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There are 12,000 described millipede species, but taxonomists estimate the number to be ten-fold greater. In the Appalachian Mountains, where I discovered ten new species of Brachoria, the “Appalachian mimic millipedes”, the pace of millipede species discovery is among the most rapid of any organismal group. The lack of taxonomic knowledge gives a false impression about millipedes’ ecological importance. In forest ecosystems, they fulfill an essential ecological service. By feeding on decaying wood, leaves, and other detritus, millipedes help break down dead vegetation thereby releasing essential nutrients which would otherwise be tied up in organic material.


The United States contains a fantastic diversity of millipedes, with a biodiversity hotspot in the Appalachian Mountains. Millipede diversity and abundance is particularly high in the limestone rich mountains and hills, perhaps because these millipedes bioaccumulate calcium carbonate as a component of their exoskeleton. Unfortunately, the limestone hills are also the target for the destructive mining practice of mountain top removal, which threatens the habitat of many narrowly endemic species. An important part of our research is describing biodiversity. The mainstay of this project is the revisionary monograph. A recently completed paper, a revisionary monograph of the mimic millipede genus Brachoria in the Appalachian Mountains, contains descriptive taxonomy and molecular systematics of 34 species, 10 of which are new.




^ Brachoria in the Tree of Life webproject:

Marek, P.E. 2010. Brachoria Chamberlin 1939. Version 16 September 2010. http://tolweb.org/Brachoria in The Tree of Life Web Project, http://tolweb.org





Most biological questions are built upon systematic and taxonomic studies. They are the fundamental building blocks for evolutionary, ecological, conservation and medical studies. For example, before studying color evolution and Müllerian mimicry in millipedes, in order to test whether shared coloration is a result of close ancestry instead of mimicry between unrelated species, I first assembled a systematic framework based on a molecular phylogeny.


Systematics also provides a framework for species discovery and description, and for updating the Linnean classification system with information about evolutionary history.


This is a molecular phylogeny of millipedes in the tribe Apheloriini that I used to name a new genus, Appalachioria; describe 10 new species of Brachoria and address questions about the evolution of warning coloration and Müllerian mimicry in these millipedes.





^ Phylogeny of the millipede tribe Apheloriini reconstructed using Bayesian inference (click to enlarge).


Harmonic mean likelihood ≥ -18315.52. Thickened, black branches denote posterior clade probability values ≥ 0.95. Boxes above branches indicate clades supported by different analyses: “Circle B”, Bayesian inference, posterior clade probability ≥ 0.95; “Circle P”, Direct optimization, node recovered in ≥ 95% of trees generated under the preferred gap-opening and extension model (summarized in a 95% majority rule consensus); and “Circle M”, Maximum likelihood, nonparametric bootstrap clade support values ≥ 95%. Numbers in boxes indicate support values < 95%. Gyalostethus monticolens branch abbreviated with a break for ease of illustration (branch length = 1.047, about 1.6X greater than shown).

 

Taxonomy and systematics

All species in the U.S.

My goal is to collect every described species of xystodesmid millipede in the United States.  Specimens will be photographed, color measured with a spectrometer, and DNA and RNA preserved.  Once DNA/RNA is preserved, specimens will be identified, labeled and preserved as a museum specimen.


Species pages will be created for each on the Tree of Life web project.

U.S. Xystodesmidae project

Checklist of U.S. Xystodesmidae (230 in U.S. - 47% completed)


green, have / red, wanted


Xystodesmidae (2 subfamilies, 1 in U.S. and 1 in Japan)

  1. Xystodesminae (9 tribes, 8 in North America and 1 in the Mediterranean rim, with representatives in eastern Asia and Central America)

  2. - Apheloriini Hoffman, 1979 (17 genera, 116 species: eastern U.S.)


  3. Apheloria Chamberlin, 1921 - 4 species: A. luminosa (Kenyon, 1893); A. montana (Bollman, 1887); A. tigana Chamberlin, 1939; A. virginiensis virginiensis (Drury, 1770); A. virginiensis corrugata (Wood, 1864); A. virginiensis butleriana (Bollman, 1889); A. virginiensis iowa Chamberlin, 1939; A. virginiensis reducta Chamberlin, 1939


  4. Appalachioria Marek and Bond, 2006 - 4 species: A. eutypa eutypa (Chamberlin, 1939); A. eutypa ethotela Chamberlin, 1942); A. falcifera (Keeton, 1959); A. separanda separanda Chamberlin, 1947); A. separanda calcaria (Keeton, 1959); A. separanda hamata (Keeton, 1959); A. separanda versicolor (Hoffman, 1963); A. turneri (Keeton, 1959)


