Gosiulus conformatus Chamberlin, 1940

Shelley, Rowland M. & Smith, Jamie M., 2016, Parajulid milliped studies XI: Initial assessment of the tribe Gosiulini (Diplopoda: Julida), Insecta Mundi 2016 (509), pp. 1-17 : 4-8

Gosiulus conformatus Chamberlin, 1940

Fig. 2–12

Gosiulus conformatus Chamberlin 1940: 10 , pl. 4, fig. 32–35. Chamberlin and Mulaik 1941: 61. Causey

1952: 203. Chamberlin and Hoffman 1958: 138. Loomis 1959: 163; 1968: 160–162, fig. 6–7. Reddell

1965: 162; 1970: 399. Hoffman 1999: 154. Ziniulus aethes Chamberlin 1940: 13 , pl 6, fig. 48–50. Chamberlin and Hoffman 1958: 149. Loomis

1976: 290. New subjective synonymy. Ziniulus medicolens Chamberlin 1940: 13–14 , pl. 6, fig. 51–52. Chamberlin and Mulaik 1941: 61. Chamberlin and Hoffman 1958: 149. Loomis 1959: 163. New subjective synonymy. Ziniulus ambiguus Loomis 1959: 163 , fig. 20–23. New subjective synonymy. Ziniulus nati Loomis 1963: 122 , fig. 20–23. New subjective synonymy. Gosiulus aethes: Reddell 1965: 162 ; 1970: 399. Hoffman 1999: 154. Gosiulus ambiguus: Hoffman 1999: 154 . Gosiulus medicolens: Hoffman 1999: 154 . Gosiulus nati: Hoffman 1999: 155 .

Type specimens. Male Lectotype ( NMNH) collected by S. and D. Mulaik on an unknown date in December 1939 south of Three Rivers, Live Oak County (Co.), Texas. Paralectotypes ( NMNH), all collected on unknown dates in December 1939 by S. and D. Mulaik, as follows: M, 3F from south of Brady , McCulloch Co .; MM, FF, juvs. from Big Spring, Howard Co .; and M, 3F from 27.2 km (12.0 mi) N Alice, Jim Wells (not Brooks) Co.

Chamberlin (1940) did not specifically designate a holotype or paratypes in the original description but merely recorded G. conformatus from four Texas localities: south of Three Rivers, Live Oak Co.; south of Brady, McCulloch Co.; 27.2 km (17.0 mi) north of Alice, Jim Wells Co. , erroneously placed in Brooks Co.; and Big Spring, Howard Co. In the vials, however, he labeled the male from Live Oak Co., the only individual in the sample, as the holotype and the males, females, and juveniles in the Howard and McCulloch Co. samples. as paratypes, all meaningless without publication. Though not labeled, the specimens from Jim Wells Co. were mentioned simultaneously, so we consider them to be additional syntypes. These samples are from two regions of Texas, McCulloch and Howard Cos. being in the westcenter some 563.2 km (325.0 mi) NW of Live Oak Co. in the southeast. Hoffman (1999) erroneously reported the male from Live Oak Co. as the holotype, so for consistency, we designate it lectotype with the material from McCulloch, Howard, and Jim Wells Cos. becoming paralectotypes.

Diagnosis. Anterior gonopod with or without short, inconspicuous coxal lobes, lateral syncoxal process apically uncinate; posterior gonopod prefemoral process with variable acuminate spine arising from caudal surface around 1/3–2/3 length; process “C” moderately long, erect, extending directly ventrad for 1/3 to 1/2 of solenomere length, configuration variable.

Descriptive notes. Male length ranging from around 20.0– 32.4 mm, maximum width 1.3–3.4 mm; 47–57 rings including collum and epiproct. Ventral margin of mandibular stipes with broad, deep, semilunar indentation. Dorsum smooth and glossy with scattered metazonital setae especially on anterior rings, caudal rings glabrous. Paraproctal rims moderately thickened; hypoproct minute. Sterna not modified. 1 st legs moderately enlarged and forcipulate, tarsi overlapping in situ. Telopodites of both pairs of gonopods projecting through aperture in situ; anterior gonopod telopodites and lateral syncoxal processes angling caudad and overhanging 8 th sternum; posterior gonopods upright, telopodites extending directly ventrad between anterior gonopod structures. Anterior gonopods ( Fig. 2–5 ) with or without short coxal lobes; telopodites leaning mediad and extending ventrad for 2/3 of lengths of lateral syncoxal processes; latter variably uncinate apically. Posterior gonopod prefemoral process ( Fig. 6–12 ) with variable spine on caudal surface at 1/3–2/3 length, with or without small additional spines ( Fig. 8 ), usually distally uncinate/falcate but occasionally rounded and swollen; process “C” ( Fig. 6, 9–12 ) extending ventrad subparallel to solenomere for nearly half its length, configuration varying from filiform ( Fig. 6, 12 ) to boletoid ( Fig. 10 ) and subclavate, solenomere usually variably sigmoid ( Fig. 6, 10–12 ), occasionally curvilinear ( Fig. 9 ), extending substantially farther ventrad than other branches to become ventralmost telopodital projection, tapering smoothly and continuously to finely acuminate tip.

