Polydesmida

Shelley, Rowland M. & Golovatch, Sergei I., 2011, Atlas of Myriapod Biogeography. I. Indigenous Ordinal and Supra-Ordinal Distributions in the Diplopoda: Perspectives on Taxon Origins and Ages, and a Hypothesis on the Origin and Early Evolution of the Class, Insecta Mundi 2011 (158), pp. 1-134 : 47-50

Order Polydesmida ( Fig. 46-50 )

Generally considered the most speciose order, though second to Chordeumatida in the number of families ( Shelley 2003 a, 2007a), Polydesmida occur natively on all continents and on islands in all oceans and seas; they are particularly dominant in the Himalayas, where over half of the ~ 200 species are polydesmidans ( Golovatch and Kime 2010). While not crossing the Arctic Circle as do Polyxenida , Polyzoniida , and Julida ( Fig. 3 , 20 , 26 ), and hence not representing the northern limit for either Diplopoda or Chilognatha, Polydesmida include the southernmost diplopod, Notonaia campbellensis Johns, 1970 (Dalodesmidea: Dalodesmidae ), on Campbell I. (52 o 36' 18"S) in the subantarctic islands of New Zealand ( Johns 1964, 1970).

We recognize 21 areas for Polydesmida ( Fig. 46 ) that vary in size from point localities in North America and Saudi Arabia to large, continuous areas in the Western Hemisphere (eastern North America to southern South America), sub-Saharan Africa, and eastern Atlantic islands to Australia / New Zealand / Oceania. The northernmost localities are Yakutat, Alaska, USA, Steinkjer, Norway, and Östersund and Längselo, Sweden, and while Polydesmida inhabit many islands, they are notably absent from Bermuda, Newfoundland, Iceland, and the Hawaiians. Works with global familial maps, all obsolete, include Jeekel (1968, Paradoxosomatidae ), ( Mauriès 1974b, Platyrhacidae sensu Hoffman (1980a)) , Hoffman (1962, 1978, Chelodesmidae , Xystodesmidae , Platyrhacidae sensu Hoffman (1980a)) , Simonsen (1990, Pyrgodesmidae , Trichopolydesmidae , Nearctodesminae , Macrosternodesminae , Fuhrmannodesmidae , Dalodesmidae , Polydesmidae , Cyrtodesmidae ), Simonsen (1992, Cryptodesmidae , Haplodesmidae ), Golovatch (1996a, Pyrgodesmidae , on a blank map without reference lines), and Djursvoll et al. (2000, Polydesmidae ).

The map of the order/nominate family by Simonsen (1990) shows Polydesmida as continuous throughout North America from southern Canada southward, but such is not the case. Continuity is through the forested biomes of eastern North America only; 13 separate areas of varying sizes exist in the west, seven of which are point localities with two being from caves. The eastern area joins with “mainland” Mexico, Central America, Caribbean and eastern Pacific Islands, and South America to form a large area extending from southern Québec and James Bay, Ontario, to southern Chile ( Isla Grande del Chiloé) and Argentina (Nequen/Rio Negro provs.). It lies generally east of Lake Superior in Canada and east of the central Plains in the US, with the western border angling south-southwestward in Texas to the Rio Grande west of the confluence with the Pecos River. The northernmost record is Pseudopolydesmus canadensis (Newport, 1844) ( Polydesmidae ), from the Albany River at James Bay, Ontario, from which the area slopes southwestward to Sault Ste. Marie and southeastward through Québec to the Gaspé Peninsula (Fig. 47). The eastern border cuts across New Brunswick and Maine to the Atlantic coast and then across Massachusetts and Connecticut to Long Island Sound. No polydesmidans are known from Long Island itself, but they do occur in southern New Jersey, Delmarva, and Cape Hatteras National Seashore and Bald Head Island, North Carolina. The eastern border then encircles the Bahamas ( Shelley 2003c, 2007c) and the Antilles, but excludes Turks and Caicos, while the western traverses northern Minnesota to Fargo, North Dakota, then heads southward through eastern South Dakota and Nebraska before angling southsouthwestward to the Rio Grande, northeastern Coahuila, and passing through Zacatecas and Nayarit to the Pacific at Jalisco.