  5. Brachoria Chamberlin, 1939 - 34 species: B. badbranchensis Marek, 2010; B. blackmountainensis Marek, 2010; B. calceata (Causey, 1955); B. campcreekensis Marek, 2010; B. cedra Keeton, 1959; B. conta Keeton, 1965; B. cumberlandmountainensis Marek, 2010; B. dentata Keeton, 1959; B. divicuma Keeton, 1965; B. electa Causey, 1955; B. enodicuma Keeton, 1965; B. evides (Bollman, 1887); B. flammipes Marek, 2010; B. glendalea (Chamberlin, 1918); B. gracilipes (Chamberlin, 1947); B. grapevinensis Marek, 2010; B. guntermountainensis Marek, 2010; B. hansonia Causey, 1950; B. hendrixsoni Marek, 2010; B. hoffmani Keeton, 1959; B. hubrichti Keeton, 1959; B. indianae (Bollman, 1888); B. initialis Chamberlin, 1939; B. insolita Keeton, 1959; B. kentuckiana (Causey, 1942); B. laminata Keeton, 1959; B. ligula Keeton, 1959; B. mendota Keeton, 1959; B. ochra (Chamberlin, 1918); B. plecta Keeton, 1959; B. sheari Marek, 2010; B. splendida (Causey, 1942); B. virginia Marek, 2010; B. viridicolens Hoffman, 1948


  6. Brevigonus Shelley, 1980 - 2 species: B. arcuatus Shelley, 1981; B. shelfordi (Loomis, 1944)


  7. Cheiropus Loomis, 1944 - 1 species: C. plancus Loomis, 1944


  8. Cleptoria Chamberlin, 1939 - 5 species: C. abbotti Hoffman, 1967; C. bipraesidens Hoffman, 1967; C. macra Chamberlin, 1939; C. rileyi (Bollman, 1889); C. robusta Shelley, 1986


  9. Croatania Shelley, 1977 - 4 species: C. catawba Shelley, 1977; C. saluda Shelley, 1977; C. simplex Shelley, 1977; C. yemassee Shelley, 1977


  10. Deltotaria Causey, 1942 - 2 species: D. brimleii brimleii Causey, 1942; D. brimleii philia (Chamberlin, 1949); D. lea Hoffman, 1961


  11. Dixioria Chamberlin, 1947 - 8 species: D. acuminata Hoffman, 1956; D. brooksi Hoffman, 1956; D. dactylifera Hoffman, 1956; D. fowleri Hoffman, 1956; D. coronata Hoffman, 1949; D. pela (Chamberlin, 1918); D. wrighti Hoffman, 1956; D. watauga Shelley, 1986


  12. Dynoria Chamberlin, 1939 - 2 species: D. icana Chamberlin, 1939; D. medialis Chamberlin, 1949


  13. Falloria Hoffman, 1948 - 17 species: F. abbreviata Shelley, 1986; F. ainsliei (Chamberlin, 1921); F. aphelorioides Shelley, 1986; F. bidens (Causey, 1942); F. crassicurvosa Shelley, 1986; F. forficata Shelley, 1986; F. fumimontis (Shelley, 1981); F. houstoni (Chamberlin, 1943); F. leucostriata (Shelley, 1981); F. lyrea (Shelley, 1981); F. mimetica (Chamberlin, 1918); F. pendulata Shelley, 1986; F. picapa (Keeton, 1960); F. prolata Shelley, 1986; F. translineata (Shelley, 1981); F. tuberosa (Shelley, 1981); F. xerophylla (Shelley, 1981)


  14. Furcillaria Shelley, 1981 - 4 species: F. aequalis Shelley, 1981; F. convoluta Shelley, 1981; F. laminata Shelley, 1981; F. thrinax (Shelley, 1982)


  15. Lyrranea Hoffman, 1963 - 1 species: L. persica Hoffman, 1963


  16. Prionogonus Shelley, 1982: 4 species: P. divaricatus Shelley, 1982; P. divergens (Chamberlin, 1939); P. haerens Shelley, 1982; P. stibarophalla (Shelley, 1981).


  17. Rudiloria Causey, 1955 - 5 species: R. guyandotta (Shear, 1972); R. kleinpeteri (Hoffman, 1949); R. mohicana Causey, 1955; R. rigida Shelley, 1986; R. trimaculata tortua (Chamberlin, 1949); R. trimaculata trimaculata (Wood, 1864).


