Ptyoiulus impressus ( Say 1821 )

Shelley, Rowland M. & Smith, Jamie M., 2016, Parajulid milliped studies XII: Initial assessment of Ptyoiulus Cook 1895 and neotype designations for Julus impressus Say 1821 and J. montanus Cope 1869 (Diplopoda: Julida), Insecta Mundi 2016 (522), pp. 1-21 : 8-11

Ptyoiulus impressus ( Say 1821)

Fig. 1–3 , 5 , 7–13 .

Julus impressus Say 1821: 102 . Brandt 1841: 101. Preudhomme de Borre 1884: 56. McNeill 1887: 324.

Bollman 1893: 144. Julus (Julus) pensylvanicus (Recte: pennsylvanicus ) Brandt 1841: 102–103. Iulus impressus: Gervais 1847: 176 ; 1859: 18. Wood 1865: 196. Provancher 1873: 418. McNeill 1888: 11. Iulus pensylvanicus: Gervais 1847: 177 ; 1859: 18. McNeill 1888: 12. Iulus pilosiscutis Wood 1864: 11 . Iulus pilosiscuta: Wood 1865: 198 . Iulus pennsylvanicus: Wood 1865: 201 , fig. 34. Julus pennsylvanicus: Preudhomme de Borre 1884: 59 . Julus pilosiscuta: Preudhomme de Borre 1884: 59 . Paraiulus impressus: Bollman 1887b: 36–37 , 44 (list). Brölemann 1922: 294–295, fig. 35–42. Blake

1931: 16–19 (list, caption, text). Paraiulus pennsylvanicus: Brölemann 1896: 45 , 68 (list, account); 1902: fig. 10–15; 1922: 291–292, fig.

22–27. Chamberlin 1940b: 58.

Paraiulus pilosiscutis: Brölemann 1896: 45 , 69 (list, account).

Parajulus impressus (not Williams and Hefner 1928): Bollman 1888a: 344; 1888b: 404; 1893: 119, 144 (text). Gunthorp 1913: 165. Bailey 1928: 20 (list). Brimley 1938: 498.

Parajulus pilosiscutus: Bollman 1887b: 44 (list).

Parajulus pennsylvanicus: Bollman 1887b: 44 ; 1888a: 344; 1888b: 404; 1888c: 106, 112; 1888d: 339; 1893: 119. Chamberlin 1918: 27. Bailey 1928: 20 (list). Williams and Hefner 1928: 126, fig. 19A, B. Brimley 1938: 498. Loomis 1944: 169.

Ptyoiulus pennsylvanicus: Chamberlin 1940a: 15 , pl. 8, fig. 71–73. Wray 1967: 153.

Ptyoiulus georgiensis Chamberlin 1943: 12 , fig. 26–27. Chamberlin and Hoffman 1958: 142. Hoffman 1999: 162. New Synonymy.

Ptyoiulus impressus: Chamberlin and Hoffman 1958: 141–142 . Filka and Shelley 1980: 13, fig. 6. Kevan 1983: 2965. Snider 1991: 181. Hoffman 1999: 162–163. Shelley 2000b: 183. Snyder 2008: 20 .

Ptyoiulus species incertis: Hoffman 1950: 31.

Aniulus impressus: Wray 1967: 153 .

Type specimens. Male neotype and numerous male, female, and juvenile paraneotypes ( ANSP) collected by H. C. Wood on an unknown date at an unspecified locality in Chester Co., Pennsylvania. One female paraneotype ( FSCA) collected by J. Oughton. 31 October 1934, in Philadelphia (Fairmont Park), Philadelphia Co., Pennsylvania.

Diagnosis. Calyx of anterior gonopod coxal process ( Fig. 5 , 7–11 ) slanting laterad, not coaxial with stem, lamellae flared and circumferentially entire, margins usually irregularly serrate to jagged, occasionally smooth or nearly so.

Color in life. Usually a mottled, subuniform dark brown, gray, or black. According to the vial label, the decapitated female from Wayne Co., Kentucky, exhibited a yellow circumferential band on most rings.

