3. Revisions and Synoptic Studies on Other Orders and Families. While the research on the Xystodesmidae and the Eurymerodesmidae was in progress, studies were also proceeding on other NA taxa to advance the goal of documenting the continental fauna. They have involved every indigenous order in North America except Polyxenida, Glomerida, and Platydesmida; the last was revised by Gardner (1975) and a distributional paper is in preparation. The entire Western Hemisphere faunas of the order Callipodida, family Paeromopodidae (Julida), and subfamily Desmoninae (Polydesmida: Sphaeriodesmidae) have been studied, and the orders Siphonophorida and Polyzoniida have been revised to the extents possible today. Though incompletely reviewed, two polydesmidan families – Nearctodesmidae and Pyrgodesmidae – and one in the order Chordeumatida (Caseyidae) have been examined in depth as have the tribe Aniulini (Julida: Parajulidae) and the representatives of the Polydesmidae occurring west of the Continental Divide. The following additional genera have also been revised: Anelus (Spirobolida: Allopocockiidae), Onychelus and Piedolus (Spirobolida: Atopetholidae), Cambala (Spirostreptida: Cambalidae), and Auturus (Polydesmida: Euryuridae).

4. Faunal Treatments. It is important to consolidate information from disparate taxonomic studies so that others can grasp total faunas and the scope of the taxa that exist in the field. This category includes detailed faunistic studies with anatomical, habitat, and other information on each species, and “annotated checklists,” which are basically taxonomic lists with a modicum of additional information. Faunistic studies have been published on the eastern Piedmont and Kings Mountain regions of North Carolina (Shelley 1978a, Filka & Shelley 1980) and Canada (the eastern, western, and central regions and the country as a whole) (Shelley 1988, 1990b, 2002a; Shelley & LeSage 1989). Checklists have been prepared for the Coastal Zone of South Carolina, North Carolina as a whole, Florida, and California (Shelley 1978b, 2000a, 2001a, 2002b).

5. Miscellaneous Research. Outside the NA focus, species and genera of particular interest have been described or redescribed in the following orders and families: the polydesmidan families Paradoxosomatidae (a form from Namibia), Chelodesmidae (forms from Ecuador, Peru, Colombia, Trinidad and Tobago, and the Bahamas), Platyrhacidae (a species ranging from Nicaragua to Panama), and Rhachodesmidae and Pyrgodesmidae (both for species in Mexico) and the polyzoniidan family Hirudisomatidae (Mexico and Nepal); additionally, the new spirobolidan family Hoffmanobolidae was proposed for a form in Mexico. Through collaborations with colleagues, the first male in the order Siphoniulida has been characterized, this being the most poorly known and rarest order in the class (Sierwald et al. 2003); the suborder Sinocallipodidea (Callipodida) also has been addressed (Shear et al. 2003) as has the lone African representative of the order Siphonophorida (Shelley and Hoffman 2004). At the request of colleagues at the Bishop Museum, Honolulu, seven papers were developed on the Hawaiian fauna, most of which is introduced (Shelley 1998a-d, Shelley & Swift 1998, Shelley et al. 1998, Shelley & Golovatch 2000); the only known elements of the Hawaiian fauna that have not been treated are the representatives of the introduced family Spirobolellidae (Spirobolida) and the indigenous genus Nannolene (Spirostreptida: Cambalidae). As exotic millipedes receive little attention in general, works were produced on introduced representatives of the family Paradoxosomatidae (Polydesmida) on Pacific Islands and two species in the family Trigoniulidae (Spirobolida) that have been introduced to islands throughout the world and also to Africa and South & Central America (Shelley 1998e, Shelley & Lehtinen 1999). The most important efforts were geared toward advancing Diplopodology in general. Dr. Shelley led a collaborative effort to prepare a second generic and familial Nomenclator (Shelley et al. 2000) and individually developed a new family-level classification for the class (Shelley 2003).

One non-scientific effort that warrants mention here is the popularized booklet on centipedes and millipedes (Shelley 1999), produced to disseminate accurate information on these arthropods to non-biologists.

