MYRIAPODOLOGICA Virginia Museum of Natural History Vol. 7, No. 1 ISSN 0163-5395 December 30, 2000 A synopsis of the Telonychopodinf a tribe of Pantanalian chelodesmid millepeds (Polydesmida: Chelodesmidae) By Richard L. Hoffman ABSTICACT The tribe Telonychopodini is considered to contain three genera restricted to the southern part of the Brazilian state of Mato Grosso: Telonychopus Verhoeff, 1951, with two species {meyeri Verhoeff and klossae Manfrediodesmiis Schubart, 1949, with the single species passarellii (Schubart), and the new genus Pantanalodesmiis, based on P. marinezae, new species from the vicinity of Pocone, Mato Grosso. Keys are provided for all taxa; male genitalia are illustrated. INTRODUCTION 111 1899, the German naturalist Hermann Meyer collected a large colorful milleped in a swamp along the Rio Cubiaba at Acorizal, Mato Grosso, the first step in an interesting sequence leading to the disclosure of a group of such species, centered on the Pantanal region around Cuiaba. Eventually Meyer’s specimen found its way, along pathways unknown to me, into the hands of Karl W. Verhoeff who perceived it to be . .eine neue, aberrante Form der Rhachidesmidae''' — so aberrant that under the name Telonychopus meyeri the species became the basonym for a new family of rhacodesmoids, the Telonychopidae. This conception was based on the apparent absence of the moveable coxal structure [cannula\ that is present in 99% of known polydesmidans. To Verhoeff, thinking in terms of isolated key characters, such an absence must Scanned with permission by Virginia Tech Insect Systematics Group 2014 (www.jointedlegs.org) 2 Myriapodologica mandate a relationship with the Middle American family RJiacodesmidae, long known and characterized largely by absence of the cannula. On the basis of this single observation, Verhoeff allied the Telonychopidae in a superfamily Rhachi- desmidea along with the Rhachidesmidae [ 52 c!] and Atopogonidae (one genus and species on New Caldedonia), and proceeded to discuss the situation in a long speculative essay. Three critical issues were not confronted by Verhoeff If the lack of a cannula was indeed a shared, derived character, why did the three groups have so little in common structurally otherwise; was the absence of the cannula simply a case of unrelated, independent loss; and why would such a association have absolutely no biogeographic precedent or known parallels? The denouement to this situation came about as follows: being suspicious of Verhoeff s reasoning, I obtained in 1962 the microscope preparation of the T. meyeri gonopods and could not see how they differed substantially from the normal chelodesmid type. Because of their large size, however, Verhoeff had separated the coxae and telopodites in order for them to fit under tlie cover glass of the preparation, inflicting some damage during this process. A year later, in 1 963, Dr, Gertrud Rita Kloss, working on the nematode parasites of Brazilian millepeds, collected several large cheiodesmids from western Mato Grosso and, after their autopsy, sent them to me for identification. The riddle was then solved instantly: the gonopods matched those of meyeri very closely, except that in their intact condition, a perfectly normal cannula was present in the usual location. In all peripheral characters, this species (which was named Telonychopus klossae) also agreed with the chelodesmid norm. So much for the family Telo- nychopidae: a fantasy based on a defective specimen. In writing this information up in my 1965 paper, I gave the same historical account as that stated above, and found the most exalted status I could justify for this genus was that of a tribe endemic to the interior of South America, embracing also the genera Catharodesmus, Man- frediodesmus, and Euthydesmus. In my 1 980 Classification, I mongrelized this tribe by adding four additional (and, as I now know, misplaced) genera, which require placement