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Notes on Afrotropical Pyrgodesmidae, 2 (Diplopoda: Polydesmida)

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Notes on Afrotropical Pyrgodesmidae, 2 (Diplopoda: Polydesmida) Arthropoda Selecta 24(4): 387–400 © ARTHROPODA SELECTA, 2015 Notes on Afrotropical Pyrgodesmidae, 2 (Diplopoda: Polydesmida) Çàìåòêè ïî àôðîòðîïè÷åñêèì Pyrgodesmidae, 2 (Diplopoda: Polydesmida) S.I. Golovatch1, A.R. Nzoko Fiemapong2, D. VandenSpiegel3 Ñ.È. Ãîëîâà÷1, À.Ð. Íçîêî Ôüåìàïîíã2, Ä. ÂàíäåíØïèãåëü3 1 Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky prospekt 33, Moscow 119071 Russia. 1 Институт проблем экологии и эволюции РАН, Ленинский пр-т, 33, Москва 119071 Россия. 2 Laboratoire de Zoologie, Université Yaoundé 1, BP812, Cameroun. 2 Лаборатория зоологии, Университет Яунде 1, BP812 Камерун. 3 Musée Royal de l’Afrique Centrale, B-3480 Tervuren, Belgique. 3 Королевский музей Цеитральной Африки, Тервурен, Бельгия. KEY WORDS: millipede, Pyrgodesmidae, taxonomy, new species, new combination, key, Cameroon. КЛЮЧЕВЫЕ СЛОВА: диплопода, Pyrgodesmidae, таксономия, новый вид, новая комбинация, ключ, Камерун. ABSTRACT. The genera Urodesmus and Mona- Material and methods chodesmus are reviewed and shown to comprise four and 18 species, respectively, all keyed, including U. Most of the material treated here derives from the cornutus sp.n., M. longicaudatus sp.n. and M. armorum collection of the Musée Royal de l’Afrique Centrale sp.n., all three from Cameroon. In addition, the West (MRAC), Tervuren, Belgium, with only a few dupli- African genus Udodesmus is recorded in Cameroon for cates retained for the collection of the University of the first time, being represented there by U. camerun- Yaoundé 1, Cameroon (UYC) or donated to the Zoo- ensis sp.n., which is unique among congeners in show- logical Museum, State University of Moscow (ZMUM), ing 5+5 undifferentiated lobulations at the anterior mar- Russia. The samples are stored in 70% ethanol. Speci- gin of the collum. mens for scanning electron microscopy (SEM) were air-dried, mounted on aluminium stubs, coated with gold and studied using a JEOL JSM-6480LV scanning РЕЗЮМЕ. Даны обзоры родов Urodesmus и electron microscope. Photographs were taken with a Monachodesmus, которые содержат теперь соответ- Leica digital camera Leica DFC 500 mounted on a ственно четыре и 18 видов. Для них всех составле- Leica MZ16A stereomicroscope. Images were pro- ны кдючи, в т.ч. U. cornutus sp.n., M. longicaudatus cessed with the Leica Application Suite software. sp.n. и M. armorum sp.n. из Камеруна. Кроме того, западноафриканский род Udodesmus впервые при- Taxonomic part водится для фауны Камеруна, где он представлен U. camerunensis sp.n., который уникален среди ви- дов рода тем, что у него 5+5 недифференцирован- On the statuses of Urodesmus Porat, 1894 and Mo- ных долей на переднем краю коллума. nachodesmus Silvestri, 1927 Urodesmus was one of the earliest genera of Pyr- godesmidae to have been proposed. Porat [1894] com- Introduction pared it directly with Pyrgodesmus Pocock, 1892, and described two species therein: Urodesmus erinaceus The mainly tropical millipede family Pyrgodesmi- Porat, 1894 and U. sexcarinatus Porat, 1894, both from dae is among the largest, but taxonomically perhaps Cameroon. However, Urodesmus was properly typified most confused in the entire class Diplopoda, currently only by Silvestri [1896] who not only beautifully re- counting almost 400 species or subspecies in 170+ described U. erinaceus from type material [Silvestri, genera, nearly 120 of which are monotypic [Jorgensen, 1927], but also designated it as the type species. Verhoeff Sierwald, 1910; updated]. [1936] ignored the above typification, as well as a cou- The present paper puts on record four new species ple of varieties which Silvestri [1927] had distinguished of Pyrgodesmidae from Cameroon. In this connection, in U. erinaceus: U. e. var. nigeriensis, from Nigeria, two genera are reviewed and rediagnosed. and U. e. var. minor, from Fernando Po Island. 388 S.I. Golovatch, A.R. Nzoko Fiemapong, D. VandenSpiegel From the very start, U. erinaceus was readily dis- cially remarkable, however, is the presence of caudad tinguished from U. sexcarinatus by the presence of a rather strongly elongated, coniform crests 19 which paramedian pair of very high, sometimes vaguely bi- conceal the tip of the epiproct from above. lobed, basally subcontiguous, anteriorly directed, round- Such conditions readily remind of those observed ish tubercles located in the central part of the collum in the rather large, mostly Afrotropical genus Mona- behind a nearly straight anterior margin with 5+5 rath- chodesmus Silvestri, 1927. The main differences from er superficial lobulations [Porat, 1894; Silvestri, 1927]. Urodesmus lie in the ornamentation of the collum and Each following metatergite in U. erinaceus shows a