Gonopodal variation. Other than slight differences in their angles, the relative lengths of the telopodites and lateral syncoxal processes, and the degree of the apical uncination, the anterior gonopods are stable and constant. Given the consistent in situ arrangement of the two gonopod pairs, the anteriors apparently function as guides to position the posterior ones and/or spermatophores for mating. As guides, the actual structural configuration of the anterior gonopods seems insignificant as long as the posterior ones are properly aligned. This apparent function, as guides for inseminating the female cyphopods, has been postulated for the aniulinine Aniulus garius (Chamberlin) [= A. bollmani Causey ] ( Matthews and Bultman 1993, McAllister et al. 2009). We believe it applies broadly to parajulids and even “bigonopodal” helminthomorph diplopods, particularly representatives of the subterclass Colobognatha, where the posterior gonopods typically project anteriad between the anterior ones in situ.

The posterior gonopods, the true reproductive structures, are highly variable. While those of a gonopod pair are virtually indistinguishable, no one pair is structurally identical to another as all three projections vary. Those of the holotype of Z. aethes , from Austin, Travis Co. ( Fig. 9 ), are “scrunched together” in less physical space than in other males. Consequently, the branches are in contact, lie over and under each other, and force each other out of position; this is also the only male we examined with a curvilinear, rather than sigmoid, solenomere. We attribute this configuration to distortion caused by the tight, more closely appressed condition of the three projections.

The relative lengths of the branches vary in practically every posterior gonopod pair, but the solenomere is always substantially longer. Its width and degree of sigmoid curvature vary, but the most notable variation is the size of the basal swelling near the origin of process “C”; the male from LaSalle Co. ( Fig. 11 ) lacks this swelling. The solenomere is the least variable projection, but process “C” is highly so. Its length, width, and configuration vary, and it may be apically acuminate or boletoid and sublinear or curve gently anteriad distally.

The prefemoral process consistently displays the diagnostic caudal spine, but its length and position on the process vary ( Fig. 6, 9, 10–12 , sp) as does practically every other aspect of the projection including overall length. One to three additional short spines may arise distad but proximal to the distal curve/ bend of the branch. The distal curve/hook is directed anteriad, opposite to the spined margin, and may be broad or narrow with variable apices; it is an abrupt, sharp bend in the male from Lubbock Co. ( Fig. 12 ).

Chamberlin authored all five available names, two in Ziniulus ( aethes and nati ) and three in Gosiulus ( conformatus , ambiguus , and medicolens ). The posterior gonopods of the type of each differ, so he apparently concluded that each represented a separate species. Assessing these five variants in the context of all lowland Texas gosiulinines reveals them to be variants of one highly variable species for which conformatus is the oldest name. By Chamberlin’s standards, we would have as many nominal species as samples with males, but clearly this is not the case. The only name that plausibly could represent a true species is aethes because of its closely appressed posterior gonopod branches and the curvilinear solenomere, but we interpret these as anomalies exhibited by this individual. More sampling in Travis Co. will reveal whether sufficient individuals show this condition to warrant taxonomic recognition, so we assign it to conformatus . An example of a localized milliped species that is surrounded by a widespread congener is Xystocheir prolixorama Shelley ( Polydesmida : Xystodesmidae ), occurring inside the range of X. d. dissecta (Wood) in northern California ( Shelley 1996).

Ecology. Gosiulus conformatus inhabits a variety of low elevation, flatland biotopes and seemingly may be expected virtually any place within its known and projected areas ( Fig. 13 ). Published habitat notations include “bottom of 90’ entrance drop, apparently washed into the cave” and “bottom of sink entrance” (both by Reddell 1965) and “leaf litter on a ledge 20 feet below the entrance” ( Reddell 1970). Comments on labels with newly examined material include “on the ground under rocks,” “cotton field,” “berlese of leaf litter,” “sandy soil, logs, pipes,” “under bags put on concrete floor” (inside an office building), and “slight preference for clay soil.”

