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Arthropoda Selecta 15 (1): 1921

ї ARTHROPODA SELECTA, 2006

Review of the millipede genus Macellolophus Attems, 1940 (Diplopoda: Polydesmida: Xystodesmidae) Обзор диплопод рода Macellolophus Attems, 1940 (Diplopoda: Polydesmida: Xystodesmidae) J.-P. Mauriиs1, S.I. Golovatch2 & R.L. Hoffman3 Ж.-П. Морьес1, С.И. Головач2 и Р.Л. Хоффман
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Dйpartement Systйmatique & Evolution, UMS 602, Musйum national dHistoire naturelle, 61 rue Buffon, CP 53, F-75231 Paris France. E-mail: collmill@mnhn.fr. 2 Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky pr. 33, Moscow 119071 Russia. E-mail: sgol@orc.ru. 2 Институт проблем экологии и эволюции РАН, Ленинский пр-т, 33, Москва 119071 Россия. 3 Virginia Museum of Natural History, 1001 Douglas Avenue, Martinsville, Virginia 24112, U.S.A. E-mail: rhoffman@vmng.net.

KEY WORDS: Diplopoda, Macellolophus, taxonomy, synonymy, Spain, Algeria, Morocco. КЛЮЧЕВЫЕ СЛОВА: Diplopoda, Macellolophus, таксономия, синонимия, Испания, Алжир, Марокко. ABSTRACT. The western Palaearctic millipede genus Macellolophus Attems, 1940 is shown to currently comprise perhaps a single valid species. Provisionally, this species is identifiable as M. rubromarginatus (Lucas, 1846), comb.n. ex Polydesmus (= M. excavatus Verhoeff, 1931, syn.n., = M. hispanicus Verhoeff, 1931, = M. panousei Schubart, 1960, syn.n., = M. breuili Ceuca, 1988, syn.n.), from southern Spain, Algeria and Morocco. The status of M. diadema (Gervais, 1836), from Gibraltar, must remain dubious until strict topotypes have become available for study but, judged from all available evidence, this name is very likely to represent a senior synonym of M. rubromarginatus. If so, Macellolophus is to be regarded as a mono- or oligospecific genus showing clear-cut tendencies to troglophily. РЕЗЮМЕ. Показано, что диплоподы западнопалеарктического рода Macellolophus Attems, 1940 в настоящее время, кажется, включают единственный валидный вид. Предварительно это вид определен, как M. rubromarginatus (Lucas, 1846), comb.n. ex Polydesmus (= M. excavatus Verhoeff, 1931, syn.n., = M. hispanicus Verhoeff, 1931, = M. panousei Schubart, 1960, syn.n., = M. breuili Ceuca, 1988, syn.n.) из Южной Испании, Алжира и Марокко. Статус вида M. diadema (Gervais, 1836) из Гибралтара остается неясным до тех пор, пока не будет изучен строго топотипический материал. Однако, судя по уже накопленным данным, весьма вероятно, что это название представляет собой старший синоним M. rubromarginatus. Если это так, то Macellolophus следует рассматривать в качестве моно- или олиготипического рода с явными склонностями к троглофилии.