in other taxa. Within the past year, however, the case has been re-opened by the discovery near Cuiba of a related, undescribed, chelodesmoid. Sent to me for identification by Prof Dr. Joachim Adis (Plon), this new material catalyzed an examination of the generic limits of Telonychopus and the results of that inquiry are set forth in the following pages. It is hoped that this initiative will be useful to Brazilian scientists working with the fauna of the Pantanal region, and wall lead to a better knowledge of the diversity of its millepeds. Hoffman; Telonychopodini 3 Family ChelodeSMIDAE Cook Tribe Telonychopodini Verhoeff Telonychopidae Verhoeff 1951, Zoo I. Anz., 146: 82. Telonychopodini Hoffman, 1965, Pap. Aviils. Zool. 17: 252; 1980, Classification of the Diplopoda, p. 151. Large chelodesmids, body length to 70 mm; paranota set high on sides, both corners rounded on segments as far back as midbody, acutely produced only on last four or five segments; ozopores in normal sequence, placed in flat peritremes continuous with segmental margins; metaterga essentially smooth to microcoriarious, with shallow transverse sulcus, no rows of tubercles; stricture deep, anterior edge sharply defined, prozonal surface coarsely reticulate; limbus with large spine-like projections ventral ly. Gonopods with large median sternal element, coxae massive, surrounding base of telopodites, much of their mass carried outside the sternal aperture; sternal apodemes large, flat, and decurved at 90° angle to coxal axis; telopodites set on coxae at a right angle, usually straight and massive, without or witli only rudimentary prefemoral process; no trace of torsion, prostatic groove visible for its entire length on mesal side. Males with low paramedian setose knobs on anterior sterna; sternum of 6th segment deeply excavate; legs without any modification. Cyphopods large and projecting ventrad outside the body, of the form shown in Figs. 12 and 13, receptacle sclerite apparently absent. As presented in my 1980 classification, this tribe was quite heterogeneous with the eight genera assigned to it. A more stringent concept now' excludes all of them except Telonychopus, Manfrediodesmus, and the new genus here described. The species of the tribe are closely similar in external features, and even the genera may not be recognized without reference to gonopod structure. Nonetheless, the latter is so diverse that annectant conditions are now difficult to postulate. lb a greater extent than in many chelodesmids, species of this tribe combine an elaborate mix of generalized and derived characters. Plesiomorphic states are manifested, e.g., in the lack of secondary sexual modifications in male sterna and legs, and the retention of prominent median gonostei nal sclerites. Specializations occur as greatly enlarged gonopod coxae, the presence of spiniform projections on the limbus, and the hypertrophied form of the cyphopods. General overall body form and the modified limbus suggest relationship with other taxa of the Parana drainage basin such as Leiodesmus and some satellite genera. Cyphopod structure is similar to that in the species of Macrocoxodesmini, but the valves face mesad rather than laterad. 4 Myriapodoiogica KEY TO GENERA OF TELONYCHOPIDAE 1. Telopodite of gonopod with an accessory subterminal process on dorsal side (Fig. 6, AP); prostatic groove carried distally on the telopodite along a lateral crest (Fig. 5, X) and visible in ventral aspect Manfrediodesmits — d’elopodite of gonopod with only two apical processes; prostatic groove confined to medial surface of telopodite and invisible in ventral aspect 2 2. Telopodite of gonopod, as seen in mesal aspect, distinctly curved dorsad, in ventral aspect, broadly conchoidal in form (Fig. 1 1); solenomere slender, sigmoidally curved (Fig. 12, S) Pantanalodesmus - Telopodite nearly straight as seen in mesal aspect, more nearly cylindrical in form (Fig. 2); solenomere