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The southern Appalachian millipeds Boraria stricta (Brölemann, 1896) and B. infesta (Chamberlin, 1918) (Diplopoda: Polydesmida: Xystodesmidae) have become established in Westchester Co., New York, and Hartford Co., Connecticut, respectively. Only three individuals are available for the latter, but B. stricta has established a reproducing population in southern New York state. This species is also recorded from Bland Co., Virginia, in the Ridge and Valley Physiographic Province. Boraria profuga (Causey, 1955) comprises two allopatric populations, one in Montgomery Co., Arkansas, and the other in Ouachita Parish, Louisiana. Distributional records and gonopod drawings are presented for these species plus B. deturkiana (Causey, 1942).
Ptyoiulus Cook 1895, the dominant parajulid diplopod genus in the eastern United States (US), comprises two species – P. impressus (Say 1821), with a slanted, fl ared, circumferentially entire, and marginally serrate apical calyx on the anterior gonopod coxal process, and P. montanus (Cope 1869), n. comb., with a smooth, upright, cupulate calyx that is open caudad and coaxial with the process’ stem. The genus occupies a broad area between the Mississippi River and Atlantic Ocean extending from southern New England, Ontario, and Michigan to the Florida Panhandle and four small disjunct ones – from Montreal, Québec, to northern Vermont, along southwestern Lake Michigan in Wisconsin and Illinois; northeastern/eastcentral Arkansas, primarily in Crowley’s Ridge physiographic feature and beside the “bootheel” of Missouri; and a point locality in northeastern Louisiana just south of the Arkansas line. A male from Chester County (Co.), Pennsylvania, is designated as the neotype of Julus impressus, as is one from Durham Co., North Carolina, for J. montanus. As both species inhabit Montgomery Co., Virginia, the type locality of J. montanus, we exercise the right of first reviser, conserve the latter name, and assign it to the species with the smooth, cupulate, and coaxial calyx. We also exercise first reviser rights and assign Parajulus ectenes Bollman 1887 to this form, thereby relegating it to synonymy under Ptyoiulus montanus. Other new synonymies include Ptyoiulus georgiensis Chamberlin 1943 under P. impressus and P. coveanus Chamberlin 1943 under P. montanus. Both Ptyoiulus and P. impressus are projected for Delaware and Rhode Island and newly reported from Québec, Connecticut, District of Columbia, Maryland, Mississippi, South Carolina, Vermont, West Virginia, and Wisconsin, and the genus and species, respectively, are newly documented from Louisiana and Arkansas; P. montanus is newly cited from Alabama, Arkansas, Georgia, Mississippi, and South Carolina. Ptyoiulus impressus occupies every state except perhaps Louisiana and is the only species in areas that were inundated during the Cretaceous and glaciated during the Pleistocene; by contrast, P. montanus inhabits a relatively narrow east/west transect through the center of the generic range. Their distribution patterns suggest an old species, montanus, being actively displaced by the younger and more successful impressus. The decurvature of the epiproct in uroblaniulinines appears to increase with age and developmental stage. A key is presented to parajulid familygroup taxa in the US and Canada east of the Rocky Mountains.
Parajulid milliped studies XI : Initial assessment of the tribe Gosiulini (Diplopoda: Julida)
(2016)
The parajulid milliped tribe Gosiulini (Diplopoda: Julida) comprises two genera – Gosiulus Chamberlin, with three projections on the posterior gonopod and two species in the southcentral/southwestern United States (US) [Arizona, Colorado, New Mexico, and Texas], and monotypic Minutissimiulus Shelley, n. gen., with two projections, in Nuevo León, Mexico. Gosiulus conformatus Chamberlin occupies the plains/fl atlands of Texas, while its congener inhabits high elevations to the west in all four US states. Both are anticipated in Mexico (Coahuila, Chihuahua, and Sonora), and G. conformatus is expected in southeastern Colorado, eastern New Mexico, and the Oklahoma panhandle. The eastern boundary of G. conformatus and the genus/tribe conforms to the western border of the Piney Woods biome in eastern Texas. As shown by the posterior gonopod drawing in the original description, Parajulus timpius Chamberlin, previously considered of “uncertain generic position or validity,” is unquestionably the oldest name for the western species. The anteriormost posterior gonopod projection, absent from Minutissimiulus, is considered the “prefemoral process,” while the “solenomere” and a third branch arise from a common base.
