MYRIAPODOLOGICA Virginia Museum of Natural History Vol. 5, No. 6 ISSN 0163-5395 March 1, 1998 Some necessary fine-tuning in the order Spirobolida (Spirobolidae, Messicobolidae, Atopetholidae) By Richard L. Hoffman ABSTRACT The following taxonomic and/or nomenclatorial changes are proposed: Oxo bolus Chamberlin, 1922, as a subjective junior synonym of Messi- cobohis Brolemann, 1913, Spirobolus reptans Porat, 1889, is transferred from Messicobolus to Hiltonius, Spirobolus Bollman, 1893, as a subjective jumor synonym; the gonopods of the type specimen of Atopetholus californiciis Chamberlm, 1918, and Hesperolus wheeleri Chamberlin, 1 91 8, are illustrated for the first time; the latter name is based on an immature male holotype and generic position thus uncertain although geogi'aphically the species is doubtless referable to Atopetholus\ Spirobolus vulvamis Karsch, 1881, is transfen'ed from Cyclothyrophorus to Centreius and the gonopods of the male lectotype illustrated for the first time; the gonopods of Cycloihyrophorus salvini Pocock, 1908, are illustrated and the genus confirmed as an atopetholid taxon. Atopetholus parvus Chamberlin, 1918, is redescribed from the ‘Tosf’ holotype and considered provisionally as type species of the nominal genus Tidolus. To I te coins Chamberlm, 1943, is resurrected from the synonymy of Centreius to accomodate the two species garcianus Chamberlm, 1 943, and fluvialis Loomis, 1968, APOLOGIA During compilation of a checklist of the millipeds of North and Middle America, I encountered a variety of taxonomic and nomenclatorial problems that are better addressed in an advance publication than in the text of the list itself Those which affect various taxa within the Spirobolida are treated in the following discussions. Scanned with permission by Virginia Tech Insect Systematics Group 2014 (www.jointedlegs.org) 64 Myriapodologica Family Messicobolidae Loomis Messicobolidae Loomis, 1968, Bull. U. S. Nat, Mus., 266: 81. Genus Messicobolus Brolemann Messicoholus Brolemann, 1913, Bull. Soc. Ent. France, p. 32, 1914, Ann. Soc. Ent. France, 83; 32. Type species, M godmani Pocock, by original designation. Oxobohis Chamberlin, 1922, Proc. U. S. Nat. Mus . 60 (8): 30. Type species: Oxobolus viriiis Chamberlin, by original description. New Synonymy! Apparently Oxobolus has survived for over 70 years of usage solely by default. There is no evidence that Chamberlin evaluated the earlier name Messicobolus when he was preparing his 1922 paper, either from Brolemann’s two publications or two well -illustrated treatments by Carl in 1918 and 1919. This is the more curious wTen it is noted that he did, however, cite Brolemann's 1914 “Essai’' in connection with Allopococlaa tylopus (cf 1922: 34), which suggests that he had seen it. In papers appearing later m his career, Chamberlin used both names to accomodate new' species, but witliout ever, to my knowledge, making a distinction between them aside from that implied by their geographic provenence. Messicobolus was used for species collected in Mexico, Oxobolus for those from further south. Nor was the situation considered by other workers, e.g. Causey (1954) and Loomis (1966, 1968). In comparing both specimens and literature accounts, 1 can find no differences whatever, at the generic level, between species placed in both the "genera"^ mentioned, and herewith consider Oxobolus a junior subjective synonym of Messicobolus . Family Spirobolidae Bollman Genus Hiltonius Chamberlin Hiltonius Chamberlin, 1918, Proc. Biol. Soc. Washington, 37: 166. Type species: H.pulchrus Chamberlin, by original designation. — Keeton, 1960, Mem. American Ent. Soc., 17: 97. . Hiltonius reptans (von Porat), new combination Spirobolus reptans von Porat, 1889, Ann Soc. Ent. Belgique, 32: 250. Two female syntypes (Inst. Roy. Sci. Nat. Belg.) from Guanajuato [Mexico] (E, Duges leg.). Spirobolus callipus Bollman, 1893, Bull. U. S. Nat. Mus., 43: 191.