Chordeumatida

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 : 42-45

Order Chordeumatida (Fig. 36, 41-44)

Chordeumatida inhabit the Tropics in both the Western Hemisphere and east/southeastern Asia, traversing the Equator in the latter; they also straddle the Tropic of Capricorn in Madagascar and occur in the South Temperate Zone in Australia, New Zealand, and South America ( Chile). They are prominent, if not dominant, in harshly cold environments and, except for the Himalayas, dominate high montane faunas of Europe and Central Asia, occurring among ice and snow in rocky crevices above the timberline in the Alps ( Geoffroy 1981, Pedroli-Christen 1993b, Read and Golovatch 1994, Spelda 1996, Kime and Golovatch 2000, Mikhaljova 2004, Golovatch and Kime 2010). We recognize 13 faunal areas ( Fig. 41 ) on all continents, with minor representation in South America and Africa (Mediterranean Coast). Sizes vary from a point locality (Kodiak I.) to large, continuous areas in North/Central America, Europe/ western Asia/Middle East/North Africa, and east/ southeastern Asia/East Indies. The seven southernmost areas, small and scattered, are in southern Chile (the only representation in South America), Madagascar, Australia, and New Figure 42. Known (solid lines) and projected (dashed lines) distributions of Chordeumatida in North and Zealand (both major islands); they represent Central America; a few records are available from the Gondwanan remnants as does that north of the dotted area, primarily the Great Basin Physiographic Equator in Sri Lanka and southernmost peninsu- Province.

lar India. Some of the most boreal chilognath records are chordeumatidans, which range northward in Scandinavia, Siberia, and Canada to areas that are beneath ice and snow for most of the year. Chordeumatida occur on Kodiak I., in the North Pacific Ocean, Madeira and the Azores in the Atlantic, Madagascar and Sri Lanka in the Indian, Indonesia and Papua New Guinea, and the following European islands: Balearic, Corsica, Sardinia, Sicily, Malta, Crete, Cyprus, and islands of the Adriatic, Ionian, and Aegean seas. While present on Newfoundland, Canada, Chordeumatida are absent from Caribbean islands, ones in the Atlantic ( Iceland, Bermuda, Canaries, Cape Verdes, Bioko, São Tomé and Principe, and Fernando Noronha), the Seychelles, Comoros, Réunion, Mauritius, Rodriguez, Maldives, Andamans, and Nicobars in the Indian, and Cocos, the Galapagos, Hawaiians, and all of Oceania in the Pacific. The boundary in the East Indies is uncertain because of a paucity of records; we encircle Papua New Guinea, but Chordeumatida plausibly occupy the geographically proximate Solomons and possibly expand southward to New Caledonia.

Kodiak I., harboring Tingupidae , is the

westernmost diplopod locality in North America/

New World ( Shelley et al. 2009a), and a large con-

tinuous area with an indefinite northern limit

spans the continent from ocean to ocean. It ex-

tends southward from the northern Alaskan Pan-

handle and the “Haines Triangle,” BC, Hudson

Bay, Ontario, and Newfoundland/Labrador, to

north of Lake Okeechobee, Florida, the Gulf Coast,

Los Angeles, California, southwestern Arizona, and

Chiriquí and Bocas del Toro provs., Panama (Fig.

42). The area thus terminates west of the Former

Canal Zone and omits southern peninsular Florida,

the Channel Islands, California, and the Yucatan

and Baja California peninsulas; we project exclu-

sion from most or all of Sinaloa and Colima and

western Sonora, Nayarit, and Jalisco. The north-

Figure 44. Distribution of Chordeumatida in central ern boundary, essentially those of the families Asia.