Because of positional homology with “process ‘C’” in Nesoressini, the last projection is accorded this name, which may also apply to the “prefemoral process” in Aniulini. Minutissimiulus biramus Shelley, n. sp., is proposed along with the following new subjective synonymies: Apacheiulus Loomis under Gosiulus; Ziniulus aethes and Z. medicolens, both by Chamberlin, and Z. ambiguus and Z. nati, both by Loomis, under G. conformatus; and A. pinalensis and A. guadelupensis, both by Loomis, under G. timpius, new combination. Ziniulus navajo Chamberlin becomes an objective synonym of P. timpius because its holotype is designated neotype of the latter. Minutissimiulus biramus Shelley is the fi rst Mexican gosiuline and “mainland” Mexican parajulid not in the tribe Parajulini.
Pandirodesmus rutherfordi, n. sp., represented by 18 individuals including eight adult males, occurs in secondary forests near Charlotteville and Speyside, Tobago, Trinidad and Tobago. Along with the type and second species, P. disparipes Silvestri, from Guyana and known only from females, the segmental legs of P. rutherfordi alternate between long (anterior pairs) and short (posterior ones), spiracular openings are on straw-like tubules, and ozopores are located on paramedian metatergal spines. These features appear to be adaptations for biotopes of loose sand, detritus, or frass, and 17 specimens, including the six juveniles, exhibit coatings of “sand grains” that are loosely cemented together and to the smooth, translucent, grayish-white exoskeleton. The tubules and spines elevate the spiracles and ozopores above the coating, thereby ensuring that they remain open and functional.
The coating, which provides camoufl age and lends strength and rigidity to the poorly sclerotized exoskeleton, is a subuniform “pavement” that covers the entire animal except the labrum/clypeus, tarsal and antennal apices, prozonae, paraprocts, and the gonopods in males. Ramose/dendritic setae, particularly on narrowly rounded podo-/antennomeres, trap “sand grains,” and the ozopore secretions apparently constitute the “glue” that cements the coating, as evidenced circumstantially by layers of “sand” between the spines on the anterior metaterga, where they are physically closest. The alternating segmental leg lengths, in part due to differing ventrolateral and ventromedial origins, appear to be an adaptation for lateral/sideways motion in which the long (anterior) legs extend laterally and pull the body to the level of the short (posterior) ones, which continue the motion while the anterior legs extend to begin the next stroke. The opposing legs perform the complementary pushing motion a fraction after the long legs initiate the pulling stroke and hence are slightly and purposefully out of sync. An adult male paratype lacks the coating, probably because it had just molted and lacked time to amass it; the juvenile female paratype of P. disparipes also is “naked,” as was, according to Silvestri, the now lost adult female holotype. Until fresh material is collected, coatings cannot be confi rmed for P. disparipes even though it shares the anatomical modifi cations that seem adaptions for such. The minute, triramous gonotelopodites of P. rutherfordi are unlike any known for a chelodesmid, so the current generic placement, in a monotypic tribe in the nominate chelodesmid subfamily, is retained. With species in both South America and the southern Antilles, Pandirodesmus/ini had to exist on both the “proto-Antillean” terrane and the adjoining part of Pangaean Gondwana before the former rifted in the Cretaceous/Paleocene, ~66 million years ago, and P. rutherfordi is a remnant of the former population that became isolated on present-day Tobago when the terrane fragmented. Affi nity between Guyanan and southern Antillean platyrhacid millipeds (Polydesmida: Leptodesmidea) suggest that Pandirodesmus/ini may occur sporadically as far north in the island chain as St. Lucia.
