Julida

Megaphyllum unilineatum, copyright Zdeněk Hyan.

Belongs within: Eugnatha.
Contains: Cylindroiulus, Leptoiulus, Julus.

The Julida are a group of millipedes with pleura fused with terga, and a gnathochilarium with meeting stipes. Members of the families Paeromopodidae and Julidae bear distinct, fine longitudinal striation all around the metazonal segments; members of other families have such striation absent or on the lower sides of the segments only (Hoffman 1990).

Characters (from Hoffman 1990): Gnathochilarium with stipes broadly in contact medially, with small promental sclerite isolated in front of median commissure and larger transverse mental plate (or two plates) at base; sterna, pleura, and terga coalesced into rigid cylindrical rings, with pleural sclerites completely coalesced with lower ends of terga, no suture line remaining; segment 3 without legs.

<==Julida [Symphyognatha, Zygocheta]
    |  i. s.: Tachypodoiulus [incl. Ischiolobus Attems 1951] TKM93
    |           `--T. niger (Leach 1815) [incl. *Ischiolobus niger Attems 1951] TKM93
    |--Paeromopodidae [Californiulidae] H90
    |    |--Paeromopus H90
    |    |--Californiulus H90
    |    `--Aprosphylosoma H90
    |--Nemasomatidae MG00
    |    |--Telsonemasoma H90
    |    |--Thalassisobates littoralis KAA-W00
    |    |--Nemasoma varicorne Koch 1847 TKM93 [=Isobates varicornis KAA-W00]
    |    |--Chelojulus Enghoff 1982 H00
    |    |    `--C. sculpturatus WA04
    |    `--Orinisobates Lohmander 1933 MG00
    |         |--O. microthylax Enghoff 1985 MG00
    |         `--O. sibiricus (Gulička 1963) [=Isobates sibiricus] MG00
    |--Blaniulidae [Isobatidae] TKM93
    |    |  i. s.: Nopoiulus kochii (Gervais 1847) KAA-W00
    |    |         Proteroiulus KAA-W00
    |    |           |--P. fuscus (Am Stein 1857) [=Blaniulus fuscus] TKM93
    |    |           `--P. varicornis KAA-W00
    |    |         Blaniulus TKM93
    |    |           |--B. guttulatus (Fabricius 1798) TKM93
    |    |           `--B. venustus A01
    |    |         Skleroprotopus Attems 1901 [Isobatini, Protoiulidae] A01
    |    |           `--*S. confucius Attems 1901 A01
    |    |--+--Virgoiulus H90
    |    |  `--Okeanobates H90
    |    `--Zosteractinidae H90
    |         |--Zosteractis H90
    |         `--Ameractis H90
    |--Parajulidae H90
    |    |  i. s.: ‘Julus’ telluster Scudder 1878 RB93
    |    |--Aliulus H90
    |    |--Aniulus bollmani KAA-W00
    |    |--Arvechambus H90
    |    |--Ethojulus H90
    |    |--Gyniulus H90
    |    |--Hakiulus H90
    |    |--Illiulus H90
    |    |--Okliulus H90
    |    |--Oriulus H90
    |    |--Pseudojulus H90
    |    |--Ptyoiulus H90
    |    |--Teniulus H90
    |    |--Uroblaniulus H90
    |    |--Apacheiulus H90
    |    |--Gosiulus H90
    |    |--Codiulus H90
    |    |--Bollmaniulus H90
    |    |--Karteroiulus H90
    |    |--Mulaikiulus H90
    |    |--Saiulus H90
    |    |--Simiulus H90
    |    `--Spathiulus H90
    `--Julidae KAA-W00
         |--Brachyiulus pusillus KAA-W00
         |--Allajulus nitidus (Verhoeff 1891) H90, TKM93 [=Cylindroiulus nitidus KAA-W00]
         |--Cylindroiulus MG00
         |--Hypsoiulus alpivagus (Verhoeff 1897) TKM93
         |--Leptoiulus TKM93
         |--Pachypodoiulus eurypus (Attems 1895) TKM93
         |--Unciger foetidus (Koch 1838) TKM93
         |--Pachyiulus fuscipes A01
         |--Glyphiulus granulatus A01
         |--Julus KAA-W00
         |--Leptophyllum (Anuroleptophyllum Attems 1901) A01
         |    `--L. (*A.) caucasicum Attems 1901 A01
         |--Pacifiiulus Mikhaljova 1982 MG00
         |    `--P. amurensis (Gerstfeldt 1859) [incl. P. imbricatus] MG00
         |--Ommatoiulus TKM93
         |    |--O. moreletti DAF07
         |    `--O. sabulosus (Linnaeus 1758) [=Schizophyllum sabulosum] TKM93
         |--Enantiulus TKM93
         |    |--E. nanus (Latzel 1884) [=Julus nanus, Leptophyllum nanum, Metaleptophyllum nanum] TKM93
         |    `--E. pelidnus (Latzel 1884) TKM93
         |--Ophyiulus KAA-W00
         |    |--O. nigrofuscus (Verhoeff 1894) TKM93
         |    |--O. pilosus (Newport 1842) TKM93
         |    `--‘Iulus (*Leucoiulus)’ tirolensis Verhoeff 1894 (n. d.) TKM93
         |--Sibiriulus Gulička 1972 MG00
         |    |--S. altaicus (Gulička 1972) [=Cylindroiulus (Sibiriulus) altaicus] MG00
         |    |--S. dentiger (Gulička 1963) [=Cylindroiulus (Sibiriulus) dentiger] MG00
         |    `--S. multinicus Mikhaljova in Mikhaljova & Golovatch 2000 MG00
         `--Megaphyllum Verhoeff 1894 MG00
              |--M. austriacum (Latzel 1884) [=Julus austriacus] TKM93
              |--M. projectum (Verhoeff 1894) [=Chromatoiulus projectus] TKM93
              |--M. sjaelandicum (Meinert 1868) MG00
              `--M. unilineatum (Koch 1838) [=Chromatoiulus unilineatus] TKM93

*Type species of generic name indicated

REFERENCES

[A01] Attems, C. Graf. 1901. Myriopodák [Myriopoden]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 275–310. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[DAF07] Department of Agriculture and Food. 2007. Common Seasonal Pests. State of Western Australia.

[H00] Hannibal, J. T. 2000. Hexecontasoma, a new helminthomorph millipede (Hexecontasomatidae n. fam.) from the Mazon Creek, Illinois, fauna (Carboniferous, North America). Fragmenta Faunistica 43 (Suppl.): 19–35.

