Melolonthinae

Hoplia coerulea, copyright Frank Vassen.

Belongs within: Scarabaeoidea.
Contains: Rhizotrogus, Automolius, Aplopsis, Maechidius, Systellopini, Liparetrus, Colpochila, Phyllotocus, Xylonichini, Dynastinae, Euchirini, Melolonthini, Rutelinae, Cetoniinae, Diphucephala, Sericini, Heteronyx, Neoheteronyx, Gnaphalopoda, Sericesthis, Scitala.

The Melolonthinae, chafers, are a cosmopolitan group of scarabaeid beetles with soil-dwelling larvae that feed on roots and hummus. Though larvae may live for up to two years, the adult life period is relatively brief and lasts only a few days or weeks. Adults of some species feed on the leaves of trees; others do not feed as adults at all (Houston & Weir 1992). As currently recognised, the melolonthines are probably not monophyletic with other phytophagous scarabaeid subfamilies such as Dynastinae, Rutelinae and Cetoniinae nested within the group.

Members of the Melolonthinae are divided between several tribes. In many of these tribes, the posterior tibia has two apical spurs that are placed above and below the tibia so that the tarsus passes between the spurs when moved from side to side, but both spurs are placed ventrally in Plectris, Pachytricha, Diphucephalini and Melolonthini. Members of the Australian genus Phyllotocidium are at least partially metallic in coloration and have a three-segmented antennal club. The Phyllotocini have deeply projecting fore coxae, large hind coxae, a slender process with a setose ending usually present on the prosternum, and strongly asymmetrical claws in the male (Britton 1978).

Characters (from Houston & Weir 1992): Head and pronotum of males unarmed; mandibles completely concealed from above; abdominal spiracles diverging, with one pair exposed beneath edge of elytra; fore coxae transverse, mesothoracic epimera not visible from above; mid coxae transverse or only slightly oblique; tarsal claws equal. Larvae with apical antennal segment about as wide as penultimate segment; mandibles withouyt transverse granular ridges forming stridulatory areas; galea and lacinia either partly fused proximally or fitting tightly together; anal cleft usually Y-shaped or angulate.