Introduction
Originally, the genus Macellolophus was proposed invalidly by Verhoeff [1931] for two new species from southern Spain: M. excavatus Verhoeff, 1931 (male and female, from a dubious locality) and M. hispanicus Verhoeff, 1931 (female, from Cartagena). Attems [1940] validated Macellolophus by selecting M. excavatus as type species. Both Verhoeff [1931] and Attems [1940] must have been so strongly impressed by the unusual looks of Macellolophus, in both species of which the head is completely covered by the collum from above, and the paraterga are conspicuously elevated above the level of the dorsum, that this genus was treated as the sole component of the family Macellolophidae. Schubart [1960] described a third species, M. panousei Schubart, 1960, from Morocco. Hoffman & Lohmander [1968] formally transferred Polydesmus diadema Gervais, 1836, from Gibraltar, to Macellolophus, while Ceuca [1988] not only added one more congener, M. breuili Ceuca, 1988, from a cave in southern Spain but he also recorded M. excavatus from a cave in Algeria. The first attempt at a review was that of Hoffman & Lohmander [1968] who revised type material of M. excavatus and gave excellent illustrations made from the lectotype. Furthermore, the Macellolophidae has since been downgraded to the rank of a separate tribe, Macellolophini, in the family Xystodesmidae [Hoffman, 1980]. Vicente [1988] revised type material of both M. excavatus and M. hispanicus and, based on extensive samples, both epigean and cavernicolous, from various parts of southern Spain, including male topotypes of M. hispanicus, she provided sufficient evidence to formally synonymize the latter species with M. excavatus.


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J.-P. Mauriиs, S.I. Golovatch & R.L. Hoffman form dark brown, the paralectotype is pale greyish, both apparently more or less strongly faded. In addition, H.W. Brцlemann identified but likewise never published a few more samples of 'Cryptodesmus rubromarginatus' belonging to the MNHNP (Collection Myriapodes-Onychophores JB 230 & 231): Espagne, prov. Valencia, part.њ Ontenyent, Cueva de la Zarra de San Blas, Bocairent, 6-4-1913, leg. H.W. Brцlemann, 1 , 1 . Espagne, prov. Valencia, part.њ Gandia, Palma, Cueva Negra de Palma, 7-4-1913, leg. H.W. Brцlemann, 1 , 1 . Algйrie, Oranais, environs dOran, leg. L. Lйger, 1 , 1 . Algйrie, Oranais, Tlemcen, grotte dAпn Fezza, leg. E. Simon, 6 , 1 .

Furthermore, in view of the highly pronounced variation range M. excavatus appears to display as regards both somatic and gonopod structures, she also suspected that M. panousei might well prove to be another junior synonym of M. excavatus. Indeed, the discriminant analysis and the numerous additional illustrations presented by Vicente [1988] show that variation in Spanish populations of M. excavatus concerns body length (1023.9 mm in males, 1725.5 mm in females), midbody pro- and metazona (0.91.7 and 1.12.1 mm in males, 1.52.2 and 1.62.5 mm in females, respectively), coloration (from moderately dark brown with yellowish paraterga to almost blackish throughout), shape of paraterga (more or less broad and more or less rounded anterolaterally), gonopods (telopodite more or less slender, with solenomere either shorter or longer in relation to prefemur, set off from latter by a more or less well-expressed lateral sulcus, and gonopod tip from more or less distinctly emarginate to subtruncate), etc. This variation seems to be purely individual, often sex-linked but with neither evident geographical patterns nor habitat inclinations (e.g., epi- versus hypogean).

Synonymy
Re-examination of all above material shows clearly that Polydesmus rubromarginatus is a member of Macellolophus, comb.n. ex Polydesmus Latreille, 1802/ 03. Moreover, this is an undisputable senior synonym of M. excavatus. Hence the new formal synonymy below: M. excavatus Verhoeff, 1931 = M. rubromarginatus (Lucas, 1846), syn.n. According to Schubarts [1960] description of M. panousei, the only characters distinguishing this species from M. excavatus are the slightly smaller body size (length 1517 mm, width of midbody pro- and metazona 1.5 and 2.12.4 mm, respectively) and the somewhat stouter gonopods. However, as noted above, since body size, proportions of the different parts of the gonopods and even the degree of expression of the lateral sulcus demarcating the prefemoral part from the solenomere prove to vary individually a great deal, we also take the opportunity to formalize the synonymy long suspected by Vicente [1988]: M. panousei Schubart, 1960 = M. rubromarginatus (Lucas, 1846), syn.n. Reading carefully the original description of M. breuili leaves little doubt, we face still one more junior synonym of M. rubromarginatus. Indeed, Ceuca [1988] depicted a ventral view of the right gonopod of what he identified as M. excavatus taken from a cave near Tlemcen (sic!), Algeria, i.e. the terra typica of M. rubromarginatus (see also topotypic material documented above). That drawing was believed to serve as the basic argument to distinguish M. excavatus from M. breuili, the latter deriving from a cave in Mбlaga, Spain. However, this time Ceuca depicted the left gonopod in mesal view, thus making a direct comparison of both species virtually impossible. So we only can compare the gonopod drawing of M. breuili with a similar figure of the M. excavatus lectotype published by Hoffman & Lohmander [1968], with the result that both look nearly the same. The distinct lateral flange on the solenomere of M. breuili as shown by Ceuca [1988] must have simply been