[H90] Hoffman, R. L. 1990. Diplopoda. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 835–860. John Wiley & Sones: New York.

[KAA-W00] Kheirallah, A.-M., A.-N. H. Aly & N. Y. Abdel-Wahed. 2000. Anamorphosis and life-history of the millipede Nopoiulus kochii (Gervais, 1847), new for Egypt. Zoology in the Middle East 21: 159–168.

[MG00] Mikhaljova, E. V., & S. I. Golovatch. 2000. A review of the millipede fauna of Siberia (Diplopoda). Arthropoda Selecta 9: 103–118.

[RB93] Ross, A. J., & D. E. G. Briggs. 1993. Arthropoda (Euthycarcinoidea and Myriapoda). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 357–361. Chapman & Hall: London.

[TKM93] Thaler, K., B. Knoflach & E. Meyer. 1993. Fragmenta faunistica Tirolensia—X. Berichte des Naturwissenschaftlich-Medizinischen Vereins in Innsbruck 80: 311–325.

[WA04] Wilson, H. M., & L. I. Anderson. 2004. Morphology and taxonomy of Paleozoic millipedes (Diplopoda: Chilognatha: Archipolypoda) from Scotland. Journal of Paleontology 78 (1): 169–184.

Last updated: 10 June 2022.

Xystodesmoidea

Yellow flat-backed millipede Cherokia georgiana, copyright Sheryl Pollock.

Belongs within: Polydesmida.

The Xystodesmoidea are a group of millipedes with usually prominent paraterga and bodies incapable of rolling into a ball.

Characters (from Enghoff et al. 2015): Length ca. 8 to more than 80 mm, exceptionally smaller. Body not capable of volvation, broad and robust, width/length ratio usually <18%, usually parallel-sided on rings 3–16. Antennae rarely with more than four apical cones, not arranged in dyads. Paraterga mostly prominent, lateral calluses normally evident; metaterga usually without setae. Gonopod coxa with cannula; telopodite with neither acccssory seminal chamber nor hairy pulvillus.

<==Xystodesmoidea H90
    |--Eurymerodesmidae H90
    |    |--Eurymerodesmus H90
    |    `--Paresmus H90
    `--Xystodesmidae [Eurydesmidae, Fontariidae] H90
         |  i. s.: Xystocheir H90
         |         Amplocheir H90
         |         Paimokia H90
         |         Anombrocheir H90
         |         Wamokia H90
         |         Motyxia H90
         |         Harpaphe H90
         |         Hybaphe H90
         |         Isaphe H90
         |         Tubaphe H90
         |         Chonaphe H90
         |         Montaphe H90
         |         Metaxycheir H90
         |         Orophe H90
         |         Stenodesmus H90
         |         Gyalostethus H90
         |         Caralinda H90
         |         Pachydesmus H90
         |         Dicellarius H90
         |         Nannaria H90
         |         Oenomoea H90
         |         Apheloria H90
         |         Rudiloria H90
         |         Dixioria H90
         |         Sigmoria aberrans H90, I92
         |         Cleptoria H90
         |         Brachoria H90
         |         Dynoria H90
         |         Lyrranea H90
         |         Deltotaria H90
         |         Stelgipus H90
         |         Cheiropus H90
         |         Croatania H90
         |         Semionellus H90
         `--Rhysodesmini H62
              |--Rhysodesmus Cook 1895 H62
              |    `--R. violaceus (Brölemann 1900) [=Fontaria violacea] K54
              |--Erdelyia Hoffman 1962 H62
              |    `--*E. saucra Hoffman 1962 H62
              |--Cruzodesmus Chamberlin 1943 H62
              |--Acentronus Chamberlin 1943 H62
              |--Boraria Chamberlin 1943 H62
              |--Howellaria Hoffman 1950 H62
              |--Cherokia Chamberlin 1949 H62
              |    `--C. georgiana MG06
              `--Pleuroloma Rafinesque 1820 H62

*Type species of generic name indicated

REFERENCES

Enghoff, H., S. Golovatch, M. Short, P. Stoev & T. Wesener. 2015. Diplopoda—taxonomic overview. In: Minelli, A. (ed.) Treatise on Zoology—Anatomy, Taxonomy, Biology. The Myriapoda vol. 2 pp. 363–454. Brill: Leiden.

[H62] Hoffman, R. L. 1962. A new genus and species in the diplopod family Xystodesmidae (Polydesmida). Proceedings of the Biological Society of Washington 75: 181–188.

[H90] Hoffman, R. L. 1990. Diplopoda. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 835–860. John Wiley & Sones: New York.

[I92] Imes, R. 1992. The Practical Entomologist. Aurum Press: London.

[K54] Kraus, O. 1954. Myriapoden aus El Salvador. Senckenbergiana Biologica 35 (5–6): 293–349.

[MG06] Mallatt, J., & G. Giribet. 2006. Further use of nearly complete 28S and 18S rRNA genes to classify Ecdysozoa: 37 more arthropods and a kinorhynch. Molecular Phylogenetics and Evolution 40: 772–794.

Myriapoda

House centipede Scutigera coleoptrata, copyright Bruce Marlin.

Belongs within: Euarthropoda.
Contains: Diplopoda, Symphyla, Pauropoda, Scolopendromorpha, Geophilomorpha, Lithobiidae.

The Myriapoda are a clade of terrestrial arthropods with often elongate bodies composed of numerous, mostly undifferentiated segments. Respiration is by means of tracheae, and Malpighian tubules are present (Grimaldi & Engel 2005). They include the centipedes (Chilopoda), members of which have the first trunk segment modified into forcipules, with the appendages forming a pincer-like structure with poison glands opening near the tips (Mundel 1990). The remaining myriapods are united in a clade Progoneata supported the presence of gonopores near the front of the body, behind the second body segment.