A trimaculate male of the diplopod genus Apheloria Chamberlin (Polydesmida: Xystodesmidae/-inae: Apheloriini) from 1.3 km (0.8 mi) west of McKenney, Dinwiddie County (Co.), Virginia, is designated the Neotype of Julus virginiensis Drury 1770, thereby stabilizing the earliest name for a North American milliped and authenticating its prior assignment to this taxon. The existing concept of Apheloria is accepted in the absence of a revisionary treatment, and a modern description of A. v. virginiensis with gonopod drawings and color photos is provided. Drury’s original account and his letter to the Virginian who sent him the original specimens are quoted verbatim to eliminate future library searches. The specific name has been associated with at least three genera, and its confusing history is clarified by summarizing works in each. Authentic localities, mapped to the extent now possible, reveal a distribution south of the James River in piedmont and coastal Virginia that extends southwestward to the Blue Ridge foothills and at least as far south in North Carolina (NC) as Greensboro, the “Triangle” (Raleigh/Durham/Chapel Hill region), and Albemarle Sound in the east. Based on the holotypes, A. aspila and A. tigana, both by Chamberlin, are placed in synonymy under A. v. virginiensis (syns. nov.), and although its status is still under review, A. waccamana Chamberlin, whose type locality is Lake Waccamaw, Columbus Co., in southeastern NC, may be the correct name for today’s A. tigana. All samples so labeled must be reexamined for misidentifications of A. v. virginiensis.
A newly discovered population of Xystocheir brachymacris Shelley, 1996 (Polydesmida: Xystodesmidae: Xystocheirini), in Placer County (Co.), California, exhibits an unusual grayish-black color dorsally with mottled, ovoid patches at paranotal bases; it cons titutes northern generic and specifi c range extensions of ~28.4 km (17.6 mi). The gonopods differ from those in the El Dorado Co. population in having shorter/acuminate prefemoral processes and blade-like, rather than spatulate, processes “B” that angle away from the solenomere instead of overhanging it. Additionally, a strong distomedial prefemoral lobe, absent from the El Dorado population, arises from the stem in Placer Co. males. Authorship of Xystocheirini is properly attributed to Hoffman, 1980.
Euryuridae Pocock 1909 and Eurymerodesmidae Causey 1951, both endemic to the eastern/central United States (US), are incorporated into Xystodesmidae Cook 1895 and reduced to subfamilies and (sub)tribes n. stats. Euryurina and Melaphina Brolemann 1916, n. stats., are sister-taxa that differ primarily in epiproctal configurations and comprise Euryurini; sister-taxa Eurymerodesmina and Nannariina Hoffman 1964, n. stats., the latter transferred from Xystodesminae, comprise Eurymerodesmini, n. stat., in which plesiomorphic forms exhibit sublinear, "stick-like," and subapically curved/bent gonopodal acropodites with moderately-long to long hairs, often with distal tufts, on their “inner” surfaces. Additional transferrals include Wamokia Chamberlin from the xystodesmine tribe Xystocheirini to Xystodesmini (= Harpaphini), and Macellolophus Attems, from Xystodesmidae to Chelodesmidae. Except for Chonaphini, the term, "prefemoral process," has traditionally been assigned to the secondary and shorter telopodital projection regardless of its position, origin, or configuration. Homology of these different structures has never been demonstrated and requires investigation, but the multitude of differences suggests that they are not such and warrant different names, for example "femoral process," for the similarly positioned branches in Devillea Brölemann and Rhysodesmus Cook (Xystodesminae: Devilleini, Rhysodesmini). The latter tribe may be polyphyletic, and new tribes may be required for components with acicular "prefemoral processes" (Boraria and Cherokia, both by Chamberlin, Gyalostethus and Erdelyia, both by Hoffman, and Pleuroloma Rafinesque) and the southeastern US genera with small-bodied species (Caralinda Hoffman and Gonoessa, Parvulodesmus, and Lourdesia, all by Shelley). Taxonomic value is accorded the "prefemoral extension/elongation," which