Identity of Polydesmus rubromarginatus Lucas, 1846
Regrettably, the collection of the Musйum national dHistoire naturelle in Paris (MNHNP) no longer contains type material of Polydesmus diadema Gervais, 1836, to verify the identity of this species versus the other nominal members of Macellolophus. So strict topotypes from Gibraltar are crucial to finally resolve the problem. The species was described as red-brownish (cinnamon) in colour, body 25 mm long [Gervais, 1836]. Fortunately, the types of Polydesmus rubromarginatus Lucas, 1846, another enigmatic species long suspected to actually represent a Macellolophus, are still available at MNHNP (Collection Myriapodes-Onychophores JB 229). The single male syntype is herewith designated as lectotype, while the single female syntype the paralectotype. The original handwritten label given for the lectotype is hard to read but it has 'Polydesmus rubro-marginatus Lucas Tlemcen' on it, with further two handwritten labels added later by H. W. Brцlemann, 'Cryptodesmus rubro-marginatus Lucas Algйrie (Brцlemann)' and 'Cryptodesmus rubromarginatus Lucas Brцlemann det. Musйum Paris Tlemcen Lucas'. The original typewritten label attached to the paralectotype reads 'Polydesmus rubromarginatus Luc. M. Lucas Tlemcen', again with further two handwritten labels added later by H.W. Brцlemann, 'Cryptodesmus rubromarginatus Lucas Algйrie (Brцlemann)' and 'Cryptodesmus rubromarginatus Lucas Brцlemanndet. Musйum Paris Lucas Tlemcen'. This species was described as generally black with red sides of segments, 21 mm long and 2.75 mm wide. The samples derived from Oran and Tlemcen, Algeria [Lucas, 1846]. The lectotype is currently rather uni-


The millipede genus Macellolophus obscured by a slightly different angle displaying the gonopod of M. excavatus. This is clear from the illustration of the same gonopod in ventral view as presented both by Verhoeff [1931] and Hoffman & Lohmander [1968]. The tip of the M. breuili gonopod is said to be a slightly swollen lobe with a minor funnelshaped extension that marks the end of the prostatic groove. Yet this description provides no distiction whatsoever from M. excavatus! Even if the subterminal lobule of the solenomere generally characteristic of M. rubromarginatus is underdeveloped to absent in M. breuili, as it has not been mentioned by Ceuca, this alone would hardly justify separation of this species from M. rubromarginatus. The slightly different body size and proportions of M. breuili as indicated by Ceuca (males 12 mm long and 2.5 mm wide, females 14.5 mm long and 3 mm wide) do not seem sufficient for specific discrimination either. In other words, based on the original description of M. breuili alone and keeping in mind the pronounced variation range of M. rubromarginatus, there seem to be no palpable differences whatever that would justify keeping this species separate from M. rubromarginatus. To summarize, we formally propose still another new subjective synonym: M. breuili Ceuca, 1988 = M. rubromarginatus (Lucas, 1846), syn.n.