(7 and ? syntypes (formerly USNM, presumed lost), from Guanajuato, Mexico (E. Duges, leg.). New synonymy! Spirobolus reptans: Pocock, 1908, Biol. Centr.-Amer., Diplop,, p. 83. Hiltonius callipus: Keeton, 1960, Mem. American Entom. Soc., 17: 114. Messicobolus reptans: Loomis, 1968, Bull. U. S. Nat. Mus., 266; 84. Hoffman: Spirobolida 65 The species represented by the two names cited above has remained enigmatic for over a century but I think the veil of obscurity may be lifted by appeal to circumstantial evidence. Bollman’s posthumously published description of callipus had been wjitten prior to his death m 1889. thus he was luiawarc of Porat's account of reptans, based on material collected at the same place by the same person who obtained the types of callipus. Insofar as I know, neither name has been mentioned in subsequent literature aside from Pocock's entry' for rep tans in the ""Biologia” until Keeton (i960: 1 14) dedueed that callipus was referable to Hiltonius and Loomis (1968: 84) arbitrarily located replans in Messicobolus. Actually, Bollman's description is fairly precise and Keeton's interpretation was well-founded. But a point by point comparison of Bollman's text with that of Porat shows a remarkable concordance, which taken into consideration with the origin of the two type senes, leaves little doubt that the two names are based upon the same species. Family Atopetholidae Chamberlin Information is provided here only to document several changes proposed in the 1980 ‘"Ciassificatioifk Genus Atopetholus Chamberlin Atopetholus Chamberlin, 1918, Proc. Biol. Soc. Washington, 31: 167. T>pe species: A.calfornicus Chamberlin, by original designation. Hesperolus Chamberlin. 1918, Proc. Biol. Soc. Washington, 37: 169. Type species: H.wheelen Chamberlin, by original designation. - Hoffman, 1980, Classification of the Diplopoda, p. 77 (as junior synonym of Afopetholus) . The identity and status of the two genera cited above remained inferential ever since their proposal, since the type species of both were not illustrated and the tvpe material itself was missing from the Museum of Comparative Zoologvy the ostensible depository How ever, following the demise of Professor R. V. Chamberlin and transfer of his myriapod collections to the U. S. National Museiun of Natural History.', the entire material of Diplopoda was placed in my hands for curation and organization. During this activity a considerable number of types long missing from several museum collections and feared to be lost, began to emerge from the intenors of large jars of miscellaneous samples (1 am convinced that Prof. Chamberlin forgot that he even had such material). 1 took the occasion to study and illustrate many of these specimens before retunimg them to the rightful ow^ner musemns, and among the lumiber were several of the enigmatic atopetholids published in the 1918 paper. 66 Myriapodologica Figs. 1-3. Atopetholus califoniicus Chamberlin. 1, right side of anterior gonopods, posterior aspect. 2, left postenor gonopod, posterior view. 3, left side of anterior gonopods, ventral aspect, showing shape of telopodite process (p). Drawings from holotype. 1 provide here gonopod di'awings for the type species of both generic names listed above. By good fortune, A. californicus is clearly congeneric with the various species subsequently described in this genus but not conspecific with any of them, as indicated by tlie details shown in Figures 1-3. We are less fortunate in the case of Hesperolns wheeleri. The male holoUpe proved to be immature, and the gonopods show little more than famih' le\nl features. The provenance of the specimen (Santa Ynez Mountains, California) from a region Hoffman; Spirobolida 67 so far known to be inhabited by only one genus of atopetholids suggests that wheeleri w ill prove to be m Atopetholus when mature male topot>pes have been studied For possible confimiatory \ alue. I gi^'e here drawings made from the holotype (figs. 4-6). Figs. 4-6, Hesperolus wheeleri Chamberlin. 