<==Myriapoda
    |  i. s.: Thyropygus NS03
    |         Atractosoma hyalops Latzel 1889 L89
    |         Arthronomalus E12
    |           |--A. brevicornis Newport 1849 E12
    |           |--A. marginatus Newport 1849 E12
    |           |--A. punctatus Newport 1849 E12
    |           `--A. puncticeps Newport 1849 E12
    |         Antrokoreana gracilipes WS10
    |--Progoneata [Dignatha] RS10
    |    |--Diplopoda AC17
    |    `--+--Arthropleuridea W05
    |       |    |--Arthropleura AC17 [Arthropleurida W05]
    |       |    |--Microdecemplicida W05
    |       |    `--Eoarthropleurida W05
    |       `--+--Symphyla AC17
    |          `--Pauropoda AC17
    `--Chilopoda [Anamorpha, Syngnatha] RS10
         |  i. s.: Eucratonyx hamatus A27
         |         Cupipes impressus A27
         |           |--C. i. impressus A27
         |           `--C. i. armatus A27
         |         Alipes grandidieri SK14
         |--Scutigeromorpha [Notostigmophora] BFM03
         |    |--Pselliodes K55 [Pselliodidae M90, Pselliodinae]
         |    |    `--P. nigrovittata (Meinert 1886) (see below for synonymy) K55
         |    `--Scutigeridae M90
         |         |--Allothereua GEW01
         |         |--Thereuopoda clunifera (Wood 1862) H04
         |         |--Scutigera M90
         |         |    |--S. coleoptrata M90
         |         |    |--S. complanata A01
         |         |    |--S. hispida A01
         |         |    `--S. linceci (Wood 1867) [=Cermatia linceci; incl. S. nubila Chamberlin 1922] K54
         |         `--Pilbarascutigera incola (Verhoeff 1925) T13
         `--Pleurostigmophora RS10
              |--+--Craterostigmus RS10 [Craterostigmomorpha BFM03]
              |  |    |--C. crabilli Edgecombe & Giribet 2008 GFB14
              |  |    `--C. tasmanianus RS10
              |  `--+--Devonobiomorpha BFM03
              |     `--Epimorpha M90
              |          |--Scolopendromorpha BFM03
              |          `--Geophilomorpha BFM03
              `--Lithobiomorpha BFM03
                   |--Lithobiidae BFM03
                   |--Anopsobius GFB14 [Anopsobiidae M90]
                   |    `--A. neozelanicus Silvestri 1909 GFB14
                   |--Gosibiidae M90
                   |--Bothropolys [Ethropolidae] M90
                   |--Pseudolithobius [Pseudolithobiidae] M90
                   |--Henicopidae M90
                   |    |--Zygethobius M90
                   |    `--Lamyctes M90
                   |         |--L. emarginatus M83
                   |         `--L. fulvicornis M90
                   `--Watobiidae M90
                        |--Malbius Chamberlin 1943 C62
                        |--Tropobius C62
                        `--Cruzobius C62
                             |--C. atoyacus Chamberlin 1942 C62
                             |--C. pococki Crabill 1962 C62
                             |--C. verus Chamberlin 1942 C62
                             `--C. viganus Chamberlin 1944 C62

Pselliodes nigrovittata (Meinert 1886) [=Scutigera nigrovittata; incl. P. colombiana Chamberlin 1921, Brasilophora trimarmorata Bücherl 1950] K55

*Type species of generic name indicated

REFERENCES

[AC17] Aria, C., & J.-B. Caron. 2017. Burgess Shale fossils illustrate the origin of the mandibulate body plan. Nature 545: 89–92.

[A01] Attems, C. 1901. Myriopodák [Myriopoden]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 275–310. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[A27] Attems, C. 1927. Myriopoden von Ambon und anderen Inseln der Banda-See. Zoologische Mededeelingen 10: 61–70.

[BFM03] Bonato, L., D. Foddai & A. Minelli. 2003. Evolutionary trends and patterns in centipede segment number based on a cladistic analysis of Mecistocephalidae (Chilopoda: Geophilomorpha). Systematic Entomology 28: 539–579.

[C62] Crabill, R. E., Jr. 1962. A new Cruzobius from Mexico (Chilopoda: Lithobiomorpha: Watobiidae). Proceedings of the Biological Society of Washington 75: 133–136.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l'Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[GEW01] Giribet, G., G. D. Edgecombe & W. C. Wheeler. 2001. Arthropod phylogeny based on eight molecular loci and morphology. Nature 413: 157–161.

[GFB14] Giribet, G., R. Fernández & S. L. Boyer. 2014. On four poorly known harvestmen from New Zealand (Arachnida: Opiliones: Cyphophthalmi: Eupnoi: Dyspnoi: Laniatores). New Zealand Journal of Zoology 41 (4): 223–233.

Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H04] Haupt, J. 2004. A new species of whipscorpion from Laos (Arachnida: Uropygi: Thelyphonidae). Senckenbergiana Biologica 83 (2): 151–155.

[K54] Kraus, O. 1954. Myriapoden aus El Salvador. Senckenbergiana Biologica 35 (5–6): 293–349.

[K55] Kraus, O. 1955. Myriapoden aus Peru, III. Senckenbergiana Biologica 36 (3–4): 173–200.

[L89] Latzel, R. 1889. Sopra alenni miriapodi cavernicoli italiani, raccolti dai Sign. A. Vacca e R. Barberi. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 360–362.

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: an annotated species list. DSIR Entomology Division Report 3: 1–98.

[M90] Mundel, P. 1990. Chilopoda. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 819–833. John Wiley & Sones: New York.

[NS03] Nardi, F., G. Spinsanti, J. L. Boore, A. Carapelli, R. Dallai & F. Frati. 2003. Hexapod origins: monophyletic or paraphyletic? Science 299: 1887–1889.

[RS10] Regier, J. C., J. W. Shultz, A. Zwick, A. Hussey, B. Ball, R. Wetzer, J. W. Martin & C. W. Cunningham. 2010. Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences. Nature 463: 1079–1083.

[SK14] Sharma, P. P., S. T. Kaluziak, A. R. Pérez-Porro, V. L. González, G. Hormiga, W. C. Wheeler & G. Giribet. 2014. Phylogenomic interrogation of Arachnida reveals systemic conflicts in phylogenetic signal. Molecular Biology and Evolution 31 (11): 2963–2984.

[T13] Taylor, C. K. 2013. Annotated bibliography for Barrow Island terrestrial invertebrates. Records of the Western Australian Museum Supplement 83: 135–144.

[WS10] Wang, M., S. Sun, C. Li & X. Shen. 2010. Distinctive mitochondrial genome of the calanoid copepod Calanus sinicus, a useful molecular marker for phylogenetic and population studies. In: China-Russia Bilateral Symposium: Proceedings of the China-Russia Bilateral Symposium of "Comparison on Marine Biodiversity in the Northwest Pacific Ocean", 10–11 October 2010, Qingdao (China) pp. 187–192. Institute of Oceanology, Chinese Academy of Sciences; A. V. Zhirmunsky Institute of Marine Biology, Far East Branch of the Russian Academy of Sciences.

