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

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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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  • Exploring Phylogenomic Relationships Within Myriapoda: Should High Matrix Occupancy Be the Goal?

    Exploring Phylogenomic Relationships Within Myriapoda: Should High Matrix Occupancy Be the Goal?

    bioRxiv preprint doi: https://doi.org/10.1101/030973; this version posted November 9, 2015. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. Exploring phylogenomic relationships within Myriapoda: should high matrix occupancy be the goal? ROSA FERNÁNDEZ1, GREGORY D. EDGECOMBE2 AND GONZALO GIRIBET1 1Museum of Comparative Zoology & Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA 2Department of Earth Sciences, The Natural History Museum, Cromwell Road, London SW7 5BD, UK 1 bioRxiv preprint doi: https://doi.org/10.1101/030973; this version posted November 9, 2015. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. Abstract.—Myriapods are one of the dominant terrestrial arthropod groups including the diverse and familiar centipedes and millipedes. Although molecular evidence has shown that Myriapoda is monophyletic, its internal phylogeny remains contentious and understudied, especially when compared to those of Chelicerata and Hexapoda. Until now, efforts have focused on taxon sampling (e.g., by including a handful of genes in many species) or on maximizing matrix occupancy (e.g., by including hundreds or thousands of genes in just a few species), but a phylogeny maximizing sampling at both levels remains elusive. In this study, we analyzed forty Illumina transcriptomes representing three myriapod classes (Diplopoda, Chilopoda and Symphyla); twenty-five transcriptomes were newly sequenced to maximize representation at the ordinal level in Diplopoda and at the family level in Chilopoda.
  • Some Aspects of the Ecology of Millipedes (Diplopoda) Thesis

    Some Aspects of the Ecology of Millipedes (Diplopoda) Thesis

    Some Aspects of the Ecology of Millipedes (Diplopoda) Thesis Presented in Partial Fulfillment of the Requirements for the Degree Master of Science in the Graduate School of The Ohio State University By Monica A. Farfan, B.S. Graduate Program in Evolution, Ecology, and Organismal Biology The Ohio State University 2010 Thesis Committee: Hans Klompen, Advisor John W. Wenzel Andrew Michel Copyright by Monica A. Farfan 2010 Abstract The focus of this thesis is the ecology of invasive millipedes (Diplopoda) in the family Julidae. This particular group of millipedes are thought to be introduced into North America from Europe and are now widely found in many urban, anthropogenic habitats in the U.S. Why are these animals such effective colonizers and why do they seem to be mostly present in anthropogenic habitats? In a review of the literature addressing the role of millipedes in nutrient cycling, the interactions of millipedes and communities of fungi and bacteria are discussed. The presence of millipedes stimulates fungal growth while fungal hyphae and bacteria positively effect feeding intensity and nutrient assimilation efficiency in millipedes. Millipedes may also utilize enzymes from these organisms. In a continuation of the study of the ecology of the family Julidae, a comparative study was completed on mites associated with millipedes in the family Julidae in eastern North America and the United Kingdom. The goals of this study were: 1. To establish what mites are present on these millipedes in North America 2. To see if this fauna is the same as in Europe 3. To examine host association patterns looking specifically for host or habitat specificity.
  • Diplopoda: Chordeumatida) Into Northern Coastal British Columbia and Southern Alaska

    Diplopoda: Chordeumatida) Into Northern Coastal British Columbia and Southern Alaska

    University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Center for Systematic Entomology, Gainesville, Insecta Mundi Florida 4-10-2009 Distribution Extensions of the Milliped Families Conotylidae and Rhiscosomididae (Diplopoda: Chordeumatida) into Northern Coastal British Columbia and Southern Alaska Rowland Shelley North Carolina State Museum of Natural Sciences Michael Medrano University of New Mexico, Albuquerque, NM William Shear Hampden-Sydney College, Hampden Sydney, VA Kristiina Ovaska Victoria, BC V9E 2B7, Canada Ken J. White British Columbia Ministry of Forests See next page for additional authors Follow this and additional works at: https://digitalcommons.unl.edu/insectamundi Part of the Entomology Commons Shelley, Rowland; Medrano, Michael; Shear, William; Ovaska, Kristiina; White, Ken J.; and Havard, Erin, "Distribution Extensions of the Milliped Families Conotylidae and Rhiscosomididae (Diplopoda: Chordeumatida) into Northern Coastal British Columbia and Southern Alaska" (2009). Insecta Mundi. 598. https://digitalcommons.unl.edu/insectamundi/598 This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Authors Rowland Shelley, Michael Medrano, William Shear, Kristiina Ovaska, Ken J. White, and Erin Havard This article is available at DigitalCommons@University of Nebraska - Lincoln: https://digitalcommons.unl.edu/ insectamundi/598 INSECTA MUNDI A Journal of World Insect Systematics 0071 Distribution extensions of the milliped families Conotylidae and Rhiscosomididae (Diplopoda: Chordeumatida) into northern coastal British Columbia and Southern Alaska Rowland M. Shelley Research Laboratory North Carolina State Museum of Natural Sciences MSC #1626 Raleigh, NC 27699-1626 USA Michael F.
  • The Millipedes and Centipedes of Chiapas Amber