Variation. So many adult males of P. impressus reside in preserved repositories that no human could begin to measure a significant sample, particularly as part of an initial, broad, generic overview with time a factor. In the few individuals with fully-developed gonopods that we did measure, lengths varied from 28.9 to 57.9 mm, and the number of rings, including the collum and epiproct, ranged from 57 to 65. From the caudal end, heavily setose rings extend anteriad for at least 15 rings and even to around midbody, and the change to denser pilosity can be abrupt and noticeable or gradual, with denser and sparser pilosities blending together over a number of rings without a detectable change. Additionally, the denser pilosities may be random, subuniform, patchy, or in sublinear, circumferential rows ( Fig. 1–3 ; Blake 1931, fig. 2 in part). The epiproct is always apically acuminate and always overhangs and extends beyond the caudal paraproctal margins, but this too varies. The structure can be relatively short and narrow, as in fig. 1 of a male from Morgan Co., Ohio; long and spiniform as in fig. 2 of one from Allen Co., Kentucky; or short and stubby, as in fig. 3 of an individual from Leon Co., Florida, and Blake (1931, fig. 2 in part) of a New England male. Gonopodal variation is minimal, subtle, and primarily involves the calyx on the anterior gonopod coxal process ( Fig. 5 , 7–11 ), which always slants laterad. However, the angle of the slant varies slightly (compare those of the neotype, Fig. 7 , and the male from Greene Co., Arkansas, fig. 11) and some calyces also lean or tilt caudad such that in caudal view one can see slightly inside the structure ( Fig. 10 of a male from Leon Co., Florida). In some individuals the caudal flange of the calyx is shorter than the anterior one, so again one can view inside the calyx in caudal aspect ( Fig. 9 of a male from Liberty Co., Florida). The degree of serration and the jaggedness of the margins vary dramatically; from lightly and shallowly wavy or nearly smooth ( Fig. 8 of a male from Rabun Co., Georgia), to a host of continuous and irregular serrations, as in most individuals that we examined ( Fig. 10–11 of males from Leon Co., Florida, and Greene Co., Arkansas, respectively). Posterior gonopods ( Fig. 12–13 ) are subuniform. Relative lengths of the two branches vary slightly as do the lengths and degrees of curvatures of the main projections, the sizes of the subterminal flanges, and the lengths and degree of curvatures of the apices distal to the latter. All these variations are trivial and clearly intraspecific.

Ecology. Say (1821) characterized the habitats as “under stones and in humid situations.” An adult female that we examined from Wayne Co., Kentucky, was decapitated and dead on a trail. Habitat notations on vial labels include “under small pieces of 2x4s in oak litter,” “yellow pan trap,” “under stove in woods,” “fungus & under rock,” “upland forest,” “ground cover in woods,” “in woods near cave,” “pitfall trap in mixed mesic forest ravine,” ”pitfall in deciduous woods,” “upland (oak) woods.” “litter around cotton,” “in junk pile,” “in house,” “mammal burrow,” “moist leaf mold in rotting logs in mixed mesophytic forest,” “in leaf litter and moist very rotten logs on ground in mixed mesophytic woods,” “beech woods,” “under bark of rotting pine log,“ “west side of hill, open woods, under rock, leaf litter, & rotten wood,” and “around pines, hardwoods.”

Distribution ( Fig. 4 , 14 ). Southern Québec, northern Vermont, southern Michigan, and southeastern Wisconsin to the Gulf Coast of the Florida Panhandle, southern Alabama and Mississippi, east central/ northeastern Arkansas, and probably the northeastern periphery of Louisiana, presently documented only for the genus. With this possible exception, P. impressus inhabits every state documented for the genus and is anticipated in Rhode Island and Delaware, where Ptyoiulus is projected. Dimensions of the areas occupied by P. impressus are the same as those of the genus, and it alone inhabits detached areas 2 and 3. As only P. impressus has dispersed southward into areas inundated in the Cretaceous, we suspect that the unassigned records in southern Alabama and Mississippi ( Fig. 14 , blue dots) also are this species.