Future Research Plans. Future taxonomic and faunistic research will focus on four main topics, though other activities may arise. Results, updates, and new information will be presented periodically on this web site.

a) The Family Parajulidae. The major focus for the next several years will be the family Parajulidae (Julida), the dominant, most taxonomically complex NA family that has largely been ignored; there is also one species in Asia (Japan & China), so the taxon exhibits a “trans-Beringian connection.” The Parajulidae may contain upwards of 200 undescribed species, and it is the only indigenous family that occurs in Alaska and every county, state, and physiographic province in the lower 48 states; in Canada, it extends from the Queen Charlotte and Vancouver Islands, British Columbia, to southeastern Québec. North-south, the family ranges from southern Alaska and James Bay, Ontario, to El Salvador, a new country where its occurrence has not been officially recorded (Fig. 3). Advancing the knowledge and taxonomy of this family is vital to the goal of documenting the continental diplopod fauna as a whole. The initial emphasis has been on the tribe Aniulini, which ranges from the Atlantic Coast to Arizona, Alberta, and Québec; five papers have been published (Shelley 2000b-c, 2001b, 2002c, 2004) in which 15 species and subspecies have been described, and a summary one is in preparation. Beyond this tribe, the plan is to work generally from east to west across the continent revising genera and tribes in the process, some of which consititute worthy graduate student research projects.  At present a paper is being developed describing a new genus, species, and tribe that occurs at high elevations on isolated mountains in central New Mexico.  This appears to be a relict form that arose from the dichotomy that split off the Aniulini; Michael Medrano, doctoral graduate student at the University of New Mexico, is assisting with this study.

Fig. 3. Distribution of the Parajulidae in North and Central America

b) “Micro-Nearctodesmids.” The genera Phreatodesmus, Tidesmus, and Oodedesmus have been erected for small-bodied polydesmidans that occur around springs and in transiently moist spots in southwestern deserts (usually during cool weather seasons), but preliminary research indicates that three taxa are not warranted. These forms display diagnostic features of the Nearctodesmidae and appear to link the large-bodied genera in the Pacific Northwest (Shelley 1994) to ones in Jalisco, Mexico, and along the coast of southern California and Baja California Norté. A related form has recently been discovered in the Olympic Peninsula of Washington.

c) Ordinal Distributions and Mappings. In the study on the Sinocallipodidea, Shear et al. (2003) presented a full distribution map of the order Callipodida. Comparable efforts are planned for the other 15 orders.

d) Fauna of the “Ark-La-Tex” Region. Collaborative research since 2001 with C. T. McAllister and students at Texas A&M University at Texarkana has greatly enhanced knowledge of the fauna of the region containing Arkansas, Louisiana, Texas, and Oklahoma. It has also significantly advanced faunal knowledge of the area between the Mississippi River and the Central Plains, where distributions are typically poorly documented. One new species, Abacion wilhelminae (Callipodida: Abacionidae) has been described (Shelley et al. 2003a) and several new species in the order Chordeumatida await description. Additionally, the ranges of a half-dozen or so species have been dramatically extended; a paper has been published on Pleuroloma flavipes (Polydesmida: Xystodesmidae)(Shelley et al. 2003b); ones are in press on Scytonotus granulatus (Polydesmida: Polydesmidae) and Virgoiulus minutus (Julida: Blaniulidae); and ones are being prepared on the genus Narceus (Spirobolida: Spirobolidae) and the tribe Pachydesmidae (Polydesmida: Xystodesmidae) west of the Mississippi River. This research will continue and expand into the Plains proper of Oklahoma and Texas.

Selected References

Filka, M. E., & R. M. Shelley. 1980. The milliped fauna of the Kings Mountain region of North Carolina (Arthropoda: Diplopoda). Brimleyana, 4:1-42.

Gardner, M. R. 1975. Revision of the millipede family Andrognathidae in the Nearctic region. Mem. Pacific Coast Entomol. Soc., 5:1-61.

McAllister, C. T., R. M. Shelley, H. Enghoff, & Z. D. Ramsey. 2005. Distribution of the milliped Virgoiulus minutus (Brandt, 1841) (Julida: Blaniulidae).  Western North American Nat., in press.

Shear, W. M., & R. M. Shelley.  2004.  Introduction of the milliped, Cleidogona nantahala Shear, in New England, USA (Diplopoda: Chordeumatida: Cleidogonidae).  Ent. News, 115(2):71-77.

_____, _____, & H. Heatwole. 2003. Occurrence of the milliped Sinocallipus simplipodicus Zhang, 1993 in Laos, with reviews of the Southeast Asian and global callipodidan faunas, and remarks on the phylogenetic position of the order (Callipodida: Sinocallipodidea: Sinocallipodidae). Zootaxa, 365:1-20.