[W05] Wilson, H. M. 2005. Zosterogrammida, a new order of millipedes from the Middle Silurian of Scotland and the Upper Carboniferous of Euramerica. Palaeontology 48 (5): 1101–1110.

Last updated: 5 November 2017.

Vanhoeffeniidae

Anterior of body, together with ventral view of gonopods, of Salvadoria alata, from Kraus (1954b).

Belongs within: Diplopoda.

The Vanhoeffeniidae has been used as a repository for a number of millipedes of uncertain affinities.

Characters (from Brues et al. 1954): Head not covered by first tergite; second tergite not enlarged; male with first pair of legs only on seventh somite modified into gonopods, coxae of gonopods widely separated but connected by broad, medially coalescent processes with median part raised and keel-shaped, gonopod without brush of hairs or seminal pouch.

<==Vanhoeffeniidae
    |--Phaneromerium K54a
    |--Cryptogonodesmus K54a
    |    |--C. fuhrmanni K54a
    |    `--C. peruvianus Kraus 1954 K54a
    |--Fuhrmannodesmus K55
    |    |--F. carli Kraus 1955 K55
    |    `--F. lividus Carl 1914 K55
    |--Olmodesmus Kraus 1954 K54a
    |    |--*O. laticeps Kraus 1954 K54a
    |    |--O. longipes Kraus 1954 K54a
    |    |--O. minimus Kraus 1954 K55
    |    |--O. robustus Kraus 1955 K55
    |    `--O. taulisensis Kraus 1954 K55
    `--Salvadoria Kraus 1954 K54b
         |--*S. alata Kraus 1954 K54b
         |    |--S. a. alata K54b
         |    `--S. a. propinqua Kraus 1954 K54b
         |--S. argentea Kraus 1954 K54b
         |--S. furcata Kraus 1954 K54b
         `--S. sagittalis Kraus 1954 K54b

*Type species of generic name indicated

REFERENCES

Brues, C. T., A. L. Melander & F. M. Carpenter. 1954. Classification of insects: keys to the living and extinct families of insects, and to the living families of other terrestrial arthropods. Bulletin of the Museum of Comparative Zoology at Harvard College 108.

[K54a] Kraus, O. 1954a. Myriapoden aus Peru, II. Senckenbergiana Biologica 35: 17–55.

[K54b] Kraus, O. 1954b. Myriapoden aus El Salvador. Senckenbergiana Biologica 35 (5–6): 293–349.

[K55] Kraus, O. 1955. Myriapoden aus Peru, III. Senckenbergiana Biologica 36 (3–4): 173–200.

Leptodesmidae

Medial (upper left) and lateral views of gonopods of Leptodesmus ornatus, together with diagram of dorsal rim of twelth trunk segment, from Kraus (1954).

Belongs within: Diplopoda.

The Leptodesmidae is a group of millipedes that is most diverse in the Americas.

Characters (from Brues et al. 1954): Males with first pair of legs on seventh somite only modified into gonopods; coxae of gonopods large, close together, connected by a narrow band, each with a prominent curved horn-shaped process; femur distinctly separated from tibia by a constriction, with a large process that is often longer than tibia and tarsus together.

<==Leptodesmidae
    |--Chondrodesmoides Kraus 1955 K55
    |    `--*C. koepckei Kraus 1955 K55
    |--Proletus K55
    |    |--P. orites (Chamberlin 1941) [=*Eucampesmus orites] K55
    |    `--P. tocachensis Kraus 1955 K55
    |--Alocodesmus Silvestri 1896 K54
    |    |--A. aporus Kraus 1954 K54
    |    |--A. dentatus K54
    |    |--A. olivaceus Kraus 1954 K54
    |    |--A. planus K54
    |    `--A. pseudolivaceus Kraus 1954 K54
    `--Leptodesmus Saussure 1859 K54
         |  i. s.: L. gounellei K54
         |         L. soleadinus K54
         |--L. (Leptodesmus) taulisensis Kraus 1954 K54
         |--L. (Desmoleptus) K54
         |    |--L. (*D.) peruanus K54
         |    |--L. (D.) ornatus Kraus 1954 K54
         |    `--L. (D.) similis Kraus 1954 K54
         |--L. (Perudesmus) K54
         |    |--L. (P.) nodosus Kraus 1954 K54
         |    `--L. (P.) titschacki K54
         `--L. (Pseudoleptodesmus) K54
              |--L. (P.) andinus Kraus 1954 K54
              `--L. (P.) bivittatus Kraus 1954 K54

*Type species of generic name indicated

REFERENCES

Brues, C. T., A. L. Melander & F. M. Carpenter. 1954. Classification of insects: keys to the living and extinct families of insects, and to the living families of other terrestrial arthropods. Bulletin of the Museum of Comparative Zoology at Harvard College 108.

[K54] Kraus, O. 1954. Myriapoden aus Peru, II. Senckenbergiana Biologica 35: 17–55.

[K55] Kraus, O. 1955. Myriapoden aus Peru, III. Senckenbergiana Biologica 36 (3–4): 173–200.

Rhachodesmidae

Medial (left) and lateral views of gonopods of Neoleptodesmus geniculatus, from Kraus (1954).

Belongs within: Polydesmida.

The Rhachodesmidae are a family of diverse and often brightly-coloured millipedes found in Mexico and Central America.

See also: Stunning Central American millipedes.

Characters (from Loomis 1964): Body incapable of being tightly rolled; segments 20; segment 1 narrower than head; pores opening from thickened outer rim of keels; gonopods with definite elongate or oval cavity in mesal face containing many setae; last segment conical at apex.

<==Rhachodesmidae
    |--Aceratophallus dux Chamberlin 1914 K54
    |--Pararchistes Pocock 1909 K54
    |--Elcarmenia Kraus 1954 K54
    |    `--*E. engelhardi Kraus 1954 K54
    |--Pammicrophallus Pocock 1909 K54
    |    |--P. ornatus K54
    |    `--P. pasaquinae Kraus 1954 K54
    |--Neoleptodesmus Carl 1903 K54
    |    |--N. aztecus K54
    |    |--N. elevatus K54
    |    |--N. geniculatus Kraus 1954 K54
    |    |--N. guatemalensis K54
    |    |--N. orizabae K54
    |    |--N. salvadorensis Kraus 1954 K54
    |    |--N. sinuatus Kraus 1954 K54
    |    |--N. sumichrasti K54
    |    |--N. vermiformis K54
    |    |--N. vertebratus K54
    |    `--N. vulcani Kraus 1954 K54
    `--Holistophallus LH62

*Type species of generic name indicated

REFERENCES

[K54] Kraus, O. 1954. Myriapoden aus El Salvador. Senckenbergiana Biologica 35 (5–6): 293–349.

