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A Revised, Annotated, Family-Level Classification of the Diplopoda

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A Revised, Annotated, Family-Level Classification of the Diplopoda Arthropoda Selecta 11 (3): 187207 © ARTHROPODA SELECTA, 2002 A revised, annotated, family-level classification of the Diplopoda Ðåâèçîâàííàÿ àííîòèðîâàííàÿ êëàññèôèêàöèÿ äèïëîïîä íà óðîâíå ñåìåéñòâà Rowland M. Shelley Ð.Ì. Øåëëè Research Lab., North Carolina State Museum of Natural Sciences, 4301 Reedy Creek Rd., Raleigh, North Carolina 27607 USA. Email: [email protected] KEY WORDS: classification, family level, Diplopoda. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: êëàññèôèêàöèÿ, óðîâåíü ñåìåéñòâà, Diplopoda. ABSTRACT: The arthropod class Diplopoda com- proposing a category at each hierarchical level. The prises two subclasses, 16 orders, and 144 families, following new ordinal-group authorship assignments which are arranged in an annotated modern classifica- are made: Polyxenida Verhoeff, 1934; Glomeridesmi- tion including alterations and higher taxa proposed since da, Platydesmida, Polyzoniida, Siphonocryptida, Spiro- publication of the last such work in 1980 (updated in bolida, Stemmiulida, Siphoniulida, Cambalidea (Spiro- 1982), which covered most taxa published through 1978. streptida), and Craspedosomatidea (Chordeumatida), The total number of families has grown by 24%, from all Cook, 1895; Siphonophorida Hoffman, 1980; Calli- 115 in 1980, and the largest and most diverse orders podida and Chordeumatida, both Pocock, 1894 (differ- remain the Chordeumatida and Polydesmida, with 47 ent publications); Polydesmida Pocock, 1887; Trigoni- and 30 families, respectively, as opposed to 35 and 28 ulidea (Spirobolida) Brölemann, 1913; and Leptodesmid- families in 1980. The suborder Siphonocryptidea Cook, ea and Strongylosomatidea (Polydesmida), both Bröle- 1895 (Polyzoniida), is elevated herein to full ordinal mann, 1916, and their junior synonyms, Chelodesmidea status, and the following ordinal-group taxa have been and Paradoxosomatidea, both Hoffman, 1967. The fol- newly proposed: the infraorder Oniscodesmoides Simo- lowing two new family-group assignments are made: nsen, 1990 (Polydesmida), and the suborders Pseudon- Sphaerotheriidae Koch, 1847, and Spirostreptidae Po- annolenidea Mauriès, 1983 (Spirostreptida) [now con- cock, 1894. As his phyla are the first proposed supra- sidered a synonym of Epinannolenidea Chamberlin, familial taxa, Brölemann is credited with the suborders 1922], Sinocallipodidea Shear, 2000 (Callipodida), and Trigoniulidea (Spirobolida) and Leptodesmidea and Heterochordeumatidea Shear, 2000 (Chordeumatida). Strongylosomatidea (Polydesmida), the first established One family published in 1976, Sakhalineumatidae Golo- in 1913 and the last two in 1916. The following new vatch (Chordeumatida), and two from 1978, Lankaso- synonymies are proposed: Nematozoniidae Verhoeff, matidae Mauriès and Megalotylidae Golovatch (both 1939, under Siphonorhinidae Cook, 1895 (Siphono- Chordeumatida), were not included in the previous phorida); Chelodesmidea Hoffman, 1967, and Sphaerio- classification, and the following 15 families have been desmidea Jeekel, 1971, under Leptodesmidea Bröle- newly proposed: Trichonemasomatidae, Chelojulidae, mann, 1916 (Polydesmida); and Paradoxosomatidea Pseudonemasomatidae, and Telsonemasomatidae, all Hoffman, 1967, under Strongylosomatidea Brölemann, by Enghoff, 1991 (Julida); Hoffmanobolidae Shelley, 1916 (Polydesmida). 