short, laminate, falcate Telonychopus Genus Telonychopus Verhoeff Telonychopus Verhoeff, 1951, Zool. Anz., 146: 86. Monotypic with a new species. Type species, T. meTm Verhoeff — Hoffman, 1965, Pap. Avuls. Zool., 17: 282. Diagnosis: With the characters of the tribe, differing from the other two genera as specified in the foregoing key, the gonocoxa is also proportionately much smaller in respect to length of the telopodite. Remarks: Although the two taxa admitted to the genus can be distinguished without difUculty, the possibility may not be excluded, that they represent elements of a single polytypic species. Their relationship can be clarified when material from additional localities around Cuiaba is available for comparison. KEY TO SPECIES OF TELONYCHOPUS Telopodite of gonopod relatively stout, solenomere without triangular basal lobe on lateral side (Fig. 1) . meyeri Telopodite of gonopod relatively slender, solenomere with triangular basal lobe on lateral side (Fig. 2, L) . klossae Telonychopus meyeri Verhoeff Figure I Telonychopus meyeri Verhoeff, 1951, Zoo!. Anz., 146; 87, tigs. 1-5. Holotype (gonopods only) (ZSBS) from Acorizal, Mato Grosso, H. Meyer leg. 23 April 1899. - Hoffman, 1965, Pap. Avuls. Zool., 17: 244, fig. 1. Hoffman; Telonychopodim 5 Material: Gonopod preparation of hoiotype; location of body, if still extant, unknown. Distribution: Known so far only from the type localit>'. Telonychopus klossae Hoffman Figures 2-3, 1 5 Telonychopus klossae Hoffman, 1965, Pap. Avals. Zook, 17: 246, figs. 2-7. Male hoiotype (MZUSP) from Santo Antonio Leverger, Mato Grosso, Brasil, M, Alvarenga et a!. leg. 26 November 1963. Materiax: Male and female paratypes (VMNH) from Chapada dos Guimaraes, Mato Grosso, 1 1-14 November 1963, leg. Alvarenga, Oliveira, and Bokermann. Remarks: The original description of this species, and illustrations of the male characters, are very detailed; the cyphopods however received scant attention. I take this opportunity to provide illustrations of these structures. Generally similar to those of Pantanalodesmus marinezae, the cyphopods of klossae are relatively more robust with prominent valvular dentations (cf. Figs. 14 and 15). Distribution: Beside the two original localities, the species is known from the Universit}' of Cuiaba campus, under Cocos nucifera, M. 1. Marques leg. 15 January 1999 (Univ. Cuiaba, lo", S. I. Golovatch det.) Genus Manfrediodesmiis Sc hu bait Manfredia Schubart, 1943, Pap. Avuls. Zool. 3: 140. Monobasic with a new' species. Type species: M passarellii, by original designation. Preoccupied by Manfredia Verhoeff, 1940. Manfrediodesmus Schubart, 1949, Rev. Brasileira Biol.., 9: 18 (new name for Manfredia Schubart, 1943, ipso facto with the same type species). Diagnosis: With the characters of the tribe, distinguished by the more elaborate structure of the gonotelopodite, particularly the tripartite distal end and the location of the solenomere on a distinct ventrolateral flange for most of its length (Fig. 5). Distribution :Th is monotypic genus is known only from the southwestern part of Mato Grosso, Brazil. Manfrediodesmus passarellil (Schubart) Figures 4-6 Manfredia passarellii Schubart, 1943, Pap. Avuls. Dept. Zool. Sao Paulo, 3:140, figs. 2, 3. Holoty'pe male, MZUSP, from Sao Luiz do Caceres, Mato Grosso, E. Garbe leg,, October 1917. 6 Myriapodologica Manfrediodesmus passarellii: Schubart, 1949, Rev. Brasil. Biol., 9: 18; 19 d8. Materjal: Male holotype and topoparatype (MZUSP) examined. Comment; The more proximal of the telopodite processes is set off from the tw'o distalmost by a distinct depression on both mesal and lateral sides (Fig. 4) which is perhaps a precursor of a cingulum. PantanalodesmiiSy new genus Type SPECIES: P. marinezae, new species. Name: A neologism derived from the Pantanal region of Brazil + the combining form -desmus, widely used in this order.. Diagnosis: With the characteristics of the tribe, Distribution: Known so far only from the localit>' of the single species. Pantanalodesmus mannez(ie, new species Figures 7~I4 Material: Male holotype and female paratype (MZUSP, Sao Paulo), two male and two female paraWpes (Univ. Cuiaba), one pair of male and female paratypes in VMNH (Martinsville), MNHG (Geneve), FMNH (Chicago), MPEG (Belem), INPA (Manaus), Zool. Mus. Moscow, Coll. Adis (Plon), from Fazenda Ipiranga, km 10 on the Transpantanal Highway near Pocone, Mato Grosso, Brasil; J. Adis et al. leg. 20 October 1998. Name: For Prof. Dr. Marinez LMarques, Universidade de Cuiaba, in recognition of her contributions to knowledge of the fauna of the Pantanal region. Diagnosis: Differing from other members of the tribe in the relatively short, broad, subconchoidal, dorsally curved telopodite and sinuously curved solenomere. Holotype: Adult male, length ca. 68 mm., body broadest near anterior end, thence parallel-sided for most of length, gradually narrowed over last four segments; width of segment 2, 10.8 mm, segment 6, 12.1 mm, segment 10, 1 1.5 mm, segment 14, 1 1 .2 mm, segment 16, 10.8 mm, segment 1 8, 7.3 mm. W/L ratio at midbody, 1 7 %. Head, antennae, paraprocts, prozona and stricture deep mahogany-red dorsally, shading into lighter browm ventral ly; metaterga light orange-brown, lateral edges of paranota and a broad lunate band on caudal margin nearly yellow. Sides of metazona, sterna, and basal podomeres light yellowish-browm, the legs darker distal ly. Prozona rather coarsely textured with isodiametric mesh; stricture deep and prominent, the anterior edge sharply defined; metaterga very finely microcoriarious. with shallow transverse sulcus, but no rows of tubercles. Limbus broad, with smooth Hoffman; Telonychopodini 1 edge dorsally, ventrolaterally and ventrally becoming prominently ciliate, the individual fimbriae originating slightly proximad of edge. Paranota large, set high on sides, continuing slope of dorsum; W/H ratio at midbody ca. 70%. Paranota of segments 2-4 directed cephaloventrad, of 5-15 basically transverse, both anterior and posterior corners evenly rounded back to about 11th, thereafter gradually more angular to acutely produced by segment 16. Scapuloral rim marginal, abruptly expanded at posterior corner of poriferous segments into oval peritreme (Fig. 9), pores in normal sequence. Posterior segments of normal chelodesmoid con- figuration, epiproct acute; paraprocts with moderate depression setting off edges; hypoproct with prominent median projection and indistinct paramedian tubercles. Sides of metazona finely and evenly granulate, with low rounded knob above base of each anterior leg. Legs attached to elevated podosterna, each with deep transverse groove and emarginate posterior surface, small subcoxal knobs present on most segments. Sternal width at midbody 2.5 mm, surface sparsely setose. Legs as long as metazonal width (11.5 mm at mid body) and robust, sparsely setose basally, tarsi densely so. Sterna of anterior segments narrow, coxae of 2nd pair in contact, 3rd nearly so, segment 4 with two small, contiguous, paramedian processes, segment 5 with two pairs in similar processes slightly more separated; segment 6 broad, deeply excavated to accomodate gonopodal apices, coxae of 6th and 7th legs elongated; legs of anterior segments densely setose ventrally. Podomeres without trace of sexual modification. Segment 7 with exceptionally large, transverse gonopodal aperture consuming entire ventral surface of both pro- and metazonum, lateral ends elevated, anterior and posterior edges not raised. Gonopod coxae enormously enlarged, half of their mass held outside the aperture; a prominent, fully sclerotized median sternal element present (Fig.l 1), broadest on ventral side, abruptly narrowed dorsally; coxosternal apodeme very large, abruptly recurved. A large, densely setose coxal apophysis present above base of cannula; lateral surface with sparse smaller subapical setae. Base