New cavernicolous millipeds of the family Cambalidae (Cambalidea: Spirostreptida) from Texas (U.S.A.) and Mexico

New Cavernicolous .Millipeds of the Family Cambalidae (Cambaliclea: SpirostrepLida) from Texas (U.S.A.) ancl Mexico ByNELL B.CAlJSEy1) lVilh plales 57 (1)-58 (2) Millipeds of the genus Cambala arc abundant in Texas caves. In three widely separated cave areas, there are two allopatric species, of which one is represented by two subspecies. Cilmbala speobia (Chamberlin), a troglobite, surely will also be found in Coahuila. Of the two subspecies of C.reddelli n. sp., the nominate subspecies is a troglophile that also OCCllI'S in epigean sites in l\'ew Mexico, and inornatlls is probably a troglobite. The last is of special interest because of the very weak development of the dOl'sal crests and pore crests. Mexicambala !'llssf:lli, n. gen. and sp., a troglobite from San Luis Potosi, is more highly modified by cave life than any other cambalid. These are the only cambalids that are known from the caves of western North America and }dexico. In the states east of Texas, Cambala minor (Bollman) and occasionally other congeners are collected in the entrance and twilight zone of ca ves. Cambala loomisi (HofTman) occurs in a cave in south-central Alabama. ~lost of the specimens that were studied in the preparation of this paper were collected by the biological division of the Texas Speleological SllI'vey, of which i\Ir. ,James Heddell is editor. The remaining specimens were collected by Dr. Hichard O. Albert, Mr. James K. Baker, Dr. T. C. Barr, Jr., and DI'. C. C. Holl'. I am grateful to all of these collectors. The deposition is mentioned in the section on each species. Genus Camhala Gr'ny Cambala Gray, 18:12, Insecta, in Griffith, The animal kingdom ... by Baron Cuvier, 15, pI. 135. Loomis, 19i18, Proc. U. S. N al. 1\[us., 86 : 37. Chamberlin and Hofman, 1958, U.S. Nal. Mus. Bull. 212:17:1. (Type species: .falas annalalas Say, by monoLypy.) Eclyills Chamberlin, (noL Holmgren 1855) Chamberlin, 1952, Enl. News, 63:10. (Type species: E. speobias Chamberlin, by monoLypy and original designaLion.) Eclomlls Chamberlin, 1952, En l. News, 63: 71. New Synonomy. 1) Department of Zoology and Physiology, Balon Houge, Louisiana, U. S. A. Louisiana Stale University, 238 Speleology I Causey This genus was summarized by Loomis in 1938. A needed revision is contemplated by HolTman (1958) to bring together the species that have been described since then. Carnbala diITers from Jll exicarnbala chiefly in the longer, slightly salient ventral margins of the collum, the fewer setae on the anal valves, and the bilobed coxite of the anterior gonopods. The species of Cambala diITer chiefly in body size, number of ocelli, amount of body pigment, prominence of dorsal crests and pore crests, segment on which crests begin, size and distribution of lobes on legs of matme males, presence or absence of tarsal claw of legpair 1 of males, and details of both pairs of gonopods. Dorsal crests begin on segments 2, 3, or 4. Pores begin on segment 5. A troglobite, C. speobia, lacks ocelli; all other species have several ocelli in 1 or 2 series. Two setae are on the mesial margin of each anal val ve. Article 2 of legpair 1 of males is about twice the width of article 3. One or 2 articles of 2 or more pairs of legs of mature males are enlarged. The articles that are modified are It and 5, 01' only article 4, of the following legpairs: 6 and 7, 4 Lhrough 7, and Lhe first 24 or so postgonopodal legs. Sternum of anLerior gonopods is somewhat rectangular and longer than it is wide. DisLal margin of coxite is bilobed; anteriOl' lobe bears macro setae ; flagellum is long. TelopodiLe is seLose on disLal sUl'face; distal margin is divided inLo two shallow lobes. Anterior .process of coxa of posterior gonopods is either simple 01' bilobed; iL bears cUl'ved macroseLae. Ectal process of coxa is smaller than anteriOl' process and eiLher simple or divided. A vertical series of minute spllrs is on mesial smface of coxa. TelopodiLe bears shorL, thick SPUl'Son apex. I am 11llcerLain wheLher the telopodite always consists of 2 articles; possibly Lhe nllmber is noL constanL. DistribuLion. The UniLed StaLes from western WashingLon and noLhern Idaho Lo the AtlanLic Coast. MosL abundanL in Lhe Appalachian ~lounLains. S p e c i e s. The numbel' is lIncerLain. Of Lhe to thaL were listed by Chamberlin and 1IofIman (1958), some are probably synonyms or subspecies of C. minor. Epigean, Lroglophilic, and Lrogiobitic species are included. KEY TO THE SPECmS OF CA.\IBALA IN TEXAS CAVI~S '1. Without ocelli. Maximum body width '1.9 mm. Central and southwest Texas. . . . . . . . . . . . . . . . . . . . . . . . .. C.speobia (Chamberlin) With ocelli. Maximum body width '1.2 mm. Distribution otherwise. (C.reddelli, n. sp.) 2 Speleology I 239 Causey 2. With distinct dorsal crests and pore crests. Ocelli usually in 2 series. ""VestTexas C. reddelli reddelli, n. subsp. Lacking distinct dorsal crests and pore crests. Ocelli in 1 series. Northwest Texas C.r. inornatus, n.subsp. The presence of a tarsal claw on legpair 1 of the male distinguishes the speeies of Cambala that are treated in this paper from the more eastern eongeners, including those in cast Texas. CAM BALA REDDELL!, new species Diagnosis. A polytypic species near C.washingtonensis Causey in the form of the gonopods; diners in the smaller body size, fewer segments, the smaller lobes on the pregonopodallegs, and the absence of lobes on the postgonopodal legs. Characterized especially by a pair of setae on the apex of the caudal tergite. Length 16 to 24 mm. Width 0.9 to '1.2 mm. 41 to 56 segments. From 9 to 2 ocelli are arranged in either 1 or 2 irregular rows. Dis t ri but ion. Ca ves and epigean sites in west and northwest Texas and I\'ew Mexico. It possibly occnrs also in Colorado. Subspecies. Two subspecies are descr.ibed here. They diner in the prominence of the crests and in the number of ocelli. An epigean collection from northern I\'ew Mexico is intermediate between them. This species is named for Mr . .James L. Heddell in recognition of his outstanding collections of troglobitic millipeds from Texas and Mexico. Cambala reddelli reddelli, new subspecies Figures 1-4 Diagnosis. A small-bodied troglophile that is like C. r. inornatlls in the form of the gonopods and lobation of Lhe legs; differs in the greater development of the dorsal crests and pore cresLs and the al'l'angemenL of the ocelli in 2 irregular series. Male holotype. Length 16 mm. WidLh 0.9 mm. 40 segmenLs, of which the last 2 are legless. Flesh-colored and wiLh red lateral glands when freshly preserved; became red-brown in alcohol. lIead is as in congeners. Antennae reach back to margin of segment 3; raLio of length of articles 'I thr01lgh 7 and width of arLicles 5 and 6 (in parentheses) is: 19:26:30:21:23(22):25(20):4. Ocelli, which are partly covered by collum, are in rows of abouL 3, 5. Body is not markedly narrowed behind head or swollen at segment 7. Collum (fig. 1) is a little longer than segments 2 (...truncated)