Shelley, R. M. 1978a. Millipeds of the eastern Piedmont region of North Carolina, U.S.A. (Diplopoda). J. Nat. Hist., 12:37-79.

_____. 1978b. Diplopoda, pp. 222-223, In: Zingmark, R. G., ed., An Annotated Checklist of the Biota of the Coastal Zone of South Carolina. University of South Carolina Press, Columbia, 364 pp.

_____. 1988. The millipeds of eastern Canada (Arthropoda: Diplopoda). Can. J. Zool., 66:1638-1663.

_____. 1990a (1989). Revision of the milliped family Eurymerodesmidae (Polydesmida: Chelodesmidea). Mem. American Entomol. Soc. No. 37:1-112.

_____. 1990b. A new milliped of the genus Metaxycheir from the Pacific Coast of Canada (Polydesmida: Xystodesmidae), with remarks on the tribe Chonaphini and the western Canadian and Alaskan diplopod fauna. Can. J. Zool., 68:2310-2322.

_____. 1994. The milliped family Nearctodesmidae in northwestern North America, with accounts of Sakophallus and S. simplex Chamberlin (Polydesmida). Can. J. Zool., 72:470-495.

_____. 1998a. Occurrence of the milliped Glyphiulus granulatus (Gervais) in the Hawaiian Islands (Spirostreptida: Cambalidea: Cambalopsidae). Bishop Mus. Occ. Paps. No. 56:36-37.

_____. 1998b. Interception of the milliped Rhinotus purpureus (Pocock) at Quarantine, and potential introduction of the order and family into the Hawaiian Islands (Polyzoniida: Siphonotidae). Bishop Mus. Occ. Paps. No. 56:54-55.

_____. 1998c. Occurrence of the milliped Trigoniulus corallinus (Gervais) on O’ahu and Kaua’i (Spirobolida: Pachybolidae: Trigoniulinae). Bishop Mus. Occ. Paps. No. 56:55-57.

_____. 1998d. Deletion of the milliped Vanhoeffenia tristriatus (Attems) from the Hawaiian fauna (Polydesmida: Dalodesmidae). Bishop Mus. Occ. Paps. No. 56:57-58.

_____. 1998e. Introduced millipeds of the family Paradoxosomatidae on Pacific Islands (Diplopoda: Polydesmida). Arthropoda Selecta, 7(2):81-94.

_____. 1999. Centipedes and Millipedes, with emphasis on North American fauna. Kansas School Nat., 45(3):1-15.

_____. 2000a. Annotated checklist of the millipeds of North Carolina (Arthropoda: Diplopoda), with remarks on the genus Sigmoria Chamberlin (Polydesmida: Xystodesmidae). J. Elisha Mitchell Sci. Soc., 116(3):177-205.

_____. 2000b. Parajulid studies II. The subgenus Hakiulus Chamberlin (Julida: Parajulidae: Parajulinae: Aniulini). Myriapodologica, 6(14):121-145.

_____. 2000c. Parajulid studies III. The genus Gyniulus Loomis (Parajulinae: Aniulini). Myriapodologica, 7(3):19-28.

_____. 2001a (2000). Annotated checklist of the millipeds of Florida (Arthropoda: Diplopoda). Insecta Mundi, 14(4):241-251.

_____. 2001b. A synopsis of the milliped genus Aniulus Chamberlin (Julida: Parajulidae: Parajulinae: Aniulini). Texas Mem. Mus. Speleol. Monogs., 5:73-94.

_____. 2002a. The millipeds of central Canada (Arthropoda: Diplopoda), with reviews of the Canadian fauna and diplopod faunistic studies. Can. J. Zool., 80:1863-1875.

_____. 2002b. Annotated checklist of the millipeds of California (Arthropoda: Diplopoda). Western North American Nat. Mon., 1:90-115.

_____. 2002c. The milliped genus Oriulus Chamberlin (Julida: Parajulidae). Can. J. Zool., 80:100-109.

_____. 2002. Narceus woodruffi Causey, a forgotten milliped species (Spirobolida: Spirobolidae).  Insecta Mundi, 16(1-3:25-29.

_____. 2003 (2002). A revised, annotated, family-level classification of the Diplopoda. Arthropoda Selecta, 11(3):187-207.