of telopodite largely enclosed within coxa, broadened prefemoral region concealed by coxal apophysis as in other members of the tribe, only a small digitiform prefemoral process evident. Telopodite (Figs. 11-13) small relative to coxal size, broad, subconchoidal, curved dorsad, mesal face concave; prostatic groove visible for entire length in mesal aspect; solenomere slender, sigmoid, basally looped; opposed by broad, spatulate apex (?tarsus; “parasolenomere”). Paratype; Adult female, body length ca. 70 mm, body subparallel-sided at about 1 1 mm. from segment 3 to 14, thereafter gradually narrowed posteriad. Coloration and most peripheral features exactly as described for male except paranota smaller and legs distinctly more slender. Cyphopods veiy large, extended far beyond ventral edge of segment 3, of the form shown by Fig. 14. 8 Myriapodologica Comments: At the t\pe locality, these millipeds were found beneath the decomposing leavess of Scheelea phalerata (Martins) (Arecaceae). In addition to the type series personally examined, additional material is known (Adis, pers. comm.) from Mato Grosso: Fazenda Retiro Novo (Pirizal) near Pocone, G. Brizolla & M. I. Marques leg. 27 X 1999, in soil under Scheelea phalerata (1 C', 1?,MZUSP). ACKNOWLEDGEMENTS The on-going initiative of Prof. Dr. J. Adis in transmitting new and interesting material of Diplopoda from Brazil is acknowledged with gratitude. Profs. Adis and S, I. Golovatch kindly provided prepublication review of the manuscript. Access to the type material of Manfrediodesmus was grairted by Drs. P. E. Vanzolini and J. L. M. Feme (MZUSP). This paper is a product of research supported by NSF (PEET) grant DB-9712438) to Drs. Petra Sierwald and W. A. Shear. REFERENCES Hoffman, R. L. 1965. Chelodesmid Studies. IT The status of the milliped Telonychopus meyeri Verhoeff, and of the family name Telonychopidae. Pap. Avuls. Dept. Zool. Sao Paulo, 17: 243-253, figs. 1-7. Schubart, O. 1943. Especies novas das familtas Strongylosomidae e Leptodesmidae da ordem Proterospermophora do interior dos Estados de Sao Paulo e de Mato- Grosso. Pap. Avuls. Dept. Zool. Sao Paulo, 3: 127-164, figs. 1-54. Schubart, O. 1949. Sobre os maiores “Proterospermophora” do Brasil. Rev. Brasil. Biol., 9(1): 17-24. Verhoeff, K. W. 1951. Eine neue, aberrante Form der RJiachidesmidae und die Familien der Rhachidesmidea. Zool. Anz,, 146: 81-88, figs. 1-5. Address of the author: Dr. Richard L. Hoffman Virginia Museum of Natural History Martinsville, Virginia 241 12, USA Hoffman; Telonychopodini 9 Figs. 1-3. Gonopods of telonychopodine species. 1. Telopodite of left gonopod of Telonychopus meyeri Verhoeff, mesal aspect. 2. Left gonopod of Telonychopus klossae Hoffman, mesal aspect, 3. The same gonopod, ventral aspect. Drawings from holotypes. 10 Myriapodologica Figs. 4-5. Manfrediodesmm passareUii (Schubart). 4. Gonopods sn situ, ventral aspect. 5. Left gonopod, mesal aspect. 6. Distal half of telopodite of left gonopod, lateral aspect. Abbreviations: s, solenomere; AC, accessory (?femoral) process; F, ventral flange with prostatic groove. Drawings from topoparatype (MZUSP). Hoffman; Telonychopodini 11 Figs. 7" 10. Panianaiodesmus marinezae, new species. Left paranoia of selected body segments. 7. Collum and segments 2 and 3. 8. Segment 5. 9, Segment 13. 10. Segment 13. Drawings from male topoparatype (VMNH). 12 Myriapodologica Figs, 1 1-13. Panianalodesmus nmrineiae, new species, male genitalia. 1 1. Conopods in situ, ventral aspect (only part of right gonopod shown). 12. Gonopods, dorsal aspect (only part of right gonopod shown), showing reduction of sternum dorsally. 13. Left gonopod, mesa! aspect. Drawings from male lopoparatype (VMNH). Hoffman; Telonychopodini 13 Figs. 14, 15. Female genitalia in telonychopodine species. 14. Panicmalodesmiis marinezae, female topoparatV'pe. 15. Telonychopus klossae, female paratype. Drawings in posterior (aboral) aspect, to same scale.