Of the 13 western areas in the US and Canada, only one is substantial in size and continuity. It extends along the Pacific Coast from Yakutat and Gustavus/Glacier Bay, Alaska (Fig. 48), to south of Ensenada, Baja California Norte, Mexico, encompassing all of the Alexander Archipelago and the Queen Charlotte, Vancouver, and associated islands but excluding the Channel Islands, California. Its inner boundary extends through Hyder, Alaska, and Stewart, Terrace, and Manning Provincial Park, BC, into Washington; from the south, it extends northward through San Diego, Riverside, and San Bernardino cos., California, angles through the Tehachapi Mts., and runs along the crest of the Sierra Nevadas and Cascades to central Oregon. There, the boundary turns eastward through the Blue Mts., Oregon, and the Bitterroots, Idaho, and nips the Continental Divide in Montana. It then heads westward/northward into Canada along the Divide and the BC/ Alberta border to Yoho National Park, where it swings westward into BC and southward into Washington. Two detached areas exist in western Canada, a point locality east of the Divide in Jasper National Park, Alberta ( Shelley 2007b), and a small one in the Shuswap Highlands, BC ( Shelley 1994a).

Polydesmida occupy nine smaller areas in the western US, one in the Rockies of eastern Idaho / western Wyoming that curves through the Wasatch Mts. to south of Great Salt Lake, Utah. A few localities cluster in the Henry Mts., Utah, and there are cave localities to the east and west in Colorado and Nevada. To the south, a slightly larger area covers most of Arizona with scattered sites on forested mountaintops and around springs and rivers, extending from the Colorado River/Grand Canyon National Park, in the northwest, to the Chiricahua and Graham Mts., in the southeast. Three regions of New Mexico are occupied – the Manzano and Sandia Mts. in the north-center, the Black Mts. and associated ranges in the southwest, and a narrow area extending from the Capitans and Sierra Blancas to the Guadalupes in Texas. Additionally, there are point localities in the Chisos Mts., Big Bend National Park, Texas, and on the Gulf of California, Baja California Sur. Other than the last site and northwestern Baja California Norte, no polydesmidans are known from the Baja Peninsula; maps showing continuous ranges here for Pyrgodesmidae ( Simonsen 1990, Golovatch 1996a) are erroneous.

Works with maps, some outdated, include Hoffman (1962, Xystodesmidae , Platyrhacidae sensu Hoffman (1980a)) , Buckett and Gardner (1968, Xystodesmidae ), Shelley (1980, 1982, 1987, 1990b, c, 1992, 1993b, c, d, e, f, 1994a, b, 1995, 1996f, g, 1997a, b, c, 2000b, c, 2003b, c, 2004, Eurymerodesmidae , Euryuridae , Nearctodesmidae , Platyrhacidae , Polydesmidae , Pyrgodesmidae , Rhachodesmidae , Sphaeriodesmidae [ Desmoninae ], Xystodesmidae ), Shear (1984, Polydesmidae ), Withrow (1988, Polydesmidae ); Whitney and Shelley (1995, Nearctodesmidae ), Shelley and Whitehead (1986, Xystodesmidae ), Shelley and Shear (2006, Nearctodesmidae ), Shear and Shelley (2007a, Macrosternodesmidae ), and Shelley and McAllister (2007, Xystodesmidae ). Ordinal limits in eastern Canada are depicted by Shelley (1988), and Hoffman (1979a) reported Chelodesmidae from Cocos Island, Costa Rica.

South American occurrence is continuous with that to the north, and we expand the border westward in the Pacific to incorporate the Galapagos and Juan Fernandez Islands, where indigenous, if not endemic, species have been recorded in Pyrgodesmidae and Dalodesmidae , respectively ( Shear and Peck 1987, Chamberlin 1957). The area encompasses the entirety of all countries but Argentina and Chile, as the southernmost records, all in Dalodesmidae , are Tsagonus muermo Chamberlin, 1957 , from de los Lagos Dist., Chile ( Chamberlin 1957), and Anaulacodesmus lacustris and Monenchodesmus inermis nahuelhuapiensis , both by Schubart, 1954, from Parque Nacional Nahuel Huapi, Neuquén and Rio Negro provs., Argentina ( Schubart 1954, Demange 1963, Mauriès 1998). We could not find records from two areas; the more northern, also blank for the class, encompasses the Atacama Desert and arid regions in Chile and adjacent Peru and Bolivia. It lies west of Lake Titicaca, as Kraus (1954) described Catharosoma titicacaensis (Paradoxosomatidae) and Cryptogonodesmus peruvianus (? Fuhrmannodesmidae ) from the western side of the lake; later, he ( Kraus 1960b) recorded Mestosoma alticolum (Attems, 1930) and described M. orobium (Paradoxosomatidae) from the lake region in general. The second blank area is in central Argentina east of the Andean Cordillera and west of Buenos Aires. Works with maps include Jeekel (1968, Paradoxosomatidae ), Demange and Silva (1976, Dalodesmidae ), Hoffman (1982b, 1990a, Chelodesmidae ), Simonsen (1990, Cyrtodesmidae ), and Vohland (1998, Aphelidesmidae ); ones with species lists and localities, particularly for southern South America, include Kraus (1956), Jeekel (1965a), and Golovatch (2005).