Loomis, H. F. 1964. The millipeds of Panama (Diplopoda). Fieldiana: Zoology 47 (1): 1-136.

[LH62] Loomis, H. F. & R. L. Hoffman. 1962. A remarkable new family of spined polydesmoid Diplopoda, including a species lacking gonopods in the male sex. Proceedings of the Biological Society of Washington 75: 145–158.

Diplopoda

Pincushion millipede Phryssonotus sp., copyright Steve & Alison.

Belongs within: Myriapoda.
Contains: Leptodesmidae, Vanhoeffeniidae, Polyxenoidea, Helminthomorpha, Pentazonia.

The Diplopoda, millipedes, are a diverse group of mostly detritivorous terrestrial arthropods with undifferentiated body segments mostly bearing two pairs of legs each. The Polyxenida, pincushion millipedes, are small soft-bodied animals with an extensive covering of tufts of barbed bristles. The remaining millipedes, in the clade Chilognatha, are characterised by a mineralised cuticle.

Characters (from Hoffman 1990): Antennae simple, eight-segmented; whole-limb mandibles present; embryonic maxillary appendages coalesced into gnathochilarium; trunk segments (diplosomites) undifferentiated, derived from fusion of two embryonic somites, generally bearing two pairs of legs; segmentation teloblastic, with proliferation zone located just anterior to last segment; trachea diffusely branched, anastomosing, opening through stigmata at lateral ends of sterna.

<==Diplopoda
    |--Polyxenida [Ancyrotricha, Polyxenida, Polyxenoidea, Pselaphognatha, Schizocephala] W05
    |    |--Polyxenoidea CND-J08
    |    `--Synxenidae [Synxenoidea] CND-J08
    |         |--Phryssonotus Scudder 1885 (see below for synonymy) CS13
    |         |    |--*P. hystrix (Menge 1854) [=*Lophonotus hystrix] CS13
    |         |    |--P. capensis (Silvestri 1923) CND-J08
    |         |    |--P. chilensis (Silvestri 1948) CND-J08
    |         |    |--P. cubanus (Silvestri 1948) CND-J08
    |         |    |--P. novaehollandiae (Silvestri 1923) [=Synxenus novaehollandiae] CS13
    |         |    |--P. orientalis (Silvestri 1900) [=Synxenus orientalis] CND-J08
    |         |    `--P. platycephalus (Lucas 1846) [=Polyxenus platycephalus] CND-J08
    |         `--Condexenus Nguyen Duy-Jacquemin 2006 CND-J08
    `--Chilognatha W05
         |--Helminthomorpha W05
         `--+--Pentazonia W05
            `--Zosterogrammidae [Zosterogrammida] W05
                 |--Zosterogrammus Wilson 2005 W05
                 |    `--*Z. stichostethus Wilson 2005 W05
                 |--Casiogrammus Wilson 2005 W05
                 |    `--*C. ichthyeros Wilson 2005 W05
                 `--Purkynia Fritsch 1899 W05
                      `--*P. lata Fritsch 1899 W05

Diplopoda incertae sedis:
  Schopfia Janssen 1940 LH02
    `--S. calceola LH02
  Leptodesmidae K55
  Vanhoeffeniidae K55
  Oniscodesmidae K55
    |--*Oncodesmus granosus (Gervais & Goudot 1844) K54a
    |--Cyrtodesmus tomentosus Silvestri 1898 K55
    |--Cyliocyrtus Cook 1898 [Oniscodesminae] K55
    |    `--C. ramosus Kraus 1955 K55
    `--Oncodesmoides Kraus 1954 K54a
         |--*O. rectus Kraus 1954 K54a
         `--O. uncinatus Kraus 1954 K54a
  Cryptodesmidae K54b
    |--Tidopterus K54b
    |--Corypherepsis K54b
    `--Apomus (Parapomus Kraus 1954) K54b
         `--A. (*P.) zilchi Kraus 1954 K54b
  ‘Curimagua’ Hoffmann 1982 non Forster & Platnick 1977 KA-Z11
  Brachydesmus superus KAA-W00
  Archispirostreptus gigas SK14

Phryssonotus Scudder 1885 [=Lophonotus Menge 1854 non Stephens 1829; incl. Koubanus Attems 1928, Kubanus Attens 1926, Schindalmonotus Attems 1926, Synxenus Silvestri 1900] CS13

*Type species of generic name indicated

REFERENCES

[CS13] Car, C. A., M. Short, C. Huynh & M. S. Harvey. 2013. The millipedes of Barrow Island, Western Australia (Diplopoda). Records of the Western Australian Museum Supplement 83: 209–219.

[CND-J08] Condé, B. & M. Nguyen Duy-Jacquemin. 2008. Classification actuelle des diplopodes pénicillates (myriapodes) avec nouvelles définitions des taxa. Bulletin de la Société Zoologique de France 133 (4): 291–302.

Hoffman, R. L. 1990. Diplopoda. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 835–860. John Wiley & Sones: New York.

[KAA-W00] Kheirallah, A.-M., A.-N. H. Aly & N. Y. Abdel-Wahed. 2000. Anamorphosis and life-history of the millipede Nopoiulus kochii (Gervais, 1847), new for Egypt. Zoology in the Middle East 21: 159–168.

[K54a] Kraus, O. 1954a. Myriapoden aus Peru, II. Senckenbergiana Biologica 35: 17–55.

[K54b] Kraus, O. 1954b. Myriapoden aus El Salvador. Senckenbergiana Biologica 35 (5–6): 293–349.

[K55] Kraus, O. 1955. Myriapoden aus Peru, III. Senckenbergiana Biologica 36 (3–4): 173–200.

[KA-Z11] Kury, A. B., & M. A. Alonso-Zarazaga. 2011. Addenda and corrigenda to the "Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones)". Zootaxa 3034: 47–68.

[LH02] Lemay, S., & J. T. Hannibal. 2002. Trigonocarpus excrescens Janssen, 1940, a supposed seed from the Pennsylvanian of Illinois, is a milliped (Diplopoda: Euphoberiidae). Kirtlandia 53: 37–40.

[SK14] Sharma, P. P., S. T. Kaluziak, A. R. Pérez-Porro, V. L. González, G. Hormiga, W. C. Wheeler & G. Giribet. 2014. Phylogenomic interrogation of Arachnida reveals systemic conflicts in phylogenetic signal. Molecular Biology and Evolution 31 (11): 2963–2984.