is absent from Eurymerodesmina; complete, encircles the acropodite, and extends for ~1/3 to 1/2 of the latter’s lengths in Euryurini; and incomplete and extends for ~1/4 to 1/3 of the "outer" acropodital surfaces in Nannariina and xystodesmine tribes. Other newly recognized taxonomic characters include the "inner" and "outer" acropodital surfaces/margins, the position on the acropodital stem of the "distal curve/bend," and the length of the "distal zone." Rhysodesmus and Sigmoria (Rudiloria) t. trimaculata (Wood) (Xystodesminae: Rhysodesmini, Apheloriini) are recorded from, respectively, Chihuahua, Mexico, and Québec, Canada, as are Xystodesmidae/-inae and, provisionally, Chonaphini, Montaphe Chamberlin, and M. elrodi (Chamberlin), the only plausible taxa for an unidentifiable juvenile from near Yahk and only 2.5 km (1.6 mi) north of the International Border. The southern periphery of interior British Columbia (BC) thus represents the second xystodesmid faunal region in BC and the third in Canada. While incorporation of Euryuridae does not affect the family’s overall distribution, that of Eurymerodesmidae fundamentally alters it by joining the formerly separate East-Nearctic and Meso-American regions into a continuous one extending, north-south, from Montréal Island, Québec, to Santa Ana Department, El Salvador, a distance of around 4,944 km (3,090 mi). Xystodesmidae also inhabit two West-Nearctic regions, one in the interior stretching from southernmost BC to northeastern Oregon and the other running along the Pacific Coast from southern Alaska to southern California. The family also occupies two Palearctic regions, each with three subregions, an eastern one spreading from Hokkaido, Japan, and the southern Maritime Province, Russia, to Taiwan; a point locality in northern Vietnam; and southern/eastern China. The second Palearctic area extends along the Mediterranean and adjoining seas from Morocco, Sardinia, and the southeastern corner of France to Cyprus and southern coastal Turkey. New locality data, references, and maps are provided along with diagnostic accounts of all reconceptualized taxa and new/revived statuses.
A simple, sublinear, "stick-like" acropodite with a curve or bend near midlength or subapically and without a secondary telopodital projection is the hypothesized plesiomorphic gonopodal condition in Xystodesmidae. This form has undergone multitudinous modifications/alterations - twists, curls, variably configured thickened and laminate expansions, reductions, bi-/trifurcations, enlargements, ornamentations, etc. that are manifested in today’s xystodesmine tribes. When Avalonia collided with Baltica 450 million years ago, ancestral xystodesmoideans on the former dispersed into the latter, penetrated and occupied vacant niches, and evolved into today’s Melaphina (Euryurinae: Euryurini) and Devilleina (Xystodesminae). A similar evolutionary burst leading to today’s Nearctic and East-Palearctic faunas occurred 10 million years later when Avalonia + Baltica collided with Laurentia to form Euramerica. Ancestral forms of Euryurinae and Xystodesminae again penetrated vacant niches and evolved; the former maintained the general gonopodal structural pattern of Melaphina but changed the epiproct from triangular to broad and spatulate, thereby creating Euryurina. The earliest xystodesmine taxa to evolve in Laurentia were Rhysodesmini and Rhysodesmus, which spread southwestward, penetrated "proto- Mexico, and left relict populations in today’s southern Appalachians. Eurymerodesmina and Nannariina arose from ancestral euryurine stock prior to the Cretaceous in western Appalachia in their present area of overlap. The former dispersed to the west and south while the latter expanded to the east and north; consequently, the Western Inland Seaway minimally impacted Nannariina while eradicating Eurymerodesmina from the inundated area. Today’s populations in the Plains and south-southeastern states therefore represent secondary dispersion in the past 50-60 million years. The Seaway also eradicated Rhysodesmus from these areas, but enough forms survived in high mountain refugia to replenish the fauna when the embayment receded.
