Great Basin Naturalist
Volume 57 Number 3 Article 5
7-31-1997
Distribution of the milliped Tylobolus utahensis Chamberlin, with remarks on T. fredericksoni (Causey) (Spirobolida: Spirobolidae)
Rowland M. Shelley North Carolina State Museum of Natural Sciences, Raleigh, North Carolina
Selena B. Bauer North Carolina State Museum of Natural Sciences, Raleigh, North Carolina
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Recommended Citation Shelley, Rowland M. and Bauer, Selena B. (1997) "Distribution of the milliped Tylobolus utahensis Chamberlin, with remarks on T. fredericksoni (Causey) (Spirobolida: Spirobolidae)," Great Basin Naturalist: Vol. 57 : No. 3 , Article 5. Available at: https://scholarsarchive.byu.edu/gbn/vol57/iss3/5
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DISTRIBUTION OF THE MILLIPED TYLOBOLUS UTAHENSIS CHAMBERLIN, WITH REMARKS ON T. FREDERICKSONI (CAUSEY) (SPIROBOLIDA: SPIROBOLIDAE)
Rowland M. Shelleyl fmd Selena B. Bauerl
ABSTRACT.-Tyloholus utahensis Chamberlin, the only representative of the genus occurring in the southwestern deserts, ranges from central Inya County, California, to the western periphelY of Kane County, Utah. This distribution roughly corresponds to the northern limit ofthe Mojave Desert ecosystem and is also shown by the milliped Piedolus utus Chamberlin (Spirobolida: Atopetholidae) and the centipede Theatops posticus (Say) (Scolopendl'Omorpha: Clyptopidae). Tyloholus frederi{;ksoni Causey, ostensibly from Douglas County, Kansas, is designated a "nomen duhium." and disregarded pending collection of fresh material; Na:rceus gordanus (Chamberlin) is deleted from South Carolina and Tennessee. T.lJloboms tmeigern~ (Wood) occurs north of the Columbia River in Klickitat County, Washington, and Hiltonius thehanus Chamberlin is referrable to Onychelus Cook in the family Atopetholidae.
Key words: Tylobolus; T utahensis; T. fredericksoni; Narcous americanus; Washington County, Utah; Inyo County, California; Hiltonius.
The subfamily Tylobolinae of the diplopod questionable reports of an additional species, family Spirobolidae comprises 2 geuera, Tylo T. fmdericksoni (Causey), in Douglas County, bolus Cook aud Hiltonius Chamberlin. Keeton Kansas, on the eastern periphery of the Cen (1960) revised both taxa, recognizing 3 species tral Plains. We report 37 additional samples of of Hiltonius in southwestern California and T. utahensis that expaud its range some 300 mi adjacent Baja California Norte, and 2 species (480 km) westward to central Inyo County, in mainland Mexico that range northward into California, east ofthe Sierras, and provide data Santa Cruz County, Arizona (Keeton 1960, on segment uumbers, lengths, aud widths. We Shelley 1995, plus unreported samples exam also designate T. fredericksoni as a "nomen ined by the 1st author). The 6th species, H. the dubium" because there is no recent, indis banus Chamberlin, occurring at Tbeba, Mari putable evidence that a representative of this copa County, Ariwna, is referrable to OnycluJlus genus occurs east of the area occupied by T. Cook in the family Atopetholidae, as the 1st utahensi,. The distribution of the Tylobolinae author has learned from examining the female in the United States and the adjacent periph holotype at NMNH (see acronyms below). In ery of Mexico is shown in Figure L Acronyms a supplemental paper on the California repre of sources of preserved study material are as sentatives of Tylobolus, Keeton (1966) cited 6 follows: species west of the crest of the Sierra Nevada, BYU-Montc L. Bean Life Science Museum, 1 of which, T. uncigerus (Wood), ranged north Brigham Young University, Provo, UT ward to the Columbia River at Portlaud, Ore CAS-California Academy of Sciences, San gon; we extend its distribution into the adja Francisco cent periphery of Washington based on a male CDFA-California Department of Food and and female from Klickitat, Klickitat County, at Agriculture, Sacramento FSCA. Tylobolus extends southward into the DC--Life Scieuces Division, Dixie College, adjacent fringe of Baja California Norte (Boll St. George, UT man 1888, Keeton 1960, Loomis 1968) and, like FSCA-Florida State Collection of Arthro Hiltonius, occupies the western interior, as T. pods, Gainesville utahensis Chamberlin inhabits Zion National LACMNH-Los Angeles County Museum Park, Washington County, Utah. There are also of Natural History, Los Angeles, CA
1North ('~1rolinu State Museum ofNatural Scien""s, Bux 2.9555, n"leigh, NC 27626·0555.
