Polyxenida

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 : 11-15

Order Polyxenida ( Fig. 3 -4)

The distribution of Polyxenida , the most poorly known of the major global/subglobal orders, is based more on inference, projection, and conjecture than are those of other taxa. Polyxenidans are best sampled by Berlese techniques, but most collectors, RMS included, search instead for readily visible chilognaths

that can be gathered by hand. Aside from Europe and the continental US, for which we document nine new states (Appendix), only scattered records exist, particularly from Asia, where they are unknown from Thailand (Enghoff 2005); for the Philippines, Y. Wang (1951) stated, “The subclass Pselaphognatha (= Penicillata) (is) represented in the Palaearctic and Oriental regions but (has) not so far been found in the Philippine Islands.” Despite these difficulties, we postulate broad, global occurrence for Polyxenida that encompasses five major areas, two smaller regions of Western Australia, and point sites in Kyrgyzstan, the Hawaiian Islands, and the Republic of the Marshall Islands (Ailinglapalap Atoll) ( Silvestri 1904; Chamberlin 1947a; Nishida 1994, 2002).

The northern borders are nebulous to varying degrees, so we show them by solid (Europe) or dashed (Asia) lines and omit that in North America because too few Canadian records exist to project a position. According to Palmén (1949), the northernmost European locality was at 63 o 30’N, but Polyxenida occur now at 65 oN on Vikna Island near Figure 4. Distributions of Penicillata and Polyxenida in Rörvik, North-Tröndelag, Norway ( Ellingsen 1910, represents northwestern North America. The dot the Meidell 1970), just south of the Arctic Circle (66 o northernmost record, from Haines, Haines Borough, 33' 39"N). It probably occurs farther north along Alaska, USA. the warmer, more moist Norwegian coast than deeper into Eurasia, but 65 oN is a convenient northern meridian for Europe if not also North America. The southernmost record, Propolyxenus patagonicus , in Santa Cruz Prov. , Argentina ( Silvestri 1903, 1905; Condé and Massoud 1974; Mauriès 1998; Nguyen Duy-Jacquemin and Geoffroy 2003; Short and Huynh 2010), is ~ 50 oS, some 16 o north of the Antarctic Circle (66 o 33' 39"S), which crosses the northern Antarctic Peninsula. Based on the known occurrence at 65 oN and the presence of southern beech ( Nothofagus ) litter layers, Penicillata / Polyxenida should be expected on Isla Grande de Tierra del Fuego and possibly even at Cape Horn (55 o 58’S). Investigating this possibility is a research objective of RMS.

In the New World, Polyxenida are known from southern Québec and the northern Alaskan Panhandle to Santa Cruz Prov. , Argentina ( Fig. 3 ). The northernmost site is Haines, Alaska (Fig. 4) (59 o 14' 09"N) ( Shelley 1990a), and in Canada, the order is also known from Nova Scotia and Ontario ( Jawlowski 1939 a, Judd 1967, Kevan 1983, Shelley 1988, 2002b). It also occupies Bermuda, and with occurrence on New Providence Island, Bahamas (an obscure record in Condé and Terver (1965)), and published localities from all Greater and several Lesser Antilles, Polyxenida can be reasonably projected for all Caribbean islands including Turks and Caicos. Fewer records exist from Mexico and Central/South America but enough to project it throughout the continental land mass southward to Santa Cruz Prov. , aside from the Atacama Desert region. The western border encompasses the Galapagos Islands ( Shear and Peck 1987), the Channel Islands of southern California ( Shelley 2002a), Vancouver and the Queen Charlotte Islands, BC, and the Alexander Archipelago, Alaska. For North America per se, the new records (Appendix) coupled with the few published localities, summarized in Condé (1996), point to widespread occurrence in the Continental US and suggest the same through border regions of adjacent Canadian provinces. Palmén (1952) did not report Polyxenida from Newfoundland, possibly because of improper sampling, but the island lies well below the northernmost European latitude and it should be expected there, farther north along the Atlantic Coast (Labrador and Québec), and maybe in southern Greenland. References with general or specific records include California ( Buckett 1964, Shelley 2002a), Florida ( Shelley 2001), Michigan ( Kane 1981, Snider 1991), Montana ( Loomis and Schmitt 1971, Nguyen Duy-Jacquemin 1976), New England ( Blake (1931), New York ( Bailey (1928), Texas ( Chamberlin and Mulaik 1941; Chamberlin 1943, 1947b; Stewart 1969), and North Carolina ( Shelley 1978, 2000a). Additional Canadian records for BC

(“mainland” and Vancouver Island), Ontario, and Nova Scotia are available in ( Kevan 1983; Shelley 1988, 1990a, 2002b).