[W05] Wilson, H. M. 2005. Zosterogrammida, a new order of millipedes from the Middle Silurian of Scotland and the Upper Carboniferous of Euramerica. Palaeontology 48 (5): 1101–1110.

Last updated: 5 November 2017.

Helminthomorpha

Reconstruction of Acantherpestes major from Scudder (1882).

Belongs within: Diplopoda.
Contains: Colobognatha, Eugnatha.

The Helminthomorpha is a clade of millipedes ancestrally characterised by the presence of more than 21 trunk segments in adults, though this has been subsequently reduced in some subclades. Crown-group helminthomorphs are also distinguished by the modification in males of at least one pair of legs on the seventh trunk segment into intromittent gonopods (Wilson 2005b).

<==Helminthomorpha [Proterandria] W05b
    |  i. s.: Siphoniulus Pocock 1894 [Siphoniulida, Siphoniulidae, Siphoniuloidea] SS03
    |           |--*S. albus Pocock 1894 SS03
    |           `--S. neotropicus Hoffman 1979 SS03
    |         Hexecontasoma Hannibal 2000 [Hexecontasomatidae] H00
    |           `--*H. carinatum Hannibal 2000 H00
    |--+--Pleurojulus Fritsch 1899 [Pleurojulida] W05b
    |  `--+--Colobognatha WA04
    |     `--Eugnatha WA04
    `--Archipolypoda [Macrosterni, Palaeocoxopleura] WA04
         |  i. s.: Anaxeodesmus Wilson 2005 W05a
         |           `--*A. diambonotus Wilson 2005 W05a
         |         Palaeodesmus Wilson & Anderson 2004 WA04
         |           `--*P. tuberculata (Brade-Birks 1923) [=Kampecaris tuberculata] WA04
         |         Albadesmus Wilson & Anderson 2004 WA04
         |           `--*A. almondi Wilson & Anderson 2004 WA04
         |         Pneumodesmus Wilson & Anderson 2004 WA04
         |           `--*P. newmani Wilson & Anderson 2004 WA04
         |         Archiulus [Archiulidae] WA04
         |         Anthracodesmus Peach 1899 WA04
         |           `--*A. macconochiei Peach 1899 WA04
         |         Pattonia WA04
         |--+--Archidesmus Peach 1882 [Archidesmida, Archidesmidae] WA04
         |  |    `--*A. macnicoli Peach 1882 WA04
         |  `--Cowiedesmus Wilson & Anderson 2004 [Cowiedesmida, Cowiedesmidae] WA04
         |       `--C. eroticopodus Wilson & Anderson 2004 WA04
         `--Euphoberiidae [Euphoberiida] WA04
              |--Euphoberia Meek & Worthen 1868 WA04
              |    `--E. horrida WA04
              |--Acantherpestes Meek & Worthen 1868 WA04
              |    |--A. major K-P83
              |    `--A. vicinus K-P83
              `--Myriacantherpestes Burke 1979 WA04
                   |--M. bradebirski Burke 1979 LH02
                   |--M. clarkorum (Burke 1973) LH02
                   |--M. excrescens (Janssen 1940) [=Trigonocarpus excrescens] LH02
                   |--M. ferox (Salter 1863) LH02
                   |--M. hystricosus (Scudder 1890) LH02
                   `--M. inequalis (Scudder 1890) LH02

*Type species of generic name indicated

REFERENCES

[H00] Hannibal, J. T. 2000. Hexecontasoma, a new helminthomorph millipede (Hexecontasomatidae n. fam.) from the Mazon Creek, Illinois, fauna (Carboniferous, North America). Fragmenta Faunistica 43 (Suppl): 19–35.

[K-P83] Kukalová-Peck, J. 1983. Origin of the insect wing and wing articulation from the arthropodan leg. Canadian Journal of Zoology 61: 1618–1669.

[LH02] Lemay, S., & J. T. Hannibal. 2002. Trigonocarpus excrescens Janssen, 1940, a supposed seed from the Pennsylvanian of Illinois, is a milliped (Diplopoda: Euphoberiidae). Kirtlandia 53: 37–40.

[SS03] Sierwald, P., W. A. Shear, R. M. Shelley & J. E. Bond. 2003. Millipede phylogeny revisited in the light of the enigmatic order Siphoniulida. Journal of Zoological Systematics and Evolutionary Research 41: 87–99.

[WA04] Wilson, H. M., & L. I. Anderson. 2004. Morphology and taxonomy of Paleozoic millipedes (Diplopoda: Chilognatha: Archipolypoda) from Scotland. Journal of Paleontology 78 (1): 169–184.

[W05a] Wilson, H. M. 2005a. A new genus of archipolypodan millipede from the Coseley Lagerstätte, Upper Carboniferous, UK. Palaeontology 48 (5): 1097–1100.

[W05b] Wilson, H. M. 2005b. Zosterogrammida, a new order of millipedes from the Middle Silurian of Scotland and the Upper Carboniferous of Euramerica. Palaeontology 48 (5): 1101–1110.

Platyrhacidae

Possible Amplinus, copyright P. Wittman.

Belongs within: Polydesmida.

The Platyrhacidae are a family of often large and colourful millipedes found in southeast Asia and Central and South America.

Characters (from Hoffman 1990): Collum wider than head, of which it covers only the posteriormost portion, and extended ventrolaterad behind mandibles; metaterga smooth and polished, without trace of sulcus; ozopores opening directly on paranotal surface; epiproct apically broadened and truncate.

<==Platyrhacidae [Euryuridae, Platyrhacoidea]
    |--Euryurus maculatus H90
    |--Auturus H90
    |--Amplinus Attems 1901 K54
    |    |--A. klugii K54
    |    |--A. palicaudatus (Attems 1901) [=Pachyurus palicaudatus] K54
    |    `--A. serratus Kraus 1954 K54
    `--Platyrhacus (Tirodesmus Cook 1896) [incl. Aymaresmus Chamberlin 1941] K55
         |--P. (T.) celinus K55
         |--P. (T.) obscurus Kraus 1955 K55
         |--P. (T.) orellanus K55
         `--P. (T.) tampocus K55

*Type species of generic name indicated

REFERENCES

Hoffman, R. L. 1990. Diplopoda. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 835–860. John Wiley & Sones: New York.

Kraus, O. 1954. Myriapoden aus El Salvador. Senckenbergiana Biologica 35 (5–6): 293–349.

Kraus, O. 1955. Myriapoden aus Peru, III. Senckenbergiana Biologica 36 (3–4): 173–200.