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regarded as a mono- to oligospecific genus showing clear-cut tendencies to cavernicoly. Cave-dwelling populations of M. rubromarginatus appear to be scattered more or less randomly over the entire geographical range of this species, apparently none of these being more than merely troglophilic. The documented localities of Macellolophus (= M. rubromarginatus (Lucas, 1846)) are as follows: 1: Gibraltar; 2: Tlemcen; 3: Cartagena; 4: Talasse MTane; 5: between Carratraca and Ardales; 6: Zuheros; 7: Calahonda; 8: Sierra de Cazorla; 9: Sorbas; 10: Calasparra; 11: Totana; 12: Bocairent; 13: Callosa dEn Sarria; 14: (Punta de) Moraira; 15: Ebo; 16: Jalon: 17: El Verger; 18: Atsubia (= Atzuvia); 19: Tous; 20: Dos Aguas (= Dos Aigьes); 21: Cabanes.
ACKNOWLEDGEMENTS. The help of Prof. Yves Coineau, the former director of the Laboratoire de Zoologie (Arthropodes) of the MNHNP, is greatly acknowledged for arranging a research visit of SG to MNHNP in November 1999.

References
Attems C. 1940. Myriopoda 3. Polydesmoidea III. Fam. Polydesmidae, Vanhoeffeniidae, Cryptodesmidae, Oniscodesmidae, Sphaerotrichopidae, Peridontodesmidae, Rhachidesmidae, Macellolophidae, Pandirodesmidae // Tierreich. Lfg.70. S.ixxxii+1577. Ceuca T. 1988. Sur quelques Diplopodes de la Pйninsule Ibйrique et du nord de lAfrique // Studia Univ. BabeкBolyai, Biol. T.33. P.4048. Gervais P. 1836. Note sur le genre Polydesmus de la classe des Myriapodes. Ann. Soc. ent. Fr., Sйr.1. T.5. 373381. Hoffman R.L. 1980. Classification of the Diplopoda // Mus. hist. nat. Genиve. 237 p. Hoffman R.L., Lohmander H. 1968. The Diplopoda of Turkey // Mitt. Hamburg. Zool. Mus. Inst. Bd.65. S.61121. Lucas H. 1846. Notes sur quelques nouvelles espиces dinsectes qui habitent les possessions franзaises du nord de lAfrique. Myriapodes // Rev. Zool. Soc. Cuvier. T.9. P.283289. Schubart O. 1960. Ein weiterer Beitrag zur Diplopoden-Fauna Marokkos // Bull. Soc. Sci. nat. phys. Maroc. T.40. Fasc.3. P.159232. Verhoeff K.W. 1931. Chilognathen aus dem Bergamasker Alpen und Nachbargebieten; auch ьber zwei neue Gattungen der Polydesmoidea aus Spanien und Japan // Zool. Jb., Syst. Bd.61. H.4. S.397522. Vicente M.C. 1988. Sinonimia entre las dos especies ibйricas del gйnero Macellolophus Attems, 1940 (Diplopoda, Polydesmida, Xystodesmidae) // Orsis. T.3. P.173181.

Concluding notes
The genus Macellolophus Attems, 1940 seems to currently contain only a single valid species. Provisionally, this species is identifiable as M. rubromarginatus (Lucas, 1846), comb. n. ex Polydesmus, with the following junior synonyms involved: M. excavatus Verhoeff, 1931, syn.n., = M. hispanicus Verhoeff, 1931, = M. panousei Schubart, 1960, syn.n., = M. breuili Ceuca, 1988, syn.n. The distribution of this species covers entire southern Spain, as well as northern Algeria and Morocco. The status of M. diadema (Gervais, 1836), from Gibraltar, is bound to remain dubious until strict topotypes have become available for study but, judging from all available evidence, including geographical one, this species is most likely to represent a senior synonym of M. rubromarginatus. On the other hand, there might be grounds to believe that still another, probably undescribed congener inhabits southern Spain [Spelda, in litt.]. If so, Macellolophus must be