231 232 GREAT BASIN NATURALIST [Volume 57
1924 at an unknown location in Zion National Park, Washington County, Utah. DIAGNOSIs.-Metazonal striae terminating ventrolaterad, well below level of ozopores; anterior gonopod telopodite apically blunt and rounded, not uncinate; posterior ganopod tibiotarsus forming right angle with distal pro jection (Figs. 2-4). VARIATION.-New material agrees closely with the holotype. The posterior gonopod tibiotarsus is slightly broader and blunter, and the denticles, which Causey (1955) and Keeton (1960) termed "prickles," cover larger areas on both the anterior and posterior gonopods. In juveniles the distal projection of the posterior gonopod is less angular, extending submediad L rather than downward or dorsad. We reexam A 1 ined the holotype and found the denticles to be stronger than shown by Keeton (1960, figs. , ,J 262-264); there is also a pronounced line or , ridge of these along the ventral surface of the I posterior gonopod telopodite that he did not show (Fig. 5). In his supplemental work on California species, Keeton (1966) tabulated meristic and Fig. L Distribution of the Tylobolinae in the United morphometric data; we present such data for States and adjacent periphery of Mexico. Solid lines (A), T utahensis (Table 1) to provide parallel accounts 'f'lloholus; dashed lines (B), Hiltonius. for all species of rylobolu.s. On the average, females are slightly longer and broader than males; adults are equivalent in length through MCZ-Museum of Comparative Zoology, Harvard University, Cambridge, MA out the range, but those on the east are nar NCSM-North Carolina State Museum of rower. The adult segment number varies from Natural Sciences, Raleigh 51 to 57, as all individuals with > 52 segments NMNH-National Museum of Natural have no legless segments except the epiproct. History, Smithsonian Institution, Wash Three females and 2 males have 51 segments, ington, DC none legless, while 3 males and 1 female with SEM-Snow Entomological Museum, Uni this count have 1 or 3 legless segments; all versity ofKansas, Lawrence individuals with < 50 segments have at least 1 UCD-Bohart Entomological Museum, without appendages. University ofCalifornia at Davis DISTRIBUTION.-Occurring from the Inyo UGA-University of Georgia Museum of Mountains on the eastern side of Owens Val Natural History, Athens. ley, Inyo County, California, to the eastern edge of Zion National Park in western Kane 'rqlobolu.s utahensis Chamberlin County, Utah, a distance of approximately 300 (Fig,. 2-5) mi (480 km; Figs. 1, 6). The milliped should rylobolu.s utahensis Chamberlin, 1925:60-61. also be expected along the Virgin River in the Keeton, 1960:131-132, figs. 262-264. northwestern corner of Mohave County, Ari Califomibolu.s utahensis: Chamberlin, 1949: zona; along this river and in other parts of 166. Chamberlin and Hoffman, 1958:162. northern Clark County, Nevada; and through out most of Lincoln County, Nevada, particu TYPE SPECIMENs.-Male holotype, female larly in canyons and gorges east of Caliente, allotype, and 2 male and 1 female paratypes Panacea, and Pioche. Specimens in Inyo (MCZ) collected by R.V Chamberlin in May County were taken in pitfall traps at elevations 1997J DISTRIBUTION OF TYLOBOWS UTAHENS!S 233
. ~-~--.-...... ~------.--....