Caseyidae , Conotylidae , and Tingupidae ( Shelley 1993a; Shear and Shelley 2007b; Shelley et al. 2007, 2009a, b), curves south-southeastward from the “Haines Triangle,” BC, through Haines, Juneau, and Hyder, Alaska, then heads eastward to Dawson Creek and Tupper, BC, and northcentral Alberta. We project it continuing curvilinearly to Ft. Severn, on Hudson Bay, Ontario, then through southcentral Québec, southern Labrador, and encircling Newfoundland and Nova Scotia ( Shelley 1988, 2002b), where Caseyidae occur; we also encircle New England even though no indigenous records exist from coastal Maine, New Hampshire, and Rhode Island. However, records (Appendix) are available from central New Jersey, northern Delaware, the “eastern shore” of Maryland, and near or on the coasts of Virginia, North and South Carolina, and Georgia, the North Carolina ones being from Hatteras Island. Chordeumatida cover northern peninsular Florida, the southernmost records being from Highlands Co. north of Lake Okeechobee ( Loomis 1966 b, Shear 1972, Hoffman 1999, Shelley 2001, Appendix). The border then extends along the Gulf Coast to Tabasco, Mexico, where it cuts across the Yucatan Peninsula but encompasses Belize, where we report the first locality (Appendix), and continues to Panama ( Loomis 1964, 1968; Shear 1972; Hoffman 1999). Substantial sampling as has taken place on Barro Colorado Island, and the lack of chordeumatidans suggests that the order is absent from the Former Canal Zone.

The western boundary curves southward from Gustavus/Glacier Bay, Alaska, and encompasses the Alexander and Queen Charlotte Archipelagos and Vancouver and associated islands, BC ( Shelley 1990a, 2002b; Shear 2004; Shelley et al. 2007, 2009b). The southernmost California localities are in the Santa Monica , San Gabriel, and San Bernardino Mts., so Chordeumatida are unknown south of metropolitan Los Angeles and are absent from Orange, Riverside, San Diego, and Imperial cos. Records are sporadic, so we project the border through Yuma, Arizona, before crossing into Mexico around Nogales. The westernmost records in northern Mexico are from western Chihuahua and Durango, so we angle the border southsoutheastward to intersect the coast at Michoacan. In addition to new records (Appendix), works detailing occurrences, with or without maps, include Cook (1904), Palmén (1952), Causey (1961), Shear (1971, 1972, 1973b), Kevan (1983), Eskov and Golovatch (1986), Shelley (1988, 1990 a, 1993a, 2002b), Gardner and Shelley (1989), Shear (1999), Shelley et al. (2007, 2009a, b), and Shear and Shelley (2007b).

The other ordinal occurrence in the Western Hemisphere, and the only one in South America, is Eudigonidae in southcentral Chile; the southernmost locality is on Isla Grande de Chiloé (Appendix), but the true southern limit is unknown. Maps are available in Golovatch (1986a) and Shear (1988), and no additional records, published or unpublished, exist. The most proximate ordinal locality is in central Panama, ~ 4,640 km (2,900 mi) to the north.

European occurrences ( Fig. 43 ), basically as depicted by Kime (2000), stretch eastward from the Azores and Madeira ( Demange 1970) to Ukraine, Belarus, the Balkan States, and Finland ( Loksina and Golovatch 1979). North/south, the area extends from southwestern Norway and southern Sweden to the Atlas Mts. and Mediterranean coast of Morocco, Algeria, and Tunisia, the only ordinal occurrence in Africa ( Pocock 1892b; Silvestri 1896; Brolemann 1920, 1921; Hoffman 1980a; Mauriès 1990; Akkari et al. 2010b). In the southeast, the distribution extends eastward through Crimea and the Caucasus, tapering into a finger along the southern Caspian Sea in northern Iran and Turkmenistan ( Enghoff and Moravvej 2005). The southern border of this extension cuts through southeastern Turkey and turns abruptly southward to include coastal Syria, Lebanon, northern Israel, and the West Bank. Vicente and Enghoff (1999) reported Ceratosphys poculifer ( Brolemann, 1920) from the Canaries but speculated that it was introduced because it had not been taken there previously. Occurrence in the Balearic Islands was reported by Enghoff and Vicente (2000). Curcic et al. (2001, 2007) and Makarov et al. (2007) mapped occurrences in the Balkans, western Turkey, and Aegean islands. Additionally, a small, detached, circular area exists in Russia west of the Ural Mts. ( Golovatch 1992a, Mikhaljova 2004).