2001 (Spirobolida); Bilingulidae Zhang & Li, 1981 (Spirostreptida: Cambalidea) [now considered a syn- ÐÅÇÞÌÅ: Êëàññ ÷ëåíèñòîíîãèõ Diplopoda âêëþ- onym of Pericambalidae Silvestri, 1909]; Paracortinidae ÷àåò äâà ïîäêëàññà, 16 îòðÿäîâ è 144 ñåìåéñòâà, Wang & Zhang, 1993, and Sinocallipodidae Zhang, êîòîðûå ïðåäñòàâëåíû â âèäå àííîòèðîâàíîé ñîâðå- 1993 (both Callipodida); and Kashmireumatidae Mau- ìåííîé êëàññèôèêàöèè, âêëþ÷àþùåé èçìåíåíèÿ è riès, 1982, Vieteumatidae Golovatch, 1984, Neocambri- âûñøèå òàêñîíû, ïðåäëîæåííûå ñî âðåìåíè ïóáëè- somatidae Mauriès, 1987, Reginaterreumatidae Mau- êàöèè ïîñëåäíåé ðàáîòû òàêîãî òèïà â 1980 ã. (îá- riès, 1988, Golovatchiidae Shear, 1992, Biokoviellidae íîâë¸ííîé â 1982 ã.), êîòîðàÿ ïîêðûâàëà áîëüøèí- Mriæ, 1992, and Altajellidae Mikhaljova & Golovatch, ñòâî òàêñîíîâ, îïóáëèêîâàííûõ ïî 1978 ã. Îáùåå 2001 (all Chordeumatida). A number of publication ÷èñëî ñåìåéñòâ âûðîñëî íà 24%, ñ 115 â 1980 ã., è dates are corrected, particularly Striariidea (Chordeu- íàèáîëåå êðóïíûìè è ðàçíîîáðàçíûìè îñòàþòñÿ matida), from Cook 1898/99 to 1896; Dalodesmidea îòðÿäû Chordeumatida è Polydesmida, ñ 47 è 30 ñå- (Polydesmida), from Hoffman 1977 to 1980; and Cam- ìåéñòâàìè, ñîîòâåòñòâåííî, â îòëè÷èå îò 35 è 28 podesmidae (Polydesmida: Leptodesmidea), from Cook ñåìåéñòâ â 1980 ã. Ðàíã ïîäîòðÿäà Siphonocryptidea 1895 to 1896. Authorships are based on the Principle of Cook, 1895 (Polyzoniida) çäåñü ïîäíÿò äî îòðÿäà. Coordination and are assigned to the first person(s) Òàêæå ïðåäëîæåíû ñëåäóþùèå èçìåíåíèÿ íà óðîâ- 188 Rowland M. Shelley íå îòðÿäà: èíôðàîòðÿä Oniscodesmoides Simonsen, da, as nearly as possible, a complete accounting of all 1990 (Polydesmida) è ïîäîòðÿäû Pseudonannolenidea supra-specific taxa established in the class through Mauriès, 1983 (Spirostreptida) [ðàññìàòðèâàåìûé ñåé- 1978, although some names proposed in that year are ÷àñ êàê ñèíîíèì Epinannolenidea Chamberlin, 1922], not included. Though dated as being published in 1979, Sinocallipodidea Shear, 2000 (Callipodida) è Hetero- it did not actually appear until June 1980 [Hoffman, chordeumatidea Shear, 2000 (Chordeumatida). Îäíî 1995], which is the official publication date for the ñåìåéñòâî, îïèñàííîå â 1976 ã., Sakhalineumatidae purposes of the Code and the 28 new taxa proposed Golovatch (Chordeumatida), è äâà â 1978 ã., Lanka- therein. A number of alterations to Hoffmans system somatidae Mauriès è Megalotylidae Golovatch (îáà â have been proposed in the past quarter century along Chordeumatida), íå áûëè âêëþ÷åíû â ïðåäûäóùèå with 18 new families and four ordinal-group taxa, êëàññèôèêàöèè. Ïðåäëîæåíû ñëåäóþùèå 15 ñåìåéñòâ: enough changes and additions in my view to justify a Trichonemasomatidae, Chelojulidae, Pseudonema- revised taxonomy to the family hierarchical level. Ad- somatidae, and Telsonemasomatidae, âñå Enghoff, 1991 ditionally, I elevate herein the suborder Siphonocryp- (Julida); Hoffmanobolidae Shelley, 2001 (Spirobolida); tidea Cook, 1895 (Polyzoniida) to full ordinal status, Bilingulidae Zhang & Li, 1981 (Spirostreptida: Cam- and I submerge the family Nematozoniidae Verhoeff, balidea) [ñåé÷àñ ðàññìàòðèâàåòñÿ êàê ñèíîíèì Peri- cambalidae Silvestri, 1909]; Paracortinidae Wang & 1939, under Siphonorhinidae Cook, 1895 (Siphon- Zhang, 1993 è Sinocallipodidae Zhang, 1993 (îáà â ophorida). Hoffmans [1980] arrangement was not Callipodida); à òàêæå Kashmireumatidae Mauriès, 1982, based on a cladistic analysis, which Enghoff [1984] Vieteumatidae Golovatch, 1984, Neocambrisomatidae provided down to the ordinal level, so this contribution Mauriès, 1987, Reginaterreumatidae Mauriès, 1988, is an amalgam of the two systems except for three orders Golovatchiidae Shear, 1992, Biokoviellidae Mriæ, 1992 that have been studied in detail: Julida [Enghoff, 1981, è Altajellidae Mikhaljova & Golovatch, 2000 (âñå â 1991], Chordeumatida [Shear, 2000a] and Polydesmi- Chordeumatida). Èñïðàâëåí ðÿä äàò îïóáëèêîâàíèÿ, â da [Simonsen, 1990]. I adopt the published arrange- ÷àñòíîñòè Striariidea (Chordeumatida) ñ Cook 1898/99 ments of the Julida and Chordeumatida, but for the íà 1896, Dalodesmidea (Polydesmida) ñ Hoffman 1977 reasons provided, I cannot accept all of the changes íà 1980 è Campodesmidae (Polydesmida: Lepto- Simonsen [1990] proposed for the Polydesmida. Hoff- desmidea) ñ Cook 1895 íà 1896. Àâòîðñòâî îñíîâàíî man recognized three subclasses Penicillata, Penta- íà ïðèíöèïå êîîðäèíàöèè è ïðèïèñàíî ïåðâûì ïåð- zonia, and Helminthomorpha nine superorders (two ñîíàì, ïðåäëîæèâøèì êàòåãîðèþ íà äàííîì óðîâíå in the Pentazonia and seven in the Helminthomorpha), èåðàðõèè. Ïðîèçâåäåíû ñëåäóþùèå èçìåíåíèÿ àâ- 15 orders, and 115 families. I adopt Enghoffs cladistic òîðñòâà íà óðîâíå îòðÿäà: Polyxenida Verhoeff, 1934; arrangement [1984] of two subclasses, Penicillata and Glomeridesmida, Platydesmida, Polyzoniida, Siphono- Chilognatha, with Pentazonia and Helminthomorpha cryptida, Spirobolida, Stemmiulida, Siphoniulida, Cam- becoming infraclasses in the latter. I see no reason not balidea (Spirostreptida) è Craspedosomatidea (Chordeu- to recognize two pentazonian superorders, Limaco- matida), âñå Cook, 1895; Siphonophorida Hoffman, morpha and Oniscomorpha, and I accept Enghoffs 1980; Callipodida è Chordeumatida, îáà Pocock, 1894 internal arrangement of the Helminthomorpha with (ðàçíûå ïóáëèêàöèè); Polydesmida Pocock, 1887; Tri- two subterclasses, Colobognatha and Eugnatha, the goniulidea (Spirobolida) Brölemann, 1913; à òàêæå latter comprising three superorders Juliformia, Nema- Leptodesmidea è Strongylosomatidea (Polydesmida), tophora, and Merocheta. The Diplopoda has grown îáà Brölemann, 1916, è èõ ìëàäøèå ñèíîíèìû, Chelo- considerably since 1978, from 115 to 144 families, a gain desmidea è Paradoxosomatidea, îáà Hoffman, 1967. of 29 families (24%); 18 are newly proposed; 12 result Ïðîèçâåäåíû ñëåäóþùèå èçìåíåíèÿ àâòîðñòâà íà from tribal and subfamilial elevations, some of which óðîâíå ñåìåéñòâà: Sphaerotheriidae Koch, 1847 è Spi- may be superfluous; and Nematozoniidae is eliminated. rostreptidae Pocock, 1894. Ïîñêîëüêó ôèëû Áð¸ëå- The most diverse orders remain the Chordeumatida and ìàííà (Brölemann) áûëè ïåðâûìè ïðåäëîæåííûìè òàêñîíàìè íàäñåìåéñòâåííîãî óðîâíÿ, åãî àâòîðñòâî Polydesmida, with 47 and 30 families, respectively,
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