In sub-Saharan Africa, Polydesmida occur generally south of the Senegal and Niger rivers, the curvilinear border passing through Mali, Niger, northern Nigeria, and southern Chad, then angling northward through Sudan to the Red Sea in Eritrea. The entire area south of this line is occupied including Bioko, except for the dotted area that encompasses western Zambia, southeastern Angola, eastern Namibia, northern Republic of South Africa, and all of Botswana. To some degree this void reflects lack of sampling, but it also corresponds to the preponderance of the generally hot, dry, and inhospitable Kalahari and Namib deserts. Polydesmida also occur on Madagascar, Sokotra, and the Comoros, Mauritius, Rodriguez, and the Seychelles Islands ( Butler 1876, 1879; Mauriès 1980b; Hoffman 1990b; Golovatch 1992b; Mauriès and Geoffroy 1999; Enghoff 2003, VandenSpiegel and Golovatch 2007). Works with maps include SchiØtz (1966, Campodesmidae ), Jeekel (1968, Paradoxosomatidae ), Hoffman (1968b, 1978, 1990b, 2004, 2005, Paradoxosomatidae , Oxydesmidae , Gomphodesmidae ), Simonsen (1990, Ammodesmidae ), Mauriès and Heymer (1996, Fuhrmannodesmidae ), Shelley and Crawford (1996, Paradoxosomatidae ), Alderweireldt and Enghoff (1998, Oxydesmidae ), and VandenSpiegel and Golovatch (2003, Ammodesmidae ). Oxydesmidae and Fuhrmannodesmidae occupy Sokotra ( Hoffman 1990 b, Golovatch and Mauriès 2007), and Cryptodesmidae occur on Bioko ( Hoffman 1975), to which we add Pyrgodesmidae (Appendix) . Chamberlin (1952) listed chelodesmid (prepodesmine) species with countries or regions of occurrence as did Jeekel (2003b, 2004b, 2006b) for Paradoxosomatidae , and valuable species lists exist in Cook (1898) and Hoffman (1953, 1968b).

Occurrence in the European part of the Euroaustralasian area, which extends around Atlantic islands and the northern periphery of Africa, follows Kime (2000) with modifications in Russia and eastern Europe. In North Africa, Polydesmida occupy the Atlantic and Mediterranean coasts of Morocco, extending inland into the Atlas Mts. and the Mediterranean coasts of Algeria, northern Tunisia, and the Benghazi region of Libya. Tabacaru (1995) and Golovatch (2007) reported Oxydesmidae from Israel, and Shelley (2009) mapped paradoxosomatid occurrences, thereby accounting for the eastern Mediterranean dip.

The European and Asian parts of this area connect through an “isthmus” in Pakistan and Afghanistan ( Jeekel 2003c) that extends westward south of the Caspian Sea, curves southward through Iraq, Syria, and Jordan, then northward through Israel to the Mediterranean. The eastern border curls northeastward through the Central Asian Republics ( Spelda et al. 1998) and around uninvestigated western China, then expands eastward as a narrow “peninsula” in Siberia atop western Mongolia. A substantial area east, north, and west of the Caspian Sea lacks records; it expands westward into eastern Ukraine and eastward into central Kazakstan, Uzbekistan, and Turkmenistan. Two sites are sufficiently detached to be shown separately, that near Riyadh, Saudi Arabia (Shelley 2009), and in the Zagros Mts., Iran.

