 436 (1885)

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2. Apteranthes munbyana (Decne. ex GILBERT, M. G. (1990). A review of Cara- Munby) Meve & Leide, Pl. Syst. Evol. lluma R. Br. and its segregates. Bradleya 234: 199 (2005) 8: 1-32. JAHANDIEZ, E. & R. MAIRE (1934). Cata- subsp. munbyana logue des plantes du Maroc (Spermatop- hytes et Ptéridophytes), vol. 3. Minerva. subsp. hispanica (Coincy) M.B. Cres- Alger. po & Mateo comb. nov. JUAN, A., L. SERRA & M. B. CRESPO ≡ Boucerosia munbyana var. hispani- (1995). Adiciones a la flora alicantina. Ac- ca Coincy, J. Bot. (Morot) 12: 250 ta Bot. Malacitana 20: 284-290. (1898) [basion.] ≡ B. hispanica LAGUNA, E., M. B. CRESPO, G. MATEO, (Coincy) Coincy, J. Bot. (Morot) 13: S. LÓPEZ, C. FABREGAT, L. SERRA, J. 336 (1899) ≡ Caralluma munbyana J. HERRERO-BORGOÑÓN, J. L. CA- var. hispanica (Coincy) Maire in Ja- RRETERO, A. AGUILELLA & R. FI- hand. & Maire, Cat. Pl. Maroc 3: GUEROLA (1998). Flora endémica, rara 582 (1934) ≡ C. munbyana subsp. o amenazada de la Comunidad Valencia- hispanica (Coincy) M. B. Crespo & na. Colecc. Biodivers. 1. Generalitat Va- Mateo, Acta Bot. Malacitana 20: lenciana. Valencia.

285. 1995. MABBERLEY, D. J. (1993). The plant.book. 4th reprint. Cambridge University Press. Cambridge. REFERENCES MEVE, U. & S. HENEIDAK (2005). A morphological, karyological and chemical BAÑARES, Á., G. BLANCA, J. GÜEMES, J. study of the Apteranthes (Caralluma) eu- C. MORENO & S. ORTIZ (eds.), Atlas y ropaea complex. Bot. J. Linn. Soc. 149: libro rojo de la flora vascular amenazada 419-432. de España. Táxones prioritarios. Dir. MEVE, U. & S. LEIDE. (2002). A molecular Gral. de Conservación de la Naturaleza. phylogeny and generic rearrangement of Madrid. the stapelioid Ceropegieae (Apocynaceae- BOLÒS, O. DE, J. VIGO, R. M. MASALLES Asclepiadoideae). Pl. Syst. Evol. 234: 171- & J. M. NINOT (2005). Flora manual dels 209. Països Catalans. 3th ed. Pòrtic. Barcelona. MEVE, U. & S. LEIDE. (2004). Subtribal BRUYNS, P. V. (1987). Miscellaneous notes division of Ceropegieae (Apocynaceae- on Stapelieae (Asclepiadaceae). Bradleya Asclepiadoideae). Taxon 53(1): 61-72. 5: 77-90. PLOWES, D.C.H. (1995). A reclassification DE COINCY, A. (1893). Ecloga plantarum of Caralluma R. Brown (Stapelieae: As- hispanicarum seu icones specierum nova- clepiadaceae). Haseltonia 3: 49-70. rum vel minus cognitarum per Hispanias SÁNCHEZ GÓMEZ, P., J. GUERRA, A. nuperrime detectarum. Masson. Paris. HERNÁNDEZ, S. FERNÁNDEZ, E. DE COINCY, A. (1898). Plantes nouvelles de COY, A. F. CARRILLO, M. J. TAMA- la flore d'Espagne (8ª note). J. Bot. (Mo- YO, J. GÜEMES & J. RIERA (1997). rot) 12: 250-252. Flora selecta de Murcia. Plantas endémi- DE COINCY, A. (1899). Plantes nouvelles de cas, raras o amenazadas. Consejería de la flore d'Espagne. J. Bot. (Morot) 13: 332- Medio Ambiente, Agricultura y Agua. Re- 338. gión de Murcia. DE COINCY, A. (1901). Ecloga quinta plantarum hispanicarum seu icones stirpium elapsis anniis per Hispanias lectarum. Masson et cie. Paris. FEINBRUN-DOTHAN, N. (1978). Flora Palaestina, vol. 3. The Israel Academy of Sciencies and Humanities. Jerusalem. (Received 17-I-2006)

18 M.B. CRESPO: Nomenclatural changes in Apteranthes (Asclepiadoideae-Apocynaceae)

Fig.1.- Apteranthes europaea subsp. negevensis (Zohary ex Feinbrun) M.B. Crespo. Drawings from FEINBRUN (1978, Plate 43 as Caralluma europaea var. judaica on the upper part, and Plate 44 as C. negevensis on the lower one).

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Fig.2.- Apteranthes munbyana subsp. hispanica (Coincy) M.B. Crespo & Mateo. Drawings from DE COINCY (1901, tab. 7, as Boucerosia hispanica, but number 13 corresponding to B. munbyana s.s.).