Published records. Overall range statements: United States east of the Rocky Mountains and Pennsylvania to Iowa and south to Florida ( McNeill 1888). Northeastern US west to Indiana and south to western North Carolina and Kentucky ( Chamberlin and Hoffman 1958, Filka and Shelley 1980). Widespread in the northeastern US ( Kevan 1983). CANADA: Ontario: Province in general ( Kevan 1983). Milton, Rattlesnake Pt. ( Causey 1952b, as Ptyoiulus sp. ). USA: USA in general ( Gervais 1847, 1859). Alabama: Alabama in general ( Hoffman 1999). Blount Co. ( Causey 1952a). Georgia: Georgia in general ( Bollman 1887b, 1893). Bibb Co., Macon ( Bollman 1888a). Butts Co., Indian Springs ( Bollman 1888a). Habersham Co., Tallulah ( Bollman 1888a). Rabun Co., NW of Clayton ( Chamberlin 1943, Chamberlin and Hoffman 1958, Hoffman 1999). Florida: Florida in general ( McNeill 1888, Bollman 1893). Escambia Co. ( McNeill 1887) . Illinois: Illinois in general ( Bollman 1887b, McNeill 1888). Indiana: Indiana in general ( Bollman 1887b; McNeill 1887, 1888). Crawford Co., Wyandotte ( Bollman 1888b). Fayette Co., Connersville ( Bollman 1888b, 1893). Franklin Co., Brookville ( McNeill 1888; Bollman 1888b, 1893). Monroe Co., Bloomington ( Bollman 1888b, 1893). Iowa: Iowa in general ( McNeill 1888). Kansas: Clay, Coffey, Cowley, Douglas, Franklin, Graham, Jefferson, Labette, Montgomery, Sumner, and Trego Cos. ( Gunthorp 1913). Massachusetts: North to the latitude of Boston ( Blake 1931); Norfolk Co., Milton, Wellesley ( Blake 1931); Plymouth Co., Hingham ( Blake 1931); Suffolk Co., Boston (Roxbury neighborhood) ( Blake 1931). Michigan: Michigan in general ( Bollman 1887b, Kevan 1983). Barry, Cass, Hillsdale, Ingham, Jackson, Kalamazoo, Kent, Lapeer, Lenawee, Livingston, Macomb, Van Buren, Washtenaw, and Wayne Cos. ( Snider 1991). Minnesota: Minnesota in general ( Bollman 1887b). Nebraska: Cuming Co., West Point ( Kenyon 1893). Lancaster Co., Lincoln, Roca ( Kenyon 1893). New Jersey: Morris Co., Rockaway ( Causey 1952a) . New York: New York in general ( Bailey 1928, Kevan 1983). Oswego Co., Oswego ( Loomis 1944). North Carolina: North Carolina in general ( Brölemann 1896, Chamberlin 1940a). Avery Co., ( Wray 1967, Filka and Shelley 1980); Grandfather Mt. and Linville ( Chamberlin 1940a, Wray 1967). Buncombe Co. ( Shelley 2000b). Burke Co. ( Shelley 2000b). Cleveland / Gaston Cos., Kings Mountain Region ( Filka and Shelley 1980). Durham Co., Duke Forest ( Brimley 1938, Wray 1967). Haywood Co. ( Shelley 2000b); GSMNP, Cataloochee, Purchase Knob ( Snyder 2008). Henderson Co. ( Shelley 2000b). Jackson Co. ( Shelley 2000b). Macon Co. ( Chamberlin and Hoffman 1958, Shelley 2000b). McDowell Co. ( Shelley 2000b). Polk Co, ( Shelley 2000b). Rutherford Co. ( Shelley 2000b). Surry Co. ( Shelley 2000b). Swain Co. ( Shelley 2000b); GSMNP, Ravensford ( Snyder 2008). Transylvania Co. ( Shelley 2000b). Wake Co., Raleigh ( Brimley 1938, Wray 1967). Watauga Co. ( Shelley 2000b). Yancey Co. ( Shelley 2000b). Ohio: Ohio in general ( Williams and Hefner 1928, Kevan 1983). Pennsylvania: Pennsylvania in general ( Brandt 1841; Gervais 1847, 1859; Wood 1865; Preudhomme de Borre 1884; McNeill 1888; Chamberlin 1940a; Hoffman 1999). Forest Co., Marienville ( Loomis 1944). McKean Co., Ludlow ( Loomis 1944). Philadelphia Co., Philadelphia vicinity ( Chamberlin and Hoffman 1958, Hoffman 1999). Susquehanna Co. ( Wood 1864, 1865; Chamberlin and Hoffman 1958; Hoffman 1999). Tennessee: Blount Co., GSMNP, Chilhowee ( Snyder 2008). Davidson Co., “beyond Glendale” ( Chamberlin 1918). Jefferson Co., Jefferson City (=Mossy Creek) ( Bollman 1888a, d). Knox Co., Knoxville ( Bollman 1888c). Sevier Co., GSMNP, Brushy Mtn., Goshen Prong, Twin Creeks ( Snyder 2008). Virginia: Page Co., Luray ( Bollman 1888a).