Distribution ( Fig. 13 ). Known only from lowland regions of Texas, extending from the western periphery of the Piney Woods biome in eastcentral Texas, approximately 280.0 km (175.0 mi) west of the Louisiana border, west- and northward to the eastern Trans Pecos and High Plains biomes, respectively, and southward to approximately 160.0 km (100.0 mi) north of the Rio Grande in Jim Wells Co. Gosiulus conformatus has not been taken south of the Rio Grande in Mexico (Tamaulipas, Nuevo León, or Coahuila states), but it occurs approximately 22.4 km (14.0 mi) from the River in Webb Co. ( Loomis 1963) and up to the watercourse itself in Maverick and Val Verde Cos.; it should be expected directly across the Rio Grande in Coahuila ( Fig. 13 , short black arrows) and perhaps even farther south. As the lowlands of westcentral Texas and the Panhandle spread west- and northward into eastern New Mexico, southeastern Colorado, and the Oklahoma Panhandle ( Fig. 13 , long black arrows); it also plausibly occurs north of the Red River in southern Oklahoma ( Fig. 13 ) and conceivably even western Kansas. We doubt that the milliped inhabits southernmost Texas and the well-sampled Rio Grande Valley because it seemingly would have been found by now.

Published records. Texas: Bexar Co. ( Loomis 1968, Hoffman 1999); and Bullis Hole ( Reddell 1970). Bexar/Comal/ Guadalupe Cos., Schertz, J. O. Vaughan Ranch ( Loomis 1959). Burnet Co. 6.0 km (5.0 mi) SE Marble Falls ( Loomis 1976, Hoffman 1999). Edwards Co., 12.8 km (8.0 mi) NE Rock Springs, Devil’s Sinkhole ( Reddell 1965). Goliad Co., Berclair ( Causey 1952). Howard Co., Big Spring ( Chamberlin and Mulaik 1941, Loomis 1968). Jim Wells Co. ( Hoffman 1999) ; 80.0 km (30.0 mi) N Alice ( Chamberlin 1940, Chamberlin and Mulaik 1941). Kendall Co., 8.0 and 17.6 km (5.0 and 11.0 mi) SSW Boerne ( Chamberlin and Mulaik 1941, Loomis 1959, Hoffman 1999). Kerr Co., Raven Ranch, 19.2 km (12.0 mi) S Kerrville ( Chamberlin 1940, Chamberlin and Mulaik 1941, Chamberlin and Hoffman 1958, Hoffman 1999). LaSalle Co., 2.7 km (1.7 mi) N Artesia Wells ( Loomis 1963, Hoffman 1999). Live Oak Co. ( Hoffman 1999); and S of Three Rivers ( Chamberlin 1940, Chamberlin and Mulaik 1941, Chamberlin and Hoffman 1958, Loomis 1968, Hoffman 1999). McCulloch Co. ( Hoffman 1999); and S of Brady ( Chamberlin 1940, Chamberlin and Mulaik 1941). Medina Co., Valdina Farms Sinkhole 30.0 km (18.8 mi) NW Rondo ( Reddell 1965). Travis Co., Austin ( Chamberlin 1940, Chamberlin and Hoffman 1958, Hoffman 1999). Webb Co., 22.4 km (14.0 mi) N Laredo ( Loomis 1963). Williamson Co., Cobb Cavern, 8.0 km (5.0 mi) N Sun City; Georgetown ( Reddell 1965).