  18. Sigmoria Chamberlin, 1939 - 17 species: S. areolata Shelley, 1981; S. australis Shelley, 1986; S. austrimontis Shelley, 1981; S. disjuncta Shelley, 1981; S. laticurvosa Shelley, 1981; S. latior latior (Brölemann, 1900); S. latior hoffmani Shelley, 1976; S. latior munda Chamberlin, 1939; S. nantahalae Hoffman, 1958; S. nigrimontis nigrimontis (Chamberlin, 1947); S. nigrimontis angulosa Shelley, 1981; S. nigrimontis intermedia (Hoffman, 1948); S. nigrimontis unicoi Shelley, 1981; S. quadrata Shelley, 1981; S. rubromarginata (Bollman, 1888); S. sigirioides Shelley, 1981; S. simplex Shelley, 1981; S. stenogon Chamberlin, 1942; S. stenoloba Shelley, 1981; S. triangulata Shelley, 1981; S. truncata Shelley, 1981; S. whiteheadi Shelley, 1986


  19. Stelgipus Loomis, 1944 - 2 species: S. agrestis Loomis, 1944; S. serratus (Shelley, 1984)

  20. - Chonaphini Verhoeff, 1941 (6 genera, 12 species: western U.S., except Semionellus placidus in eastern U.S.)


  21. Chonaphe Cook, 1904 - 4 species: C. armata (Harger, 1872); C. evexa Shelley, 1994; C. remissa Chamberlin, 1949; C. schizoterminalis Shelley, 1994


  22. Metaxycheir Buckett & Gardner, 1969 - 1 species: M. prolata Buckett & Gardner, 1969


  23. Montaphe Chamberlin, 1949 - 2 species: M. elrodi (Chamberlin, 1913); M. paraphoena Shelley, 1994


  24. Selenocheir Shelley, 1994 - 3 species: S. arcuata Shelley, 1994; S. directa Shelley, 1994; S. sinuata Shelley, 1994


  25. Semionellus Chamberlin, 1920 - 1 species: S. placidus (Wood, 1864)


  26. Tubaphe Causey, 1954 - 1 species: T. levii Causey, 1954

  27. - Nannariini Hoffman, 1964 (2 genera, 26 species: eastern U.S.)


  28. Nannaria Chamberlin, 1918 - 25 species: N. austricola Hoffman, 1950; N. castanea (McNeill, 1887); N. conservata Chamberlin, 1940; N. davidcauseyi Causey, 1950; N. depalmai (Causey, 1950); N. domestica Shelley, 1975; N. equalis Chamberlin, 1949; N. ericacea Hoffman, 1949; N. fowleri Chamberlin, 1947; N. laminata Hoffman, 1949; N. minor Chamberlin, 1918; N. missouriensis Chamberlin, 1928; N. morrisoni Hoffman, 1948; N. oblonga (Koch, 1847); N. ohionis Loomis & Hoffman, 1948; N. rutherfordensis Shelley, 1975; N. scutellaria Causey, 1942; N. shenandoah Hoffman, 1949; N. simplex Hoffman, 1949; N. tennesseensis (Bollman, 1889); N. terricola (Williams & Hefner, 1928); N. wilsoni Hoffman, 1949


  29. Oenomaea Hoffman, 1964 - 1 species: O. pulchella (Bollman, 1889)

  30. - Orophini Hoffman, 1964 (3 genera, 2 species: 1 northwestern U.S., 2 China - Kiulinga, Pamelaphe)


  31. Orophe Chamberlin, 1951 - 2 species: O. cabinetus Chamberlin, 1951; O. unicus Loomis, 1953

  32. - Pachydesmini Hoffman, 1980 (3 genera, 9 species: eastern, central U.S.)


  33. Dicellarius Chamberlin, 1920 - 5 species: D. atlanta (Chamberlin, 1946); D. bimaculatus bimaculatus (McNeill, 1887); D. b. fictus (Chamberlin, 1943); D. b. lamellidens (Chamberlin, 1931); D. okefenokensis (Chamberlin, 1918); Dicellarius sternolobus Loomis, 1969; D. talapoosa talapoosa (Chamberlin, 1939); D. t. separandus Shelley, 1984


  34. Pachydesmus Cook, 1895 - 2 species: P. clarus (Chamberlin, 1918); P. crassicutis crassicutis (Wood, 1864); P. c. denticulatus Chamberlin, 1946; P. c. duplex Chamberlin, 1939; P. c. incursus Chamberlin, 1939; P. c. laticollis (Attems, 1899); P. c. hubrichti Hoffman, 1958; P. c. retrorsus Chamberlin, 1921; P. c. adsinicolus Hoffman, 1958


  35. Thrinaxoria Chamberlin & Hoffman, 1950 - 2 species: T. bifida (Wood, 1864); T. lampra (Chamberlin, 1918)

  36. - Rhysodesmini Brölemann, 1916 (11 genera, 26 species: eastern, central, southwestern U.S. - Rhysodesmus 3 species eastern and southwestern U.S., 67 species Central America; Stenodesmus 1 species southwestern U.S., 4 species Mexico)


  37. Boraria Chamberlin, 1943 - 4 species: B. deturkiana (Causey, 1942); B. infesta (Chamberlin, 1918); B. profuga Causey, 1955; B. stricta (Brölemann, 1896)