_____. 2003. A new polydesmid milliped genus and two new species from Oregon and Washington, USA, with a review of Bidentogon Buckett and Gardner, 1968, and a summary of the family in western North America (Polydesmida: Polydesmidae).  Zootaxa, 296:1-12.

_____. 2003. Redescription of the milliped Amphelictogon subterraneus bahamiensis Chamberlin, 1918, with an assessment of the family Chelodesmidae in the Bahamas (Polydesmida: Leptodesmidea). Zootaxa, 180:1-8.

_____. 2004a (2002). Parajulid studies V. The genera Pseudojulus Bollman and Arvechambus Causey (Parajulinae: Aniulini). Insecta Mundi, 16(4):191-204.

_____.  2004b.  The milliped family Pyrgodesmidae in the continental United States, with the first record of Poratia digitata (Porat) from the Bahamas (Polydesmida).  J. Nat. Hist., 38(9):1159-1181.

_____, & D. R. Whitehead. 1986. A reconsideration of the milliped genus Sigmoria, with a revision of Deltotaria and an analysis of the genera in the tribe Apheloriini (Polydesmida: Xystodesmidae). Mem. American Entomol. Soc. No. 35:1-223.

_____, & L. LeSage. 1989. Les mille-pattes (Diplopoda) de l’est du Canada: récoltes futures et perspectives de recherche. Rev. Ent. Québec, 34(1, 2):6-9.

_____, & S. F. Swift. 1998. The milliped order Julida in the Hawaiian Islands. Bishop Mus. Occ. Paps. No. 56:38-43.

_____, & P. T. Lehtinen. 1999. Diagnoses, synonymies, and occurrences of the pantropical millipeds, Leptogoniulus sorornus (Butler) and Trigoniulus corallinus (Gervais) (Spirobolida: Pachybolidae: Trigoniulinae). J. Nat. Hist., 33:1379-1401.

_____, & S. I. Golovatch. 2000. The milliped family Haplodesmidae in the Hawaiian Islands, with records of Prosopodesmus jacobsoni from Florida and Louisiana (Diplopoda: Polydesmida). Bishop Mus. Occ. Paps., 64:48-49.

_____, & R. L. Hoffman.  2004. A contribution on the South African milliped genus Nematozonium Verhoeff, 1939 (Siphonophorida: Siphonorhinidae). African Ent., 12(2):217-222.

_____, & H. Enghoff.  2004.  Titsona tida Chamberlin, 1962, a synonym of Choneiulus palmatus (Nĕmec, 1895) (Diplopoda: Julida: Blaniulidae), with new North American locality records.  Ent. News, 115(2):101-107.

_____, S. B. Bauer, & S. F. Swift. 1998. The milliped family Paradoxosomatidae in the Hawaiian Islands (Diplopoda: Polydesmida). Bishop Mus. Occ. Paps. No. 56:43-53.

_____, P. Sierwald, S. B. Kiser, & S. I. Golovatch. 2000. Nomenclator generum et familiarum Diplopodorum II. A List of the Genus and Family-Group Names in the Class Diplopoda from 1958 through 1999. Pensoft Publishers, Sofia, Bulgaria, 167 pp.

_____, & G. B. Edwards. 2002. Introduction of the milliped family Rhinocricidae in
Florida (Spirobolida).  Ent. News, 113(4):270-274.

_____, C. T. McAllister, & J. L. Hollis. 2003a. A new milliped of the genus Abacion Rafinesque, 1820 from Arkansas, U. S. A. (Callipodida: Abacionidae). Zootaxa, 170:1-7.

_____, _____, & S. B. Smith. 2003b. Discovery of the milliped Pleuroloma flavipes (Polydesmida: Xystodesmidae) in Texas, and other records from west of the Mississippi River.   Ent. News, 114(1):2-6.

____, ____, & Z. D. Ramsey. 2005. Discovery of the milliped, Scytonotus granulatus (Say, 1821) in Oklahoma, with a new record from Alabama and a review of its distribution (Polydesmida: Polydesmidae).  Western North American Nat., in press.

Sierwald, P., W. A. Shear, R. M. Shelley, & J. E. Bond. 2003. Millipede phylogeny revisited in the light of the enigmatic order Siphoniulida. J. Zool. Syst. Evol. Res., 41:87-99.