Material examined. Texas: Bandera Co., Bandera, Shaw Ranch, M, F , 2 February 1962, G. Marion ( FSCA). Bee Co., Beeville, M, 10 October 1895, E. Swain ( NMNH). Bell Co., 19.2 km (12.0 mi) N Temple, 2M, 3 November 1927, O. F. Cook ( FSCA); and SW Belton, 3M, January 1931, O. F. Cook ( FSCA). Bexar Co., San Antonio, 7M, 26 November 1912 ( FSCA), 2M, 9 December 1930, O. F. Cook ( FSCA), and M, 27 December 1965, T. Stewart ( FSCA); and 28.8 km (18 mi) N San Antonio, 2 January 1943, W. S. Ross ( FSCA). Brown Co., Lake Brownwood St. Pk., MM, FF, juvs., 27 November 2002, C. T. McAllister ( FSCA, UTIC). Burnet Co., Marble Falls, 2M, F, 15 February 1975, J. C. Loomis ( FSCA); rest area W side Marble Falls, M, 13 March 1982, J. C. Cokendolpher ( FSCA); Eckhardt Root Cave, 11.0 km (6.9 mi) S Bertram, M, F, 17 April 1990, M. Reyes ( UTIC); and Simon Says Sink, 10.0 km (6.3 mi) S Bertram, M, 12 November 1990, J. R. Reddell, M. Reyes ( UTIC). Caldwell Co., Maxwell, M, 1 March 1963, R. O. Albert ( FSCA). Comal Co., New Braunfels, FF, 18 December 1954, L. Hubricht ( FSCA); Hendricks Cave nr. New Braunfels, F, 19 March 1960, W. J. Gertsch ( FSCA); 6.4 km (4.0 mi) E Bulverde, Kappelman Cave, M, F, 15 March 1964, J. R. Reddell, W. Russell ( FSCA); 32.0 km (20.0 mi) W San Marcos, Espinajo Diablo, 2M, 8-14 March 1995, W. E. Steiner, J. M. Swearingen ( NMNH); and Barely Cave, Camp Bullis, M, 12 January 2000, P. Sprouse ( UTIC). Dallas Co., Dallas, MM, FF, May-June 1955 ( FSCA). Duval Co., 6.4 km (4.0 mi) NW San Diego, FF, 8 January 1961, R. O. Albert ( FSCA); and NE corner, 0.8 km (0.5 mi) W Jim Wells Co. line, along Parrits Cr., MM, FF, 4 April 1962, R. O. Albert ( FSCA). Goliad Co., Goliad, M, 2 January 1952, ( FSCA). Grimes Co., Navasota, M, 2 November 1961, S. Hopkins ( FSCA). Hays Co., 27.2 km (17.0 mi) W San Marcos, F, 2 March 1995 and 6.4 km (4.0 mi) NW Kimberley, Twin Sister Peaks, 3M, 3F, 9 March 1995, W. E. Steiner, J. M. Swearingen, J. R. Ott, E. Silverfine ( NMNH). Howard Co., Big Spring St. Pk., MM, FF, 24 December 2003, C. T. McAllister ( FSCA). Hunt Co., Greenville, F, 8 February 1932 ( FSCA). Irion Co., 9.6 km (6.0 mi) SSE Mertjon, M, F, 18 November 2005, C. T. McAllister ( FSCA). Jim Wells Co. , 1.6 km (1.0 mi) N Brooks Co. line, 2M, 24 April 1961, J. F. Quinlan ( FSCA); Alice, MM, FF, November 1961 and M, FF, 22 January 1967, R. O. Albert ( FSCA); and 27.1 km (17.0 mi) N Alice, M, 3F, S. and D. Mulaik ( NMNH). Johnson Co. , along county road 1434, M, 27 March 2010, C. T. McAllister ( FSCA). Karnes Co. , 1.6 km (1.0 mi) N Falls City, 2M, F, 3 April 1961, J. F. Quinlan ( FSCA); and Falls City, 2M, F, 17-19 April 1961, J. F. Quinlan ( FSCA). Lavaca Co., Hallettsville, M, 19 January 1962, M. E. Key ( FSCA). Live Oak Co., George West, M, F, 4 December 1960, R. O. Albert ( FSCA); and 24.0 km (15.0 mi) N Sandia, M, juvs., 18 February 1961, R. O. Albert ( FSCA). Lubbock Co., Yellow House Canyon, MM, FF, juvs., 30 March and 26 April 1973, T. R. Mollhagen ( FSCA). Mason Co., 8.0 km (5.0 mi) SW Mason, M, F, 8 November 1964, J. R. Reddell ( FSCA). Maverick Co., Eagle Pass, MM, FF, 19 October 1932, O. F. Cook ( FSCA). Nolan Co., Sweetwater, M, F, 2 November 1932, H. C. McNamara ( FSCA). Randall Co., Palo Duro Canyon St. Pk., 24.0 km (15.0 mi) E Canyon, MM, FF, juvs., 28 April 1962, R. O. Albert ( FSCA); and Canyon, M, 1 November 1993, W. D. Sissom ( NCSM). Reagan Co., 13.6 km (8.5 mi) NW Barnhart, along US hwy. 67, M, F, 11 November 2005. C. T. McAllister ( FSCA). Real Co., Prade Ranch on Frio R., F, 21 April 1962, R. O. Albert ( FSCA). Runnels Co., Miles, along US hwy. 67, M, 2F, juv., 23 December 2006, C. T. McAllister ( FSCA). San Patricio Co., Sinton, 2M, F, 11 December 1960, F, juv. ( FSCA). Shackleford Co., between Albany and Lueders, M, F, 8 November 1927, O. F. Cook ( FSCA). Stonewall Co., between Swenson and Peacock, 8 November 1927, O. F. Cook ( FSCA). Sutton Co., Caverns of Sonora headquarters, 7M, 8F, 3 juvs., 18 November 2005, C. T. McAllister ( FSCA). Taylor Co., 16.0, 24.0, and 40 km (10.0, 15.0, and 25.0 mi) SW Abilene, MM, FF, juvs., 1 March 1944, H. S. Dybas ( FSCA). Terrell Co., 16.0 km (10.0 mi) SE Sanderson, 2M, F, 30 October 1943, W. S. Ross ( FSCA). Tom Green Co., 56.0 km (35.0 mi) NW San Angelo, March Ranch, M, 22 November 1970, D. L. Rambo ( FSCA). Travis Co., S of Austin on TX hwy. 135, along Onion Cr., M, 23 January 1976, J. Richter ( UTIC); and 2.8 km (1.75 mi) S Longhorn Dam, M, 2 March 1973, J. T. Moore ( UTIC). Uvalde Co., Uvalde, M, F, 10 October 1927, O. F. Cook ( FSCA); Laguna, 5M, 2F, 25 December ( AMNH); 4.2 km (2.6 mi) W Uvalde, M, F, 12 April 1940, S. and D. Mulaik ( NMNH); 32.0 km (20.0 mi) NW Rondo, Valdina Farms Sinkhole, M, F, 12 January 1964, J. R. Reddell ( FSCA); and 35.2 km (22.0 mi) NW Uvalde, Mason Ranch, M, F, 1 February 1967, Miles, R. Tandy, R. Ballinger ( FSCA). Val Verde Co., Del Rio, 2M, 19 October 1932, O. F. Cook ( FSCA); 8.0 km (5.0 mi) E Shumba, near Comstock, M, 8F, 4 juvs., S. and D. Mulaik ( NMNH); and 16.0 km (10.0 mi) SE Del Rio, M, F, 22 March 1978, O. F. Franke, T. B. Hall, J. V. Moody ( UTIC). Williamson Co., TWAS A Cave, Cedar Park, 2M, 2F, 16 April 1989, W. Elliott, J. R. Reddell, M. Reyes ( UTIC); Garden of Sinks Cave, Cedar Park, 2M, F, 13 February 1990, J. R. Reddell, M. Reyes ( UTIC) and M, 16 February 1990, J. R. Reddell ( UTIC); Chaos Cave, 3.0 km (1.9 mi) N McNeil, 2M, F, 14 April 2000. J. R. Reddell, M. Reyes ( UTIC); and Ranch at Deer Cr., Cedar Park, Jumbled Rocks Cave,.M, 2F, 10 April 2001, M. Warton ( UTIC). Wilson Co., 19.2 km (12.0 mi) W Falls City, M, F, 2 May 1961, J. F. Quinlan ( FSCA); and 3.2 km (2.0 mi) NW Falls City, juv., 25 February 1961, J. F. Quinlan ( FSCA).