4, left side of antenor gonopods, anterior aspect. 5, led side of anterior gonopods, posterior aspect. 6. posterior gonopod, posterior aspect. 68 Myriapodologi ca Genus Centrelus Cook Centrelus Cook, 1911, Proc. U. S. Nat. Mus., 40; 154. Type species: C. falcatus Cook, by original designation. Smissurobolus Carl, 1918, Rev. Suisse Zool,, 27; 389. Type species; Julus me farms DeSaussure, 1860, by original designation. Synonymized by Hoffman, 1980. Nec: Centrelus, sensu Hoffman & Orcutt, 1960: 123, and Loomis, 1968: 79. The holotype of C. falcatus, mentioned as missing from the USNM collection already in 1960, has never been located, mandating the collection of topotypes at Guanajuato, Mexico for an defimtive concept of the status of this species and the genus based on it: Hoffman & Orcutt (1960; 123) attempted to reconcile the verbal account of Centrelus with some of the atopetholids then known to them, with completely incorrect conclusions. Subsequent experience with Mexican atopetholids led, during preparation of the “Classification'' in 1978, to a complete reconsideration of the two names cited above, taking into account stmctural features mentioned bv the two authors as well as biogeographic probabilities, led me to propose the indicated generic synonymy. As a result, the generic name Toltecolus (Chamberlin, 1943) was revived for the species treated as Centrelus by Hoffman & Orcutt (1960) and Loomis (1968) (it is a curious fact that Loomis consistently misspelled the name as Centre llus in several papers, although using the single 'T” in correspondence). Although the examination of male topotypes of falcatus from Guanajuato is prerequisite for a final disposition of the name, it now^ appears that about eight nominal species can be referred to Centrelus^ on the basis of the elongated coxal processes of the 3rd leg pair and simple '"canoe-shaped" telopodites of the posterior gonopods. A list of these species is given below^ to establish the new' combinations. I think it likely that zacatecus may be removed to a different genus when the gonopods have been exammed carefully, and the same may be true for nigrescens. Species: C. boreus (Loomis, 1966); C. falcatus Cook, 1911; C. neglectus (Carl, 1919); C. nietanus (DeSaussure, 1860); C. nigrescens (Chamberlin, 1923); C. spinosus (Loomis, 1968); C. vulvanus (Karsch, 1881), and C. zacatecus (Chamberlin, 1947). Centrelus vulvanus (Karsch), new combination Figures 7-10. Spiroboius vulvanus Karsch, 1881, Zeitschr. Ges. Naturwiss. (3) 6; 55. 2cf(7 1? ST (ZMB 882, vidi!) from Puebla (probably the city), Mexico. One of the males has been isolated and labeled as the lectotype, the otlier tw'o speemrens as lectoparatypes. Cyclothyrophorus vulvanus: Pocock, 1908, Biol Centr.-Amer., Diplopoda, p. 86. Hoffman: Spirobolida 69 Cycloihyrophorus valvanns [sic!]; Loomis, 1968, Bull. U. S. Nat, Mus. 266, p, 80. Working oiiK’ from Karsch’s verbal description, Pocock showed remarkable insight in considering this species to be closely related to DeSaussure's Spiroholus nietanus, which he likewise knew only from a short description. Gonopod drawings that I made from the lectoh^e of vidvanus indicate that the posterior gonopods are vei>' similar to those of metanus as illustrated by Carl (1919: fig. 16), and the relative Figs, 7-10. Centrelus vuhanus (Kai'sch). 7, right side of anterior gonopods, anterior aspect. 8, lelt posterior gonopod, postenor aspect, 9, coxae and coxal processes of 3rd pair of legs, oblique posterolateral aspect. 