5 2
0'··.'., ,.;":'-:-" '...... •...., ~~~.,:.'',' "
3
Figs. 2-5. Tylobolu.s utalwnsis: 2, anterior gonopods of a male from Saline Valier, Inyo County, California, anterior view; 3. left posterior gonopod of the same, anterior view; 4, the same, caudal view; 5, right posterior gonopod ofholo type. anterior view. Scale line =: 1.00 nun for aJlfigures. of2296-6560 ft. In addition to the types, spec 40 km) S Saline Valley, 2M, 29 April 1975, imens were examined as follows: A.R Hardy (CDFA), and Death Valley at Pk, CALlFORNlA, Inyo Co., Inyo Mts, Lead Can Panamint Mts, Johnson Cyn, F, 2 June 1961, yon (Cyn), F, 9 March-13 August 1981, D. R Waner ( MNH). Giuliani (CAS), Inyo Mts, Hunter Cyn, M, F, NEVADA: Nye Go., Nuclear Test Site, Ranier 9 June 1980-26 May 1981, D. Giuliani (CAS); Mesa, 2M, juv, date unknown, D.B. Thomas Inyo Mts, Willow Cr, F, 16 September 1976, (LACMNH, NCSM). collector unknown (LACMNH); Saline Valley, UTAH: Washington Co., 12 mi (19.2 km) NW 11 different stations but exact locations un St. George, nr Baker Darn, 3M, 2F, 16 April known, lOM, 17F, juv, 5 April 1959-14 June 1971, B. K. Carrell (DC), Pine Valley, F, 26 May 1960, B. Banta (CAS); Saline Valley, 10 stations 1971, Prothero (DC); Snow Cyn St Pk, M, 16 along Grapevine Cyn Rd but exact locations April 1982, RW Baumann (BYU); Pintura, F, 8 unknown, 16M, 6F, 15 August 1959-7 May March 1941, J. & W. Ivie (NMNH), Motaqua, 1960, B. Banta (CAS); 21 and 25 mi (33.6 and M, 17 April 1933, M. Zuie (NMNH), Oak Grove 234 GREAT BASIN ATUMUST [Volume 57
TAIJLE ]. Meristic lmd mOT})hOlOehic dara on Tylnbolus utahensis. Individuals are listed in d~ndjng order according to segment counts, which include the epiproct; the number orIegless segments in addition to the epiproct, if any, is shown in parentheses. Measurements l:I.fC in mm. Averages are for specimens \vith no additional legless segments. Males -Females Segments Length Max. width Segment!!" Length Max. width CA, Hunter Canyon 5[([) 41.7 4.0 49([) 35.8 3.9
CA, Lead Canyon 50(2) 27.4 3.[
CA, \'Villo\\' Creek 51(3) 36.1 3.4
CA. Sal.ill~ Valley 57 70.3 5.1 55 86.7 8.3 " " 55 73.S 5.6 55 48.8 5.6 " " 55 72.7 7.0 54 76.4 7A " " 55 67.2 7.2 54 71.4 6.6 " " 515 62.S 6.8 54 65 ..\ 6.6 " " 54 62.6 6.8 54 64.9 7.2 " " 54 59.3 6.0 54 62..1 6.6 " " 53 65.1 li8 54 61.2 7.6 " " 53 54.2 5.5 53 74.2 8.2 " • 53 48.8 5.6 53 71.0 6.1 " " 52(1) 46.2 5.[ 53 60.8 6.7 " " 52 64.8 6.5 51 47.1 5.3 " " 51 46.1 5.5 46(4) 22.6 2.8
Averages 63.8 6.7 64.4 6.7
CA, Grapevine Canyon Road 5.\ 60.3 5.9 .56 65.2 7.7 " " .\5 59.6 6.2 .56 50.6 5.2 " " 515 59.1 5.7 54 720 7.6 " " 55 48.0 5.2 54 63.S 6.7 " " 54 60.4 6.1 53 66.8 7.9 " " 54 56.l 6.0 53 64.6 6.9 " " 54 55.0 6.2 52 56.3 5.9 " " 53 77.8 6.7 " " 53 56.4 6.1 " " '90, 68.9 7.1 " " 52 58.3 7.0 " " 52 57.8 6.3 • " 52 57.6 6.0 Averages 59.6 6.2 62.8 6.8
NY, Ranier Mtlsn 53 59.2 5.7
UT, Zion National Park 53 62.1 5.3 53 54.8 4.7 " " 52 67.5 5.5 52 76.9 6.7 " " 52 66.8 5.5 52 71.5 6.7 " " 52 65.3 5.6 52 48.8 4.1 " " 52 48.8 4.1 51 64.3 6.2 " " 51 67.5 5.8 " " 51 49.l 4.5 " " 51(3) 36.5 0.0' - " " 51(3) 31.2 3.1 Averages 61.0 5.2 63.3 5.7 1997] DISTRIBUTION OF TYWBOWS UTAHENSIS 235
blechrostriatus Shelley and Bauer (Julida: Pae romopodidae), occurring in Saline Valley and along the eastern slope of the Sierra Nevada (Shelley and Bauer 1997), and Idrionaria dineh Shelley, kuown only from Washington County •• • • •• (Shelley 1996). •., • • • Tylobolus fredericksoni (Causey) 6 • Spirobolus uncigerus Wood: Cragin, 1885:145. Kenyon, 1893:16. Tylobolus uncigerus: Gunthorp, 1913:164; 1921:88. Califomibolus fredericksoni Causey, 1955:78, 80, figs. Ie, 4-5. Chamberlin and Hoffman, 1958:161. Tylobolus fredericksoni: Keeton, 1960:132-133. Fig. 6. Distribution of Tylobolus utalwnsis. For over a century records have existed of a mysterious tyloboline on the eastern periph Cpgd, F, 2 June 1988, L.L. Smithee (BYU); and ery ofthe Central Plains. The confusion began Zion Nat Pk, exact site unknown, MM, FF, with Cragin (1885), who reported Spirobolus May 1924, \eM. Tanner (NCSM), 3F, 1927, "uncigerus Wood (?)" from Topeka, Shawnee A.M.D. (NMNH), M, July 1931, w.J. Gertsch County, Kansas, stating that there were "impor (NMNH), and behind visitor's center, F, 30 tant differences" with uncigerus, but the speci April 1991, C.S. Crawford (NCSM). Kane Co., men was closer to this species than to any other. Zion Nat Pk, east entrance, M, 3 March 1967, Kenyon (1895) repeated Cragin's citation, in G.R Knowlton (NMNH). cluding the parenthetical question mark, for an REMARKs.