Remarks. The holotype of the synonym, Ziniulus medicolens , is an unusually large-bodied gosiulinine.

Comparing Fig. 1 and 13 reveals that the entire distribution of G. conformatus was submerged at the height of the Western Interior Seaway in the Cretaceous/Paleocene periods (65–100 mya) while the western part of the range of G. timpius (El Paso, western New Mexico, and Arizona) was land in eastern Laramidia as was at least most of the distribution of Nesoressini . Consequently, G. conformatus appears substantially younger than its more anatomically stable congener, and its greater variability may reflect insufficient time to stabilize.

Figures

Figure 1. Known (solid lines) and projected (dashed lines) distribution of Gosiulini plotted against the maximal extent of the Western Inland Seaway in the Cretaceous Period, ca. 65–100 mya. The approximate distribution of Nesoressini (Shelley and Medrano 2006) is shaded in red.

Figure 13. Distributions of species of Gosiulini. Black, G. conformatus. Red, G. timpius. Green, M. biramus.

Figures 2–5. Gosiulus conformatus, left anterior gonopods. 2) Holotype, anterior view. 3) The same, lateral view. 4) Male from Travis Co., anterior view. 5) The same, lateral view. at, anterior gonopod telopodite; lsp, lateral syncoxal process.

Figures 6–12. Gosiulus conformatus, left posterior gonopods (sub)medial views, all from Texas. 6) Lectotype, Live Oak Co. 7) Distal halves of prefemoral process and solenomere of paralectotype from McCulloch Co., lateral view. 8) Distal half of prefemoral process of male paralectotype from Jim Wells Co., lateral view. 9) Telopodital projections of male from Travis Co. 10) The same of male from Kerr Co. 11) The same of male from LaSalle Co. 12) The same of male from Lubbock Co. C, process “C”; pfp, prefemoral process; s, solenomere; sp, subbasal prefemoral spine.