Colobognatha

Head of Siphonophora barberi, from Chamberlin (1922).

Belongs within: Helminthomorpha.

The Colobognatha are a clade of millipedes characterised by the presence of elongate, sucking mouthparts. They are divided between three orders, among which the Platydesmida have the pleurites fused to the tergites (Sierwald et al. 2003) and a well-defined median longitudinal suture along the metaterga (Hoffman 1990). The Polyzoniida and Siphonophorida can be distinguished by the presence (Polyzoniida) or absence (Siphonophorida) of ocelli.

Synapomorphies (from Sierwald et al. 2003): Mouthparts elongate, with gnathochilarial palps absent; adult males with ninth leg modified into an accessory gonopod, tenth leg into a functional gonopod; adults curl protectively around eggs; first larval instar with four leg pairs.

<==Colobognatha [Typhlogena]
    |--Platydesmida SS03
    |    |--Platydesmus Lucas 1843 [Platydesmidae] K54b
    |    |    |--P. lankesteri Brölemann 1905 K54b
    |    |    `--P. marmoreus Pocock 1903 K54b
    |    `--Andrognathidae H90
    |         |--Andrognathus H90
    |         |--Ischnocybe H90
    |         |--Mitocybe H90
    |         |--Gosodesmus H90
    |         `--Brachycybe lecontii H90
    `--+--Polyzoniida [Ommatophora, Polyzoniidea] SS03
       |    |  i. s.: Petaserpes Cope 1870 W05
       |    |--Siphonotus [Siphonotidae] H90
       |    |    `--S. purpureus [=Rhinotus purpureus] H90
       |    |--Hirudisomatidae H90
       |    |    |--Euzonium H90
       |    |    |--Hypozonium H90
       |    |    `--Octoglena H90
       |    `--Polyzoniidae H90
       |         |--Bdellozonium H90
       |         |--Buzonium H90
       |         |--Calzonium H90
       |         |--Pizonium H90
       |         |--Polyzonium Brandt 1837 W05
       |         |    `--P. germanicum RS10
       |         `--Angarozonium Shelley 1998 MG00
       |              `--A. amurense (Gerstfeldt 1859) (see below for synonymy) MG00
       `--Siphonophorida SS03
            |--Siphonorhinidae H90
            |    |--Illacme plenipes H90, GE05
            |    `--Siphonorhinus H90
            `--Siphonophoridae H90
                 |--Siphonacme H90
                 `--Siphonophora Newport 1845 K54a
                      |--S. andecola Kraus 1954 K54a
                      |--S. barberi K54b
                      |--S. gracilicornis K54a
                      |--S. lafloridae Kraus 1955 K55
                      |--S. porcullae Kraus 1954 K54a
                      |--S. progressor K54b
                      |--S. sabachana K54b
                      `--S. trifini Kraus 1954 K54b

Angarozonium amurense (Gerstfeldt 1859) [=Platydesmus amurensis, Polyzonium amurense; incl. Po. cyathiferum Mikhaljova 1981] MG00

*Type species of generic name indicated

REFERENCES

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H90] Hoffman, R. L. 1990. Diplopoda. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 835–860. John Wiley & Sones: New York.

[K54a] Kraus, O. 1954a. Myriapoden aus Peru, II. Senckenbergiana Biologica 35: 17–55.

[K54b] Kraus, O. 1954b. Myriapoden aus El Salvador. Senckenbergiana Biologica 35 (5–6): 293–349.

[K55] Kraus, O. 1955. Myriapoden aus Peru, III. Senckenbergiana Biologica 36 (3–4): 173–200.

[MG00] Mikhaljova, E. V., & S. I. Golovatch. 2000. A review of the millipede fauna of Siberia (Diplopoda). Arthropoda Selecta 9: 103–118.

[RS10] Regier, J. C., J. W. Shultz, A. Zwick, A. Hussey, B. Ball, R. Wetzer, J. W. Martin & C. W. Cunningham. 2010. Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences. Nature 463: 1079–1083.

[SS03] Sierwald, P., W. A. Shear, R. M. Shelley & J. E. Bond. 2003. Millipede phylogeny revisited in the light of the enigmatic order Siphoniulida. Journal of Zoological Systematics and Evolutionary Research 41: 87–99.

[W05] Wilson, H. M. 2005. Zosterogrammida, a new order of millipedes from the Middle Silurian of Scotland and the Upper Carboniferous of Euramerica. Palaeontology 48 (5): 1101–1110.

Last updated: 25 May 2017.

Spirostreptidae

Spirostreptus ibanda, copyright Gey-stern.

Belongs within: Eugnatha.

The Spirostreptidae is a family of often large millipedes.

Characters (from Hoffman 1990): Stipes of gnathochilarium widely separated by large central mentum, but lingual lamellae in contact, not separated by mentum or promentum; lateral ends of collum with several parallel engraved striations; sterna, pleura and terga coalesced into rigid cylindrical rings, with pleural sclerites completely coalesced with lower ends of terga, no suture line remaining; segment 3 without legs.

<==Spirostreptidae [Spirostreptidea]
    |  i. s.: Charactopygus frequens H02
    |         Orthoporus C90
    |           |--O. cobanus L62
    |           |--O. flavior C90
    |           |--O. kiemi Loomis 1962 L62
    |           |--O. luchicolens L62
    |           |--O. ornatus C90
    |           `--O. paxillicauda Loomis 1962 L62
    `--Spirostreptinae K55
         |--Spirostreptus ibanda H02
         |--Epistreptus K55
         |    |--E. (Epistreptus) K55
         |    |    |--*E. (E.) oscenus Silvestri 1897 K55
         |    |    `--E. (E.) silvestrii Kraus 1955 K55
         |    `--E. (Microtrullius) vagans (Chamberlin 1922) (see below for synonymy) K54
         `--Scaphiostreptus [incl. Andineptus Chamberlin 1941] K55
              |  i. s.: S. rugiceps K54
              `--S. (Scaphiostreptus) K55
                   |--S. (S.) absconsus (Chamberlin 1922) [=Orthoporus absconsus] K54
                   |--S. (S.) apheles (Chamberlin 1941) [=Andineptus apheles] K55
                   |--S. (S.) broelemanni (Verhoeff 1941) K55
                   |--S. (S.) delectus (Chamberlin 1941) [=Andineptus delectus] K55
                   |--S. (S.) huallagae Kraus 1955 K55
                   |--S. (S.) perditus (Chamberlin 1941) [=Andineptus perditus] K55
                   |--S. (S.) pongus (Chamberlin 1941) [=Andineptus pongus] K55
                   `--S. (S.) salvadoricus Kraus 1954 K54

Epistreptus (Microtrullius) vagans (Chamberlin 1922) [=Gymnostreptus vagans; incl. G. pacificus Chamberlin 1922, Epistreptus (Microtrullius) pacificus] K54

*Type species of generic name indicated

REFERENCES

[C90] Crawford, C. S. 1990. Scorpiones, Solifugae, and associated desert taxa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 421–475. John Wiley & Sones: New York.