-Tylobolus utalwnsis demonstrates individual from Weeping Water, Cass County, 2 important distribution patterns among North Nebraska. He also reported the abundant east American myriapods. The occurrences of the Nearctic spirobolid, Narceus aTMricanus (Beau genus and subfamily in coastal California and vois) [cited as Spirobolus marginatus (Say)], southwestern Utah are reiterated by the tribe from adjacent Sarpy County. Thus, Kenyon's Tynommatini and subfamily Tynommatinae in record of uncigerus is plausibly a misidentifi the callipodoid family Schizopetalidae, as Ty cation ofthis common species. Gunthorp (1913, fWmma Loomis occurs around San Francisco/ 1921) questioned Cragin's record as being geo San Pablo and Monterey bays, and Idrionaria graphically improbable and deleted uncigerus Shelley inhabits Washington County, Utah (Shel from the Kansas fauna; he suggested that ley 1996). Secondly, 2 millipeds and 1 cen mutilated specimens of N. americanus from tipede are now known to occur between cen Topeka [cited as Arctobolus marginatus (Say)] tral Inyo County, California, and Washington may have been misinterpreted. Thirty-four years County, a range that corresponds roughly to later Causey (1955) proposed Californibolus the northern limit of the Mojave Desert eco fredericksoni for 2 males ostensibly collected system. The otber milliped is Piedolus utus in 1949 in Douglas County, Kansas, and an Chamberlin (Spirobolida: Atopetholidae), which immature female taken in 1950 from Monroe was previously known only from St. George, County, Iowa. Her illustrations of the antero Washington County (Hoffman and Orcutt 1960), ventral corner of the mandible and both pairs but can now be recorded from Inyo County of gonopods resemble the conditions in un based on 2 males from Panamint Springs (UCD). cigerus, and all other Kansas spirobolids that Similarly, the scolopendromorph centipede, she examined were N. americanus (cited as N. Theatops posticus (Say) (Cryptopidae), occurs oklahomae Chamberlin). Keeton (1960) exam in Saline Valley; the Nuclear Testing Site, Nye ined the holotype, transferred C. fredericksoni County, Nevada; and St. George, Utah (Shel into Tylobolus, and noted that the vial con ley 1990, 1997). Two additional millipeds that tained fragments of 2 specimens, the anterior may demonstrate this pattern are Califomiulus end of a small male and the caudal end of a 236 GREAT BASIN NATURAUST [Volume 57
larger individual. As the go napods appeared Keeton 1959, 1960, DeylUp and Franz 1994). to be from an immature specimen, Keeton There is no recent, credible evidence that withheld commentary on the status of T. fred another form exists in the eastern two-thirds ericksoni pending discovery of an adult male. of the continent, and the locality of T freder In our reexmnination ofthe holotype, we found icksoni mal' represent a laheling error. lOe dis what appeared to be 1 highly fragmented tribution pattern of species along the Pacific specimen with no legless segments at the cau Coast and in the western interior, and another dal end. Its gonopods are .ccurately figured by some 1000 mi (1600 km) to the east in the Causey and Keeton and are open to different eastern Central Plains, is not demonstrated by interpretations. Keeton (1960) thought they another diplopod genus and thus seems im were incompletely developed but gave no ex plaUSible for Tylobolus. We therefore designate planation for this opinion. They seem mature T fredericksoni as a "nomen dubium" and rec to u.s, and while similar to those of T. uncigertl$ ommend disregarding the species until its and plausihly representing a variant, they