    The Millipedes and Centipedes of Chiapas Amber

    14 4 ANNOTATED LIST OF SPECIES Check List 14 (4): 637–646 https://doi.org/10.15560/14.4.637 The millipedes and centipedes of Chiapas amber Francisco Riquelme1, Miguel Hernández-Patricio2 1 Laboratorio de Sistemática Molecular. Escuela de Estudios Superiores del Jicarero, Universidad Autónoma del Estado de Morelos, Jicarero C.P. 62909, Morelos, Mexico. 2 Subcoordinación de Inventarios Bióticos, Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, Tlalpan C.P. 14010, Mexico City, Mexico. Corresponding author: Francisco Riquelme, [email protected] Abstract An inventory of fossil millipedes (class Diplopoda) and centipedes (class Chilopoda) from Miocene Chiapas amber, Mexico, is presented, with the inclusion of new records. For Diplopoda, 34 members are enumerated, for which 31 are described as new fossil records of the orders Siphonophorida Newport, 1844, Spirobolida Bollman, 1893, Polydesmida Leach, 1895, Stemmiulida Pocock, 1894, and the superorder Juliformia Attems, 1926. For Chilopoda 8 fossils are listed, for which 3 are new records of the order Geophilomorpha Pocock, 1895 and 2 are of the order Scolopendromorpha Pocock, 1895. Key words Miocene, Mexico, Diplopoda, Chilopoda. Academic editor: Peter Dekker | Received 14 May 2018 | Accepted 26 July 2018 | Published 10 August 2018 Citation: Riquelme F, Hernández-Patricio F (2018) The millipedes and centipedes of Chiapas amber. Check List 14 (4): 637–646. https://doi. org/10.15560/14.4.637 Introduction Diplopoda fossil record worldwide (Edgecombe 2015). Two other centipedes have been reported in Ross et al. The extant species of millipedes and centipedes distributed (2016). Other records of millipedes have been mentioned across Mexico have been studied since the initial reports in the literature but are questionable because of a lack of Brand (1839) and Persbosc (1839).
  • Ordinal-Level Phylogenomics of the Arthropod Class

    Ordinal-Level Phylogenomics of the Arthropod Class

    Ordinal-Level Phylogenomics of the Arthropod Class Diplopoda (Millipedes) Based on an Analysis of 221 Nuclear Protein-Coding Loci Generated Using Next- Generation Sequence Analyses Michael S. Brewer1,2*, Jason E. Bond3 1 Department of Environmental Science, Policy, and Management, University of California Berkeley, Berkeley, California, United States of America, 2 Department of Biology, East Carolina University, Greenville, North Carolina, United States of America, 3 Department of Biological Sciences and Auburn University Museum of Natural History, Auburn University, Auburn, Alabama, United States of America Abstract Background: The ancient and diverse, yet understudied arthropod class Diplopoda, the millipedes, has a muddled taxonomic history. Despite having a cosmopolitan distribution and a number of unique and interesting characteristics, the group has received relatively little attention; interest in millipede systematics is low compared to taxa of comparable diversity. The existing classification of the group comprises 16 orders. Past attempts to reconstruct millipede phylogenies have suffered from a paucity of characters and included too few taxa to confidently resolve relationships and make formal nomenclatural changes. Herein, we reconstruct an ordinal-level phylogeny for the class Diplopoda using the largest character set ever assembled for the group. Methods: Transcriptomic sequences were obtained from exemplar taxa representing much of the diversity of millipede orders using second-generation (i.e., next-generation or high-throughput) sequencing. These data were subject to rigorous orthology selection and phylogenetic dataset optimization and then used to reconstruct phylogenies employing Bayesian inference and maximum likelihood optimality criteria. Ancestral reconstructions of sperm transfer appendage development (gonopods), presence of lateral defense secretion pores (ozopores), and presence of spinnerets were considered.