Deletions: Iowa, Kansas, Minnesota, Nebraska, and Durham and Wake Cos., North Carolina. Gonopod illustrations were not published for records from the first four states, and RMS has not found vouchers in any repository. All are also well outside the ranges we document ( Fig. 4 , 14 ) for the genus and species, so we consider them misidentifications of other parajulids, probably aniulinines. Likewise, Durham and Wake Cos., in central North Carolina where P. montanus is common, are also outside the range of P. impressus ( Fig. 14 ), and we do not believe the latter occurs in this part of the state.

Figures

Figure 4. Distributions of Ptyoiulus/Ptyoiulinae plotted against the maximal extent of the Western Interior Seaway during the Cretaceous Period, Mesozoic Era. All land areas shown are in the eastern land mass, Appalachia, which was separated from western Laramidia by the vertical, latitudinal arm of the Seaway (not shown), which segregated Ptyoiulus from bollmaniulinines in Laramidia but is irrelevant to Ptyoiulus itself. The Seaway spread eastward south of Appalachia and when it receded, P. impressus spread southward into the formerly inundated area. 1, large, contiguous main range area. 2, small, disjunct northernmost population extending from Montreal, Québec, to northern Vermont. 3, disjunct population along the southwestern coast of Lake Michigan. 4, disjunct area in northeastern/eastcentral Arkansas. 5, point locality in northeastern Louisiana vouchered by females. The transect denoted by the red lines near the center of the distribution is the band occupied by both P. montanus and P. impressus; the latter occurs alone to the north and south.

Figure 14. Distributions of species of Ptyoiulus. Black dots, P. impressus. Open black circle in southeastern Pennsylvania, denoted by the black arrow, is the neotype locality. Red dots, P. montanus. Open red circle in central North Carolina, denoted by the red arrow, is the neotype locality. Blue dots, geographically significant samples lacking adult males and presently unidentifiable to species.

Figures 1–3. Pilosity and epiproct variation on caudalmost rings of Ptyoiulus impressus. 1) Specimen from Morgan Co., Ohio. 2) Specimen from Allen Co., Kentucky. 3) Specimen from Leon Co., Florida. e, epiproct; h, hypoproct; lp, left paraproct.

Figures 5–6. Ptyoiulus spp. anterior gonopods, caudal views. 5) P. impressus. 6) P. montanus. at, anterior gonopod telopodite; ca, calyx; cp, coxal process stem.

Figures 7–13. Gonopodal variation in P. impressus. 7) Distal extremity of right anterior gonopod of neotype, caudal view. 8) The same of male from Rabun Co., Georgia. 9) The same of male from Liberty Co., Florida. 10) The same of male from Leon Co., Florida. 11) The same of male from Greene Co., Arkansas. 12) Right posterior gonopod of male from Rabun Co., Georgia, lateral view. 13) The same of male from Greene Co., Arkansas. Abbreviations as in Fig. 5–6.