also presence in the Plains is confirmed by a show enough differences to be reasonably freshly collected male with unimpeachable interpreted as representing a distinct species. locality data, or until the rnilliped is discovered Today, 47 yr after its description, T. Feder in the principal gene.ic area, probably in Cali icksoni is still questionable, and no potential fornia, and an accurate locality can be reported. tylobolines have been collected from Kansas, Nebraska, or Iowa doring this time. Its osten ACKNOWLEDCMENTS sible occurrence in the Central Plains contin ues to generate confosion, as Enghoff (1995, We thank R.W Brooks and I.S. Ashe (SEM) family-clade no. 13) recorded Tylobolus from for loaning the holotype of C. fredemksoni, the east- and west-Nearetic based on this and L. Leibensperger (MCZ) for loaning the species. To our knowledge, the only definite types of T. ulahensis. The following curators, spirobolid records from Kansas are ofN. amer collection managers, and university faculty icanus (cited as N. oklahomae Chamberlin) from loaned specimens from the indicated reposito Douglas County (Causey 1955) and N. annu ries: BYV, R.W Baumann; CAS, C.E. Gris laris Rafinesque from Osage County (Keeton wold; CDFA, A.R. Hardy; DC, A.H. Barnum; 1960). We have seen 1 authentic Kansas spiro LACMNH, the late C.L. Hogue; NMNH, J.A. bolid, a juvenile from Lawrence, Douglas Coddington; UCD, the late R.O. Schuster; and County (VGA), tb.t is clearly referrable to UGA, G.L. Smith. Narceus. Aside from south Texas, which is occu pied by Anelus richardsoni (Pocock) (A1lopo LITERATURE CITED cockiidae) and 1 or more representatives of the Atopetholidae (Hoffman and Orcutt 1960, BOLLM!\J\, C. l8BS. Notes on North American Julidae. Shelley and Hoffman 1995), only 4 spiro Annals of the New York Academy of Science 4: S'.5-44. holoids definitely occur in the V nited States CAUSEY, N.B. 1955. Spirobolidae (Spirobolida: Oiplopoda) east of the Rocky Mountains: N. americanus, east of the Rocky Mountains. Journal of the Kansas which is widespread east of the Central Plains Entomological Society 28;69--80. and extends northward into Quebec (Shelley _ ...... 1957. Floridobolw, a new milliped g~nus (Spiro 19~~); N. gordanus (Chamberlin), in peninsular bolidae). Proceedings of the Biological Society of z Washington 70:205-208. Florida ; Chiwbolus spinigerus (Wood) (Spiro CHA;-,HlEHLIN, R.\~ 1925. Notes on some centipeds and bolidae), ranging from southern South Car millipeds from Utah. Pan-Pacific Entomologist 2: olina to the south Florida Keys (Keeton 1960); ~. and Floridobolus penneri Causey (Floridoboli . 1949. On some western millipeds of the order -"'Spirobolida. Journal of the Washington Academy of dae), endemic to the Lake Wales Ridge, High Sdence 39:163-169. lands ,md Polk counties, Florida (Causey 1957, CllA~mERLI~, R.\~, AND R.L. HOFFMAN. 1958. Checklist of the millipeds of North America. United States National Museum BuUetin 212;1-236. CRAGIN, F.W 1885. rlrst contribution to a knowledge of 2Kl....,t ENCHOFF, H. 1995. Historical biogeography of the Hol SHF.LLEY, R.M. 19SH. Thtl mil1ipOOs of eastern Canada arctic: area relationships, ance.~tral areas, and disper (Arthropoda; Diplopoda). G..1.nadian Journal of Zool sal ofnou-marine animals. Cladistics lL223--263. oj.,'Y 66;1638-1663. CUNTHORp, H. 1913. Annotated list of the Diplopoda and __,,' 1990. The centipcde, Theatops pusticus (Say), in Chilopoda, with a key to tile Myriapoda of Kansas. the southwestern United Slates and Mexico (Scolo Kansas University Science Bulletin 7:]61-182. pendromorpha; Cryplopidae), Canadian Journal of _-,,,,' 1921. Cmgin's collection of Kansas Myriapoda. Zoology 68;2631-2644. Canadiml Entomologist 53:87-9L __,,' 1995. The milliped family Hirudisomillidae in the HOFr"MAN, RL., AND B.S. ORCU'Ii. 1960. A syllopsis ofthe New World (PolY-.loniida). Brimleyana 23:103-.143. Atopctholidac, a family ofspiroboloid millipeds. Pro , 1996. The milliped order Callipodida in western ceedings ofthe United Stotes ~'ational Museum 111: --N"orth America (Schi;wpetalidac: Tynommatin