Polyxenida have been collected less frequently in Mexico /Central America. In the former, they are known only from Baja California Sur, Nuevo León, and Veracruz ( Loomis 1966 a, 1968; Hoffman 1999; Bueno-Villegas et al. 2004), and in Central America, they have been reported from Guatemala, Costa Rica, and Panama ( Condé and Terver 1964; Loomis 1964, 1968). A map and table documenting Ancistroxenus Schubart, 1947 , and Lophoturus Brolemann, 1931 , in Nicaragua and Costa Rica is available in Nguyen Duy-Jacquemin (2002), which also covers the Greater and Lesser Antilles. In addition to Hoffman (1999), works lacking maps that report Polyxenida for the New World include Cuba ( Loomis 1933, Condé and Terver 1965; de la Torre y Callejas 1974; Gonzalez Oliver and Golovatch 1990); Hispaniola ( Haiti and the Dominican Republic including Beata Island) ( Loomis 1936, Condé and Terver 1965, Pérez- Asso and Pérez-Gelabert 2001, Pérez-Gelabert 2008); Puerto Rico including Mona Island ( Chamberlin 1950 a, Vélez 1967); Jamaica ( Loomis 1975; Condé and Terver 1979); Cayman Islands (Grand Cayman) ( Hounsome 1994); Bermuda ( Condé 1972); Bahamas (New Providence) ( Condé and Terver 1965); US Virgin Islands (St. John) ( Loomis 1970); St. Eustatius ( Condé and Terver 1965, 1979); St. Martin ( Condé and Terver 1979); St. Barthélemy ( Condé and Terver 1979); St. Kitts ( Loomis 1933); Guadeloupe ( Condé and Terver 1979, Nguyen Duy-Jacquemin 2002); Martinique ( Nguyen Duy-Jacquemin 2002); St. Vincent ( Loomis 1933, Condé and Terver 1979); the Grenadines (Mustique) ( Loomis 1933, Condé and Terver 1979); Tobago ( Nguyen Duy-Jacquemin 2002); and Trinidad ( Nguyen Duy-Jacquemin 2002).

South American countries with records, but not maps, include Surinam ( Condé and Terver 1964), Brazil ( Mello-Leitão 1925, Nguyen Duy-Jacquemin and Condé 1967, Brandão et al. 1991, Diniz and Brandão 1993), and Peru ( Condé 1980), and the map and table of occurrences in Nguyen Duy-Jacquemin (2002) covers northern Brazil, Colombia, Venezuela, Surinam, and French Guiana. The southernmost record, along the Santa Cruz River, Santa Cruz Prov. , Argentina, is ~ 800 km (500 mi) south of that of Chilognatha , in Parque Nacional los Alerces, Chubut Prov.

We enclose the Azores, Cape Verde, and Canary islands ( Enghoff 1993b, Vicente and Enghoff 1999); Polyxenida are unknown from Iceland, but the 65th parallel crosses the northern tip of the island and occurrence is plausible. Nearly all of Europe is occupied, or potentially so, including Great Britain, Ireland, and all Mediterranean and Aegean Islands. The northern border drops southward in Finland, swings eastward in Russia to near Moscow, and then angles southward into the Caucasus region and south of the Caspian Sea in northwestern Iran ( Enghoff and Moravvej 2005). In North Africa, the range spreads southward into the Sahara Desert in southern Algeria and probably extends directly southwestward to the Atlantic, though no records exist. Condé (1954a) documented Polyxenida in Morocco, and we project it for Libya, Egypt, the Gaza Strip, Israel, and northern Syria and Iraq. Enghoff (2006) cited localities in western Turkey, some mapped by Short and Huynh (2010), and we project the order throughout the country.

Kime (2000) did not map European Polyxenida , so we scoured papers for records. Ones with maps document occurrences in Scandinavia ( Palmén 1949, Meidell 1970, Enghoff 1976, Ishii 1988, Andersson et al. 2005), British Isles ( Blower 1985), Switzerland ( Pedroli-Christen 1993a), Hungary ( Korsós 1994a), and Belgium ( Kime 2004). Documentations without maps include Balearic Islands ( Enghoff and Vicente 2000), Balkan Peninsula ( Ceuca 1992, Mauriès et al. 1997, Curcic et al. 2002), France ( Geoffroy 1996), Germany ( Schubart (1925, 1934a), Italy (Stasser and Minelli 1984, Foddai et al. 1995), Luxembourg ( Kime 1996), Netherlands ( Jeekel 1953), Poland ( Jawlowski 1938, 1939b; Jedryczkowski 1982), Portugal ( Ladeiro 1943), and Switzerland ( Pedroli-Christen 1992). References documenting Polyxenida in eastern Europe, European Russia, and the former Soviet Republics include the Carpathians ( Loksina and Golovatch 1979), “Russian Plain” ( Loksina and Golovatch 1979; Golovatch 1992a), Georgia ( Lohmander 1936, Loksina and Golovatch 1979, Kobakhidze 1965, Talikadze 1984), Crimea ( Loksina and Golovatch 1979; Golovatch 1990 a, 2008), and Ukraine ( Chornyi and Golovatch 1993).