following metaterga, these in the latter genus showing paramedian pair of similarly high, often heavily earth- 1+1 high dorsal protuberances which are rather weakly encrusted, apically largely vaguely bi- or trilobate, most- split on segment 19 and usually fail to conceal the tip ly subvertical and clearly separated tubercles growing of the epiproct from above (Figs 1A, 2A–I). The distri- increasingly strongly inclined caudad only in the last bution patterns of ozopores and tergal ornamentations few segments. The paraterga are prominent, set very along the body remain basically the same, only poros- low, subhorizontal, lying almost level to venter, vague- teles in Monachodesmus are exceptionally absent from ly bi- or trilobate at the lateral margin, devoid of ante- segment 16. Each dorsal crest on segments 2–4 is com- ro- and caudolateral lobulations. The pore formula is posed of two tubercles, thereafter, like all lateral crests, normal (5, 7, 9, 10, 12, 13, 15–19), very evident poros- of three tubercles. The paraterga in Monachodesmus teles are only present in paraterga 5, 7, 9, 10, 12, 13, 15 species are only seldom poorly developed, nearly want- and 16. The dorsal protuberances of segment 19 are ing, while many species show small intercalary granu- clearly directed caudad, short to moderately elongate, lations between and below the main (2+2) metatergal even if clearly split medially, then leaving the tip of the crests. In addition, the epiproct in Monachodesmus is epiproct fully or nearly exposed in dorsal view. The often fully concealed from above by the medially fused gonopods in U. erinaceus are deeply sunken inside a ultimate dorsal crests, although in some species it re- prominent gonocoel, the telopodites are massive, com- mains up to fully exposed due to the rather short and plex, deeply bi- or tripartite, including a solenomere. independent ultimate crests. All possible transitions in Such a concept of Urodesmus has since remained the degree of fusion and caudad protrusion of the ulti- largely intact. Only Attems [1940] correctly added Sty- mate crests are thereby observed. lodesmus camerunensis Silvestri, 1927, from Came- In general, the ultimate crests (usually 19th, depend- roon, while Stylodesmus horridus Cook, 1895, and its ing on the number of body segments) being very con- further two congeners from western Africa, have been siderably elongated/extended caudad is a common trait removed [Mauriès, Maurin, 1981; Golovatch, Vanden- among tropical Pyrgodesmidae. There can hardly be Spiegel, 2014]. As a result, Urodesmus currently en- any doubt that this condition may have appeared many compasses only the following few species: Urodesmus times in the course of the family’s evolution. Putting erinaceus Porat, 1894 (the type species), from Came- special emphasis on this character alone has resulted in roon, Fernando Po Island, and Nigeria, U. camerunen- such speciose artificial assemblages as the genera Lo- sis (Silvestri, 1927), from Cameroon, and U. simplex phodesmus Pocock, 1892, Myrmecodesmus Silvestri, Silvestri, 1927, from Nigeria. 1910, Monachodesmus and some others. Thus, Lo- Urodesmus serratus Verhoeff, 1936, described phodesmus is actually based on L. pusillus Pocock, from Sri Lanka as the type, and sole constituent, spe- 1894, a species from Flores, Indonesia, but numerous cies of the subgenus Attemsocyphus Verhoeff, 1936 congeners have since been described not only from [Verhoeff, 1936; Attems, 1940], seems best to be re- tropical parts of Asia, but also from Africa and, espe- moved from Urodesmus, albeit largely based on zoo- cially, America. Hoffman [1999] emphasized that no- geographical grounds. Here we follow Verhoeff [1937] one must take Lophodesmus seriously at the present. not only because he promoted Attemsocyphus to the The situation concerning Myrmecodesmus seems to be rank of a full genus, but he also provided meaningful slightly better, as the genus is restricted to the Ameri- illustrations of the collum, caudal body end and gonop- cas [Hoffman, 1999], whereas Monachodesmus remains od structure to further document the differences of A. largely Afrotropical, albeit a species from the Philip- serratus from Urodesmus spp. These diffferences also pines has also been included therein [e.g. Attems, 1940]. concern the presence of intercalary granulations be- We suggest treating all Afrotropical species for- tween the main crests which never occur in the African mally described in or assigned to Lophodesmus as ac- counterparts. tually belonging to Monachodesmus. Among the Afri- One new species is added here to Urodesmus. can Lophodesmus, only L. cristulifera Brolemann, 1920, In contrast, the collum in U. sexcarinatus shows from Zanzibar, and L. escherichii Silvestri, 1911, from 2+2 and 2+2 central, rather small tubercles behind an Eritrea, show small intercalary
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