10, coxa of 4lh pair of legs, postenor aspect. Drawings from leciotype, fig. 9 enlarged. 70 Mynapodol ogica proximity of Puebla to Cuernavaca admits the possibility that the two names are synonyms The superficial similarity of the coxal processes of the third legs to the posterior gonopods is noteworth> : both appendages are simple, falcate, and deepK concave on one surface. The anterior gonopods (Fig. 7) are notably for the extensive development of membrane on the anterior surface of the coxae, the distinct separation between the transverse clement of the sternum and the ventrally-produced median triangular part. An especially curious feature is the apparent coalescence of the coxa with the teiopodite (unless it can be demonstrated that the coxa is bounded medially by the abundant membranous folds, m which case the teiopodite w'ould be unusual for its long proximal extension beside the stemmn) The elongate apical processes of the 3nd coxae of males, curv’ed poster! ad and concave on their posterior side, are illustrated (Fig. 9). Genus Cydothyrophorus Pocock CycUnhyrophorus Pocock, 1908, Biol, Centr.-Amer Diplopoda, p. 83. Type species; C sahim Pocock. by original designation. This genus of somewTat unusual atopetholids has been essentially an orphan for most of its existence, not being assigned to its correct family until the 1968 Checklist by Loomis. Working solely from Pocock 's origmal description. Hoffman & Orcutt (1960: ] 13) decided that the genus was either not an atopelholid or a member of a disjunct subfamily group. But onl>’ a few years later, I received freshly collected material from Colima which agreed very closely with Pocock’s account of C. salvini, and which permitted a close examination of gonopod structure. On becoming convinced that tlie species is in fact an atopetholid, I transmitted the information to Mr. Loomis, who w'as at that time compiling his 1968 Checklist, and Cyclothyro- phorus thereupon came to rest in its correct position. Pocock had included three other Mexican spiroboloids in his genus, entirely on the basis of published verbal descriptions. One of them, Julus nietemus DeSaussure, was later redescribed by Carl (1919: 390), who designated it the type species of the new genus Saussurobohis. The second, Spirobolus heieropygus DeSaussure & Hiunbcit, was never mentioned by Carl (although the type material must also have been at Geneve), and in the lack of definite information, the species was allowed to phorese as a species of Cydothyrophorus in the Loomis Checklist (1968: 80). The third was Spirobolus vulvcmus Karsch, described from Puebla and never mentioned by anybody after Pocock’s diffident inclusion, until Loomis continued its sojourn in Cydothyrophorus. Eventually I studied the male holotype in the Berlin museum, and found that this species is probably referable to Centrelus, q.v. the species entry' above under that genus.. Although the gonopods embody the traditional atopetholid characteristics, the Hoffman: Spirobolida 71 combination of extensive, lightly sclerotized intercoxal membrane and the apically enlarged and ramose posterior gonopod imparts a distinctive generic personality to C. salvini. Despite special attention to the point, I was unable to establish the occurrence of a prostatic groove, or its equivalent, on the posterior gonopod. Cyclothyrophorus salvini Pocock Figures 11-14 Figs. 11-14. CycloibyrophofiAs salvini 1 1, nght gonopods, anterior aspect. 12,righl gonopods, postenor aspect. 13, left posterior gonopod, posterior aspect. 14, the same gonopod, apical third enlarged, anterior aspect. 72 Myriapodologi ca Cyclothyrophoms salvmi Pocock, 1908, Biol. Centr.