Polyxenida probably inhabit oases and high elevation inselbergs across the Sahara so as to occur subcontinuously from northern Scandinavia to southernmost Africa. It would be the only diplopod order to do so, but lacking published records from the desert heartland, we recognize a separate sub-Saharan region that extends into both the Atlantic and Indian Oceans and encloses Madagascar ( Condé and Nguyen Duy-Jacquemin 1962, Nguyen Duy-Jacquemin and Condé 1967, Enghoff 2003). The northern border traverses Sierra Leone, Liberia, Ivory Coast, Nigeria, Chad, Central African Republic, southern Sudan, and Ethiopia, reaching the Red Sea at the Eritrea / Djibouti border. Excluding Sokotra, it swings around the Seychelles ( Golovatch and Korsós 1992), Rodriguez ( Condé and Nguyen Duy-Jacquemin 1962; Nguyen Duy-Jacquemin and Condé 1967), Mauritius ( Condé and Nguyen Duy-Jacquemin 1962, Mauriès and Geoffroy 1999, Enghoff et al. 2004), and Réunion islands ( Condé and Nguyen Duy-Jacquemin 1962, Nguyen Duy-Jacquemin and Condé 1967, Enghoff 2003) in the Indian Ocean and encompasses St. Helena in the Atlantic ( Enghoff et al. 2004). Documented sub-Saharan occurrences include: Botswana ( Condé 1959, Hamer 1998), Ethiopia ( Chalande 1908, Ribaut 1922), Kenya ( Chalande 1908, Ribaut 1922, Nguyen Duy- Jacquemin and Condé 1967), Liberia ( Turk 1947), Mozambique ( Condé 1959), Republic of South Africa ( Condé 1959; Nguyen Duy-Jacquemin and Condé 1967; Hamer 1998), and Uganda ( Nguyen Duy-Jacquemin and Condé 1967). The polyxenidan from Lehututu, in the Kalahari Desert, is the only milliped record from Botswana, so we add the first chilognath, an unknown spirostreptidean ( Spirostreptidae ) from Ngamiland (Appendix).

In Asia, there is an isolated record in Kyrgyzstan ( Read and Golovatch 1994), and we join ones from India (Almora and Bombay) ( Turk 1947, Nguyen Duy-Jacquemin and Condé 1967, Golovatch and Martens 1996), Nepal ( Golovatch and Martens 1996), and Sri Lanka ( Pocock 1892a) into a separate region on the Indian Subcontinent, which likely connects with that in east/southeastern Asia, whose inner/western border is nebulous. We merge the latter with eastern Australia / Tasmania, New Zealand, and western Oceania, but the Marshall and Hawiian Islands are too remote. The northernmost Asian sites are in Honshu, Japan, and the Maritime Prov., Russia, and we incorporate the eastern periphery of China, all of the Indochina and Malay peninsulas, and most of Thailand, although (Enghoff 2005) cited no polyxenidans. The area extends eastward around the Bonin and Ryukyu Islands, Taiwan, all of Indonesia, Papua New Guinea, the Solomon Islands, New Caledonia, and Tonga. The only map, in Nguyen Duy-Jacquemin and Condé (1982), depicts records in the text. Other works with records cover Cambodia ( Enghoff et al. 2004), China (D. Wang and Mauriès 1996; provinces and localities are not provided), Indonesia (Java) ( Nguyen Duy-Jacquemin and Condé 1967, Murakami 1975a), Japan (mainland and Ryukyu Islands) ( Nguyen Duy-Jacquemin and Condé 1967; Murakami 1975a, b; Ishii 1988; Tsurusaki 2002), Eastern Malaysia (Sabah / Sarawak) ( Condé and Nguyen Duy-Jacquemin 1984, Lewis 1984a), Papua New Guinea (“mainland” and the Bismarck Archipelago) ( Enghoff et al. 2004), Asian Russia (Mikhaljova 1993, 1998, 2004, 2010), Singapore ( Ishii 1988), Taiwan ( Korsós 2004), and Vietnam ( Nguyen Duy-Jacquemin and Condé 1967, Enghoff et al. 2004). We project Polyxenida for the Philippines and eastern Myanmar.

Australian records, from the Australian milliped website and Short and Huynh (2010), include a coastal strip from Cape York to Tasmania and southeastern South Australia. Additional areas, both in Western Australia, are in the Kimberley Region and along the west/southwestern coasts.

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).