The biogeographic significance of Diplopoda is substantiated by 50 maps documenting indigenous occurrences of the 16 orders, the three Spirostreptida s. l. suborders – Cambalidea, Epinannolenidea, Spirostreptidea – and all higher taxa including Diplopoda itself. The class is indigenous to all continents except Antarctica and islands/archipelagos in all temperate and tropical seas and oceans except the Arctic; it ranges from Kodiak Island and the northern Alaskan Panhandle, United States (USA), southern Hudson Bay, Canada, and near or north of the Arctic Circle in Iceland, continental Scandinavia, and Siberia to southern “mainland” Argentina, the southern tips of Africa and Tasmania, and Campbell Island, subantarctic New Zealand. The vast, global distribution is interrupted by sizeable, poorly- or unsampled areas including the Great Basin, USA; the Atacama Desert region of Chile and neighboring countries; southern South American islands; the central Kalahari and Sahara deserts; the Gobi Desert, Mongolia, and all of north-central and western China; from north of the Caspian Sea, Russia, to central Kazakhstan; and the “Outback” of central Australia. Five Arabian countries lack both samples and published records of indigenous diplopods – Bahrain, Kuwait, Oman, Qatar, and United Arab Emirates – as do Turks and Caicos, in the New World, and Mauritania and possibly Egypt, Africa. New records, including the first for Chilognatha from Botswana and the first specific localities from Northern Territory, Australia, are cited in the Appendix. Increased emphasis on mappings in taxonomic research is warranted along with investigations of insular “species swarms” that constitute a microcosm of the early evolution of the class. The largest “species swarm” in the Diplopoda is Diplopoda itself!
Motyxia Chamberlin is comprised of eight species of bioluminescent xystocheirine millipeds in which the gonopodal solenomere arises at different positions, from basally and subbasally on the acropodite to being fused with the companion acropodal branch and detaching proximad or near midlength. Previous synonymies of Amplocheir Chamberlin and LuminodeslnllS Loomis and Davenport under Motyxia are confirmed as is its assignment to the tribe Xystocheirini, which is redefined. Component species are 111. Iwnw Chamberlin, the type species, monica Chamberlin, sequoiae (Loomis and Davenport), tularea (Chamberlin), sequoia (Chamberlin), pior Chamberlin, porrecta Causey and Tiemann, and tiemanni Causey. Motyxia sequoia is comprised oftwo races, the nominate and sequoia alia Causey and Tiemann; sequoia ollae Causey and Tiemann is properly a subspecies of tularea. 1I1otyxiapiorform secea is an invalid name without standing in nomenclature, and M. tejona Chamberlin, and M. expansa and exilis, both by Loomis, are placed in synonymy under M. monica, the oldest name for the southernmost species, as Polydesllws dissectus Wood is referrable to Xystocheir Cook. The bioluminescence is a continuous, neon-white glow of the entire dorsal surface including the antennae and legs. Its visibility at night suggests a warning function analogous to aposematic coloration. The phenomenon may observe a circadian rhythm, and controlled photoperiod experimentation may be productive.
Localities are documented for the milliped Abacion texense (Loomis, 1837) (Callipodida: Abacionidae) whose distribution forms both the northern and southern ordinal limits in the Western Hemisphere. The westernmost component of Abacion Rafinesque, 1820, A. texense is the only milliped species whose range spans the Mississippi and Pecos rivers and the Rio Grande. Distribution extremes are in Hennepin County (Co.), Minnesota, in the north; Terrell and Potter cos., Texas, in the west; Alcorn Co., Mississippi, in the east; and southwestern Tamaulipas, Mexico, in the south. Occurrences are projected for southeastern South Dakota, northwestern Alabama, and the southwestern periphery of Tennessee. The type series of A. texense consists solely of the male holotype, so a neotype will be needed if this individual is ever lost, because no paratypes were officially designated.