  38. Caralinda Hoffman, 1978 - 4 species: C. beatrix Hoffman, 1978; C. causeyae Shelley, 1983; C. dactylifera Shelley, 1983; C. pulchritecta Shelley, 1979


  39. Erdelyia Hoffman, 1962 - 1 species: E. saucra Hoffman, 1962


  40. Cherokia Chamberlin, 1949 - 1 species: C. georgiana georgiana (Bollman, 1889); C. g. ducilla (Chamberlin, 1939); C. g. latassa Hoffman, 1960


  41. Gonoessa Shelley, 1984 - 5 species: G. aciculata Shelley, 1984; G. cingulata Shelley, 1984; G. clavata Shelley, 1984; G. dentata Shelley, 1984; G. furcata Shelley, 1984


  42. Gyalostethus Hoffman, 1965 - 1 species: G. monticolens (Chamberlin, 1951)


  43. Lourdesia Shelley, 1991 - 1 species: L. minuscula Shelley, 1991


  44. Parvulodesmus Shelley, 1983 - 1 species: P. prolixogonus Shelley, 1983


  45. Pleuroloma Rafinesque, 1820 - 4 species: P. cala (Chamberlin, 1939); P. flavipes Rafinesque, 1820; P. pinicola Shelley, 1980; P. plana Shelley, 1980


  46. Rhysodesmus Cook, 1895 - 3 species: R. chisosi Shelley, 1989; R. restans Hoffman, 1998; R. texicolens (Chamberlin, 1938)


  47. Stenodesmus De Saussure, 1859 - 1 species: S. tuobitus (Chamberlin, 1910)

  48. - Sigmocheirini Causey, 1955 (2 genera, 4 species: California)


  49. Ochthocelata Shelley, 1995 - 1 species: O. adynata Shelley, 1995


  50. Sigmocheir Chamberlin, 1951 - 3 species: S. calaveras Chamberlin, 1951; S. furcata Shelley, 1995; S. maculifer (Chamberlin, 1941)

  51. - Xystodesmini Hoffman, 1980 (7 genera, 5 species: western U.S., eastern Asia - Koreoaria, Levizonus, Riukiaria, Xystodesmus, Yaetakaria)


  52. Harpaphe Cook, 1904 - 3 species: H. haydeniana haydeniana (Wood, 1864); H. h. cummingsiensis (Verhoeff, 1944); H. h. inlignea Chamberlin, 1949; H. h. maurogona Buckett & Gardner, 1968; H. h. lanceolata Buckett & Gardner, 1968; H. h. scotia (Chamberlin, 1941); H. pottera Chamberlin, 1949; H. telodonta (Chamberlin, 1943)


  53. Isaphe Cook, 1904 - 2 species: I. convexa Cook, 1904; I. tersa (Cook, 1904)

  54. - Xystocheirini Hoffman, 1980 (5 genera, 30 species: California)


  55. Anombrocheir Buckett & Gardner, 1969 - 2 species: A. bifurcata Gardner & Buckett, 1969; A. spinosa Buckett & Gardner, 1969


  56. Motyxia Chamberlin, 1941 - 8 species: M. kerna Chamberlin, 1941; M. monica Chamberlin, 1944; M. pior Chamberlin, 1941; M. porrecta Causey & Tiemann, 1969; M. sequoia sequoia (Chamberlin, 1941); M. sequoia alia Causey & Tiemann, 1969; M. sequoiae (Loomis & Davenport, 1951); M. tiemanni Causey, 1960; M. tularea tularea (Chamberlin, 1949), M. tularea ollae Causey & Tiemann, 1969


  57. Parcipromus Shelley, 1995 - 3 species: P. cooki (Causey, 1995); P. gigantoarboricolus Shelley, 1995; P. tiemanni Shelley, 1995


  58. Wamokia Chamberlin, 1941 - 8 species: W. dentata Buckett & Gardner, 1968; W. discordis Buckett & Gardner, 1968; W. falcata Buckett & Gardner, 1968; W. hoffmani Buckett & Gardner, 1968; W. placera Chamberlin, 1941; W. remota Buckett & Gardner, 1968; W. sierrae Buckett & Gardner, 1968


  59. Xystocheir Cook, 1904 - 9 species: X. bistipita Shelley, 1996; X. brachymacris Shelley, 1996; X. dissecta dissecta (Wood, 1867); X. d. microrama Shelley, 1996; X. d. taibona Chamberlin, 1912; X. modestior modestior (Chamberlin, 1941); X. m. haerens Shelley, 1996; X. prolixorama Shelley, 1996; X. reducta (Causey, 1955); X. solenofurcata Shelley, 1996; X. stenomacris Shelley, 1996; X. stolonifera stolonifera Shelley, 1996; X. s. uncinata Shelley, 1996

^ Appalachioria eutypa eutypa from Grandfather Mountain, Avery Co. North Carolina