The eastern and southern Asian section expands curvilinearly northeastward through central China and Manchuria into Khabarovsk Prov., Russia, crossing into the Sea of Okhotsk north of Sakhalin Island. It then turns southeastward through the Kurile Islands and traverses the Pacific east of Bonin, the Marshall Islands, Micronesia, Fiji, and New Zealand. Fiji has no endemic or presumably indigenous Paradoxosomatidae ( Jeekel 1972) , but he ( Jeekel 1980) regarded their Cryptodesmidae as native, so we draw the Oceania border east of Fiji and Chatham Island, New Zealand (Appendix). From the isthmus in Central Asia, the border curves southward through western India to encompass the entire Indian Peninsula, Sri Lanka, all of southeastern continental Asia, the Andaman and Nicobar Islands, and all of Indonesia and Papua New Guinea. We continue this area through eastern and southern Australia, including nearly all of Victoria and Capital Territory and about halfway into South Australia. Polydesmida occupy the westernmost coastal region of Western Australia, a small interior area in the Kimberley Region, and Darwin vicinity ( Paradoxosomatidae , Haplodesmidae , Appendix), the first precise diplopod localities from Northern Territory.

A bibliographic catalog on Platyrhacidae ( Jeekel 2007) is available for the Asian/Australian part of this area. Papers with maps regarding the continent and islands include Jeekel (1965b, Paradoxosomatidae ), Hoffman and Lohmander (1968, Paradoxosomatidae , Xystodesmidae ), Johns (1979, Polydesmida as a whole), Hoffman (1973, 1975, 1997, Cryptodesmidae , Platyrhacidae ), Golovatch (1979, 1986b, 1987b, c, 1988, 1990c, d, 1992b, 1994b, c, 1996b, Fuhrmannodesmidae , Opisotretidae , Paradoxosomatidae , Polydesmidae ), Simonsen (1990, Haplodesmidae [= Doratodesmidae ], Opisotretidae ), Golovatch and Enghoff (1994, Paradoxosomatidae ), Tanabe and Shinohara (1996, Xystodesmidae ), Mikhaljova (1998, Paradoxosomatidae , Polydesmidae ), Tanabe (2002, Xystodesmidae ), Mesibov (2002, 2003a, b, c, 2004a, b, c, 2005a, b, Dalodesmidae , Haplodesmidae ), Golovatch and Nguyen Duc (2006, Platyrhacidae ). Jeekel (1973, 2000) listed species from the Solomon Islands, and Attems (1914a), Jeekel (1967), and Hoffman (1980a) reported Eustrongylosoma (Paradoxosomatidae) from Pohnpei, Federal Republic of Micronesia. In addition to Enghoff (2006) and Enghoff and Moravvej (2005), Iranian and Caucasus records are available in Golovatch (1981a), and Jeekel (2006c) published New Zealand dalodesmid localities. We report Dalodesmidae (P. Johns, pers. comm. to RMS) from Chatham I. in the South Pacific Ocean, ~ 1,136 km (710 mi) east of the South Island. To the best of our knowledge, this is the first report of Polydesmida from this archipelago.

Figures

Figure 3. Known (solid lines) and projected (dashed lines) indigenous distributions of the Subclass Penicillata and Order Polyxenida. The arrow denotes the record from the Federal Republic of the Marshall Islands; the upright question mark (?) denotes the unsubstantiated record of Propolyxenus patagonicus (Silvestri, 1903) from Santa Cruz Prov., Argentina (Mauriès 1998); and the inverted question mark (¿) indicates uncertain occurrence on the Yucatan Peninsula, Mexico. Other parameters as in Fig. 1. Too few records exist from Canada to project a northern boundary in North America, and the isolated record from central Asia (Kyrgyzstan) is too distant to be combined with an area where records cluster. No records are available from the dotted areas in India and North America, and the only one from that in Africa is Chilexenus schultzei (Attems, 1909) from Botswana (Hamer 1998).

Figure 26. Distribution of the Order Julida.

Figure 45-46. Distributions. 45) Distribution of the Order Stemmiulida. 46) Known (solid lines) and projected (dashed lines) distributions of the Superorder Merocheta and the Order Polydesmida, parameters as in Fig. 1. No records or samples are available from the dotted areas.

Figure 49-50. Distributions. 49) Distributions of Merocheta/Polydesmida in Europe, North Africa, the Middle East, and western Asia; the dotted line circumscribes an area lacking both records and samples. 50) Distributions of Merocheta/Polydesmida in Central Asia; the dotted line circumscribes an area lacking both records and samples.

Figure 19-20. Distributions. 19) Distribution of Platydesmida in Europe, the Middle East, and North Africa. 20) Known (solid lines) and projected (dashed lines) distribution of the Order Polyzoniida, parameters as in Fig. 15.