Hoffman, R. L. 1990. Diplopoda. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 835-860. John Wiley & Sones: New York.

[H02] Hunt, D. J. 2002. The African Rhigonematoidea (Nematoda: Rhigonematida). 1. Trachyglossus quintus gen. n., sp. n. (Xustrostomatidae fam. n.) with redescription of Rhigonema multipapillatum (Skrjabin, 1916) and designation of a neotype. Nematology 4: 671–686.

[K54] Kraus, O. 1954. Myriapoden aus El Salvador. Senckenbergiana Biologica 35 (5-6): 293–349.

[K55] Kraus, O. 1955. Myriapoden aus Peru, III. Senckenbergiana Biologica 36 (3-4): 173–200.

[L62] Loomis, H. F. 1962. Two unusual Central American spirostreptid milliped species. Proceedings of the Biological Society of Washington 75: 47–52.

Eugnatha

Gathering of Epinannolene, copyright Karl Questel.

Belongs within: Helminthomorpha.
Contains: Chordeumatida, Polydesmida, Spirobolida, Julida, Spirostreptidae.

The Eugnatha are a clade within the millipedes characterised by modification of the eighth pair of legs in adult males into a pair of functional gonopods, and the fusion of the pleurites to the tergites. Major clades within the Eugnatha include the Nematophora, characterised by the possession of spinnerets, and molar cusps on the mandibles (Sierwald et al. 2003).

<==Eugnatha
    |--Nematophora SS03
    |    |--Stemmiulida [Monocheta] WA04
    |    `--Coelocheta SS03
    |         |--Chordeumatida WA04
    |         `--Callipodida WA04
    |              |--Abacion magnum H00
    |              |--Delophon H90
    |              |--Tetracion H90
    |              |--Texophon H90
    |              |--Tynomma H90
    |              |--Diactis H90
    |              |--Heptium H90
    |              |--Colactis H90
    |              `--Apfelbeckia Verhoeff 1896 H00
    |                   `--A. hessei Verhoeff 1929 UO05
    `--+--Polydesmida WA04
       `--Juliformia [Juloidea] SS03
            |--+--Spirobolida SS03
            |  `--Julida SS03
            `--Spirostreptida SS03
                 |--Spirostreptidae H90
                 |--Epinannolenidea H90
                 |    |--Choctella [Choctellidae] H90
                 |    |    `--C. cummingsi H90
                 |    `--Epinannolene K55
                 |         |--E. alticola (Silvestri 1898) K55
                 |         |--E. bifasciata (Silvestri 1898) K54
                 |         |--E. cylidricaulis Kraus 1954 K54
                 |         `--E. flagellosa Kraus 1955 K55
                 `--Cambalidea H00
                      |  i. s.: Protosilvestria Handschin 1944 H00
                      |--Cambalopsidae H90
                      |    |--Plusioglyphiulus SGS04
                      |    `--Cambalopsis nordquisti A27
                      `--Cambalidae [Leioderidae, Nannolenidae] H90
                           |--Cambala hubrichti H00
                           |--Agastrophus B13
                           |--Hypocambala B13
                           |--Nannolene H90
                           |--Paiteya H90
                           |--Titsona H90
                           |    |--*T. sima C62
                           |    `--T. tida Chamberlin 1962 C62
                           |--Buwatia H90
                           |--Platydere H90
                           |--Pharodere H90
                           |--Endere H90
                           |--Leiodere H90
                           |--Odachurus H90
                           |--Alakene H90
                           |--Doilene H90
                           |--Tigolene H90
                           |--Amastigogonus Brölemann 1913 B13
                           |    `--*A. tasmanianus Brölemann 1913 B13
                           `--Dimerogonus Attems 1903 B13
                                `--*D. orophilus B13

*Type species of generic name indicated

REFERENCES

[A27] Attems, C. 1927. Myriopoden von Ambon und anderen Inseln der Banda-See. Zoologische Mededeelingen 10: 61–70.

[B13] Brölemann, H. W. 1913. The Myriapoda in the Australian Museum. Part ii. Diplopoda. Records of the Australian Museum 10 (6): 77–158, pls 14–18.

[C62] Chamberlin, R. V. 1962. Millipeds from the Nevada Test Area. Proceedings of the Biological Society of Washington 75: 53–56.

[H00] Hannibal, J. T. 2000. Hexecontasoma, a new helminthomorph millipede (Hexecontasomatidae n. fam.) from the Mazon Creek, Illinois, fauna (Carboniferous, North America). Fragmenta Faunistica 43 (Suppl): 19–35.

[H90] Hoffman, R. L. 1990. Diplopoda. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 835–860. John Wiley & Sones: New York.

[K54] Kraus, O. 1954. Myriapoden aus Peru, II. Senckenbergiana Biologica 35: 17–55.

[K55] Kraus, O. 1955. Myriapoden aus Peru, III. Senckenbergiana Biologica 36 (3–4): 173–200.

[SGS04] Schwendinger, P. J., G. Giribet & H. Steiner. 2004. A remarkable new cave-dwelling Stylocellus (Opiliones, Cyphophthalmi) from peninsular Malaysia, with a discussion on taxonomic characters in the family Stylocellidae. Journal of Natural History 38: 1421–1435.

[SS03] Sierwald, P., W. A. Shear, R. M. Shelley & J. E. Bond. 2003. Millipede phylogeny revisited in the light of the enigmatic order Siphoniulida. Journal of Zoological Systematics and Evolutionary Research 41: 87–99.

[UO05] Ubick, D., & R. Ozimec. 2005. On the harvestman genus Lola Kratochvíl (Opiliones: Laniatores). Natura Croatica 14 (3): 161–174.

[WA04] Wilson, H. M., & L. I. Anderson. 2004. Morphology and taxonomy of Paleozoic millipedes (Diplopoda: Chilognatha: Archipolypoda) from Scotland. Journal of Paleontology 78 (1): 169–184.

Last updated: 25 May 2017.