While absent from sub-Saharan Africa, Chordeumatida occur on Madagascar; distributions were mapped by Mauriès (1994, 1997b) in addition to the species list of Enghoff (2003). No other island off Africa harbors this order.

No prior, consolidated map exists for chordeumatidan distributions in Asia, so we combined species and generic maps with unmapped records. We recognize four areas in central Asia ( Fig. 41 , 44) beginning with Sri Lanka and southernmost peninsular India (Tamil Nadu and perhaps part of Kerala), also Gondwanan in origin. The next small area is in Nepal and may pervade adjacent India and China ( Shear 1979, 1987; Golovatch 1986b; Mauriès 1988b; Golovatch and Martens 1996). To the northwest, a larger, irregular area arises in Kashmir that curves through mountainous terrain in Pakistan, Afghanistan, Tajikistan, Kyrgyzstan, and western China (Xinjiang) to eastern Kazakhstan ( Shear 1979, Read and Golovatch 1994, Golovatch and Martens 1996). Finally, a slender, transverse, curvilinear area exists in Siberia atop the Mongolian and Chinese borders from northeastern Kazakhstan to the Amur River Region of Far Eastern Russia ( Golovatch 1980c, Shear 1990, Shelley 1993 a, Mikhaljova 1998, 2004, 2010). The western corner of the Siberian and the northern one of the Central Asian areas are only ~ 480 km (300 mi) apart, but continuity has not been demonstrated.

A large, irregularly crescent-shaped area covers eastern and southern, continental and insular Asia and eastern Australia. It extends, north/south, from southern Kamchatka, Sakhalin Island, and the Maritime Prov., Russia, through southcentral China, eastern Myanmar, southern Indonesia, and eastern Papua New Guinea, and along the east coast of Australia to Tasmania. It encompasses the Kurile and Ryukyu Islands, Japan, Taiwan, the Philippines, and the Korean Peninsula, where the western border angles southward into the Yellow and East China seas before expanding westward into China at the Yangtze Delta. From there, the border extends westward into Sichuan Prov. then curves through eastern Tibet into Myanmar and southward along the Irrawaddy River to the Andaman Sea (east of the Anadaman and Nicobar Islands) and Indian Ocean around Sumatra, Java, and southern Indonesia. Publications with records and/or partial maps include Murakami and Kawasawa (1976), Loksina and Golovatch (1979), Golovatch (1986b), Eskov and Golovatch (1986); Shear (1990, 1999), Mikhaljova (1993, 1998, 2000, 2004, 2010), Shear and Tanabe (1994), Mikhaljova and Basarukin (1995), Shear et al. (1997), Mikhaljova and Nefediev (2002), Mikhaljova and Korsós (2003), Korsós (2004), Enghoff et al. (2004), and Enghoff (2005).

In Australia. we project continuous occurrence along the entire eastern coasts of Queensland, New South Wales, and Victoria, and all of Tasmania. Chordeumatida also occupy a small, detached area in the southwestern corner of Western Australia. While now obviated by the Australian and Tasmanian websites, Golovatch (1986a) and Shear and Mesibov (1997) mapped Metopidiotrichidae in Tasmania, and though no maps are presented, Shear (2002) addressed Metopidiothrix Attems, 1907 , geographically. Shear (1999) also mapped the superfamily Heterochordeumatoidea globally, including areas in Australia and New Zealand, for which we encircle both major islands although records appear to be lacking from the North Island. We exclude Campbell I., subantarctic New Zealand, because of Johns’ (1964) suspicion that the Schedotrigona sp. occurring there was introduced from New Zealand proper. Golovatch’s (1986a) work on the then superfamily Conotyloidea is also relevant.

Figures

Figure 41. Known (solid lines) and projected (dashed lines) distributions of the Order Chordeumatida, parameters as in Fig. 1; a few records are available from the dotted area in North America. The upper arrow denotes the Tingupidae record from Kodiak Island, Alaska, USA, and the lower one indicates the known and projected ranges of Eudigonidae in Chile.

Figure 43. Distribution of Chordeumatida in Europe, North Africa, and the Middle East.