-Amer., Diplopoda, p. 84, pi. 7, figs. 6a-d. Type material presumed lost, from Amula, Edo Guerrero, Mexico - Loomis, 1968. Bull. U. S. Nat Miis. 266: 80. Like most of the new miliipeds described in the “Biologia'' after Pocock's career change (to become superintendent of the London zoo), the tvpes (and only known specimens of this species) were never returned to the British Museum and have to be considered as lost. In 1968 it w'as my good fortune to receive extensive collections of Mexican miliipeds from Drs. G, E. Ball and D. R. Whitehead, and this material contained several males wbich matched the original description of saivini in everv respect. Gonopod illustrations made from one of the males (from 19 miles north of Ciudad Colima. July 26, 1963. in WINH) are reproduced here as Figures 1 M4, Despite previous doubts (Hoffman & Orcutt, 1960: 113) about the correct placement of Cyciothyrophorus\ the Colima material was adequate to demonstrate the atopetholid facies of the type species. The Colima population extends the known range of saivini approximately 400 km northwest along the Sierra Madre del Sur. Genus Tidolus Chamberlin Tidolus Chamberlin, 1949, Joiirn. Washington Acad. Sci., 43: 169. Type species: Atopetholus parvus Chamberlin, 1918, by original designation. The generic diagnosis of this taxon was based solely upon the brief verbal description of A. parvus published in 1918, and without better knowledge of the gonopod structure, was essentially meaningless. Ironically, it appears that Professor Chamberlin actually had the holotype of parvus m his possession since the mid- 1920s, misplaced in a large jar of imscellaneous material taken aw'ay from the Museum of Comparative Zoologv'. When curating the diplopod material after its transfer to the National Museum of Natural History', I eventually came across this specimen and made some drawings of the gonopods before returning it to the Harvard museum. It is clear that parvus - lacking the accessory^ process on the telopodite of the anterior gonopod - is not a species of Atopetholus It is less obvious how the species, and its nominal genus, remain to be distinguished from otlier regional genera such as Waitchelus. Pending a revision of the family, I recognize Tidolus provisionally as a valid monotypic genus. Tidolus parvus (Chamberlin) Figs. 15-18 Atopetholus parvus Chamberlin, 1918, Proc. Biol. Soc. Washington, 31: 170. o' HT (MCZ), from Claremont, Los Angeles Co., California. Hoffman: Spirobolida 73 Tidoius parvus: Chamberlin, 1949, Journ. Washington Acad Sci., 39: 169. - Hoffman & Orcutt, 1960, Proc. U. S. Nat. Mus., 11 1; 1 14. The anterior gonopods represent the configuration t>pical of the Atopetholinae, with a distinct, sclerotized median sternal projection essentially continuous with the slightly arched basal sternal element (Fig. 15). Nor does this gonopod show any generic apomorphy in posterior aspect (Fig. 16). The posterior gonopod however is singular in being bent at nearly a right angle near its midlength, with a prominent bidentate lobe on its inner surface just proximad to the curv ature (Figs. 17, 18). Among the documents which I inlicritcd along with the collection of H. F. Loomis after his demise in 1976 is a complete description and gonopod drawings of a species identified (and I think correctly) as parvus. This information was obtained from two male specimens taken by 0. F. Cook in Topango (or Topanga) Canyon, northwest of Santa Monica, California, on 30 December 1927. I have no knowledge where these specimens may be, if still extant, as they w'crc not in the material that remained in the Loomis collection after he dispersed much of it to several museums. But 1 consider the illustrations accurate enough to be dcfmitive. Topanga Canyon is approximately 50 miles west of Claremont, suggesting that parvus ma> be/have been widespread - if sporadic - in the Los Angeles region Figs. 15-16. Tidoius parvus (Chamberlin), 15, nght side of anterior gonopods, anterior aspect. 16, right side of anterior gonopods, posterior aspect. 74 Myriapodologica Figs. 17-18. Ti dolus panm (Chamberlin). 1 7, right postenor gonopod, posterior aspect. 1 8, The same gonopod, oblique anterior aspect, enlarged. Drawings from holotype, Genus Toltecolus Chamberlin Toltecolus Chamberlin, 1943, Bull. Univ. Utah, 34(7): 27. Type species; T. garcianus Chamberlin, by original designation. Centrelus [nec Cook, 191 IJ Hoffman & Orcutt, 1960, Proc. U. S. Nat. Mus. Ill: 123, Incorrect syiion^miy! One result of the re-assessment of Centrelus (q.v., supra) was the necessity to resurrect Toltecolus as the valid group name for a eureline incorrectly assigned to Centrelus,, this had been done without explanation in the “Classification”, p. 77. The general facies of body form and posterior gonopod of both T. kerrensis and T. fluvialis strongly resemble those of Eurelus soleatus. Probably the single feature most supportive of separate generic status for Toltecolus is the occurrence of a distinct, tliin fleshy flap on the antenor side of the gonocoxal apex. The development of large marginal spines on the lower metazonal margins is a relative character, probably of only specific value. For the present I retain Toltecolus as a recognizable generic taxon but without the conviction that another opinion might award the same status to the two species involved. Hoffman: Spirobolida 75 RL'FERENCES Bollman, C. H. 1893. Notes upon a collection of myriapods belonging to the U. S National Museum, pp, 190-200, in: The Mynapoda of North America. Bull. U. S. Nat. Mus.,46: 1-210, Brolemann, H. W. 1913. Un nouveau systeme de Spirobolides. Bull. Soc. Entom, France for 1913: 476-478. Brolemann. H. W. 1914. Etude sur les Spirobolides. Ann, Soc. Entom France, 83; 1-38. Carl, J. 1918, Miscellanees diplopodologiques. Rev. suisse Zool., 26; 417-468. CarL J, 1919. Revision de quelques Spirobolides du Museum de Geneve. Rev, suisse Zool., 27; 377-404. Causey, N. B. 1954. New Mexican and Venezuelan millipeds in the collection of the Illinois State Natural Histon' Survey. Proc. Biol. Soc. Washington, 67: 55-68. Chamberlin, R, V. 1918. New spiroboloid diplopods. Proc. Biol. Soc. Washington, 31: 165-170. Chamberlin, R. V. 1922. The millipeds of Central America. Proc. U. S. Nat. Mus., 60(8); 1-75. Chamberlin, R. V. 1943. On Mexican millipeds. Bull. Univ. Utah, 34(7); 1-103. Chamberlin, R. V, 1949. On some western millipeds of the order Spirobolida. Joiirn. Washington Acad, Sci., 39: 163-169. Cook, O. F. 1911. Notes on the distribution of millipeds in southern Texas, with descriptions of new genera and species from Texas, Arizona, Mexico, and Costa Rica. Proc. U. S. Nat. Mus., 40: 147-167. Hoffman, R, L. 1980. Classification of the Diplopoda, Geneve, Museum d’Histoirc Naturelle. 236 pp. Hoffman, R. L. & B. S, Orcutt, i960. A synopsis of the Atopetholidae, a family of spiroboloid millipeds. Proc. U. S. Nat. Mus., Ill: 95-166. Karsch, F. 1881. Neue Juliden des Berliner-Museums, als Prodromus einer Juliden- Monographie. Zeitschr. Natunv., 54: 1-79. 76 Myriapodologica Keeton, W. T. 1960. A taxonomic study of the milliped family Spirobolidae (Diplo- poda: Spirobolida). Mem. American Entom, Soc., 17: 1-146. Loomis, H. F. 1968. A checklist of the millipeds of Mexico and Central America. Bull. U. S. Nat. Mus., 266: 1-137. von Porat, C. O. 1889. Ober einige exotischen luhden des BRisster-Museums. Ann. Soc. Entom. Belgique. 32: 205-256. Pocock, R. 1, 1903-1910, Diplopoda, m: Biologia Centrali- Americana, Chilopoda and Diplopoda, pp. 41-217 [fasicle treating spiroboloids dated 1908], London, Taylor & Francis. Address of the author: Dr. Richard L. Hoffman Virginia Museum of Natural History 1001 Douglas Avenue Martinsville, VA 241 12, USA