<==Melolonthinae [Chasmatopterinae, Melolonthidae, Melolonthides, Sericinae]
    |--Hoplia [Hopliini] A06
    |    |--H. aureola [incl. H. aureola var. sibirica] C01
    |    |--H. coerulea (Drury 1773) B14
    |    |--H. equina LeConte 1880 A06
    |    |--H. farinosa [=Melolontha farinosa] G20
    |    |--H. graminicola (Fabricius 1792) A06
    |    |--H. parvula C01
    |    `--H. sulphurea Chevrolat 1846 E12
    |--Phyllotocini [Phylloticides, Phyllotocina] A06
    |    |--Phyllotocus A06
    |    `--Anthotocus Britton 1957 A06, HW92
    |         |--*A. luridus (Macleay 1864) HW92 (see below for synonymy)
    |         |--A. antennalis (Lea 1919) [=Phyllotocus antennalis] HW92
    |         |--A. cribriceps (Lea 1919) [=Phyllotocus cribriceps] HW92
    |         |--A. fugitivus (Lea 1920) [=Phyllotocus fugitivus] HW92
    |         `--A. perissus Britton 1957 HW92
    |--+--Xylonichini A06
    |  `--+--Chasmatopterus [Chasmatopterini] A06
    |     |    |--C. annamariae Branco 2004 F05
    |     |    `--C. hirtus (Illiger 1803) A06
    |     |--Pachydemini A06
    |     |    |--Tanyproctus persicus (Ménétries 1832) A06
    |     |    `--+--Elaphocera A06
    |     |       |    |--E. capdeboni Schauffuss 1882 A06
    |     |       |    `--E. tangerina Kraatz 1888 A06
    |     |       `--+--Acoma Casey 1881 A06, N02
    |     |          |--Europteron gracile Marseul 1867 A06
    |     |          `--Pachypus [Pachypodinae] A06
    |     |               `--P. candidae (Petagna 1786) A06
    |     `--+--Dynastinae A06
    |        |--Euchirini A06
    |        |--Melolonthini A06
    |        |--+--Rutelinae A06
    |        |  `--Cetoniinae A06
    |        `--Pleocominae [Pleocomidae] MW15
    |             |--Crecotoma Nikolajev 2002 [Cretocomini] N02
    |             |    `--*C. tologoica Nikolajev 2002 N02
    |             `--Pleocoma Le Conte 1856 [Pleocomini] N02
    |                  `--P. australis Fall 1911 B14
    `--+--Diphucephalini [Diphucephalides] HW92
       |    |--Diphucephala A06
       |    `--Cunderdinia Lea 1916 HW92
       |         |--*C. variabilis Lea 1916 HW92
       |         `--C. setistriata Lea 1930 HW92
       `--+--+--Sericoides [Sericoidini] A06
          |  `--+--+--Athlia rustica Erichson 1835 A06
          |     |  `--+--Sericini A06
          |     |     `--Ablaberini [Camentini] A06
          |     |          |--Ablabera hirsuta Blanchard 1850 A06
          |     |          |--Cyrtocamenta A06
          |     |          |    |--C. pygidialis Frey 1968 A06
          |     |          |    `--C. pygmaea Brenske 1897 A06
          |     |          `--+--Empecamenta buettikeri Ahrens 2000 A06
          |     |             `--Camenta A06
          |     |                  |--C. elongata Frey 1960 A06
          |     |                  `--C. westermanni Harold 1878 A06
          |     `--Heteronycini HW92
          |          |--Heteronyx A06
          |          |--Neoheteronyx HW92
          |          |--Callabonica Blackburn 1895 HW92
          |          |    `--*C. propria Blackburn 1895 HW92
          |          |--Cubidens Britton in Houston & Weir 1992 [=Paraheteronyx Britton 1988 non Moser 1924] HW92
          |          |    `--*C. unguiculatus (Burmeister 1855) (see below for synonymy) HW92
          |          |--Eurychelus Blanchard 1850 HW92
          |          |    `--*E. marmoratus Blanchard 1850 HW92
          |          |--Nepytis Erichson 1842 HW92
          |          |    `--*N. russula Erichson 1842 HW92
          |          |--Proborhinus Britton 1988 HW92
          |          |    `--*P. cornutus (Blackburn 1910) [=Heteronyx cornutus] HW92
          |          |--Odontotonyx Macleay 1871 HW92
          |          |    |--*O. brunneipennis Macleay 1871 HW92
          |          |    `--O. ruficeps Lea 1919 HW92
          |          |--Webbella Britton 1988 HW92
          |          |    |--*W. firma (Blackburn 1909) [=Heteronyx firmus] HW92
          |          |    `--W. labralis (Blackburn 1908) HW92 (see below for synonymy)
          |          |--Acheilo Britton 1988 HW92
          |          |    |--*A. capitalis (Blackburn 1910) [=Heteronyx capitalis] HW92
          |          |    |--A. clypeatus Britton 1988 HW92
          |          |    `--A. pedarius (Blackburn 1910) [=Heteronyx pedarius] HW92
          |          `--Pseudoheteronyx Blackburn 1892 HW92
          |               |--*P. baldiensis (Blackburn 1892) [=Heteronyx baldiensis] HW92
          |               |--P. basicollis Lea 1919 HW92
          |               |--P. creber Blackburn 1908 [incl. P. seticollis Lea 1919] HW92
          |               |--P. helaeoides Blackburn 1892 HW92
          |               |--P. laticollis Blackburn 1908 HW92
          |               `--P. puncticollis Lea 1919 HW92
          `--Scitalini HW92
               |--Gnaphalopoda HW92
               |--Sericesthis HW92
               |--Scitala HW92
               |--Colpochelyne Britton 1987 HW92
               |    `--*C. dux Britton 1987 HW92
               |--Idanastes Britton 1987 HW92
               |    `--*I. abditus Britton 1987 HW92
               |--Phorine Britton 1987 HW92
               |    `--*P. anomala (Blackburn 1907) [=Byrrhomorpha anomala] HW92
               |--Synchilus Britton 1955 HW92
               |    `--*S. gisleni Britton 1955 HW92
               |--Xyrodes Britton 1987 HW92
               |    `--*X. calorata (Blackburn 1907) [=Frenchella calorata] HW92
               |--Protelura Britton 1987 HW92
               |    |--*P. nana Britton 1987 HW92
               |    `--P. guttata Britton 1987 HW92
               |--Hadrops Britton 1987 HW92
               |    |--*H. halei Britton 1987 HW92
               |    `--H. flavus (Lea 1919) [=Engyops flavus] HW92
               |--Colpochilodes Blackburn 1898 HW92
               |    |--*C. raucipennis Blackburn 1898 HW92
               |    `--C. peregrina Britton 1987 HW92
               |--Xyrine Britton 1987 HW92
               |    |--*X. carnei Britton 1987 HW92
               |    `--X. inusitatus Britton 1987 HW92
               |--Byrrhomorpha Blackburn 1892 HW92
               |    |--*B. verres Blackburn 1892 HW92
               |    |--B. basicollis Lea 1919 HW92
               |    `--B. ponderosa Blackburn 1892 [incl. B. rudis Lea 1919] HW92
               |--Ophropyx Britton 1987 HW92
               |    |--*O. hispida (Blackburn 1898) [=Frenchella hispida] HW92
               |    |--O. approximans (Blackburn 1898) [=Frenchella approximans] HW92
               |    `--O. ciliata (Boisduval 1835) HW92 (see below for synonymy)
               |--Xyroa Britton 1987 HW92
               |    |--*X. polita Britton 1987 HW92
               |    |--X. alta Britton 1987 HW92
               |    |--X. amoena Britton 1987 HW92
               |    `--X. nitida Britton 1987 HW92
               |--Homolotropus Macleay 1871 [=Homalotropus (l. c.)] HW92
               |    |--*H. luridipennis Macleay 1871 HW92
               |    |--H. metallicus Britton 1970 HW92
               |    |--H. sagus Britton 1970 HW92
               |    |--H. sericeus Britton 1970 HW92
               |    `--H. taylori Britton 1970 HW92
               `--Telura Erichson 1842 A06, HW92
                    |--*T. vitticollis Erichson 1842 HW92
                    |--T. alta Britton 1987 HW92
                    |--T. imparilis Britton 1987 HW92
                    |--T. monticola Britton 1987 HW92
                    `--T. petiolata Britton 1987 HW92

Melolonthinae incertae sedis:
  Pyronota T27
    |--P. festiva T27
    |--P. laeta B35
    `--P. refulgens Dupont in Boisduval 1835 B35
  Ancistrosoma klugii Curtis 1835 B14
  Polyphylla B14
    |--P. decemlineata (Say 1824) B14
    `--P. fullo B28
  Chlorochiton suturalis [=Stethaspis suturalis] L19
  Rhizotrogus M86
  Chnaunanthus S38
  Comophorina Strand 1928 (see below for synonymy) HW92
    `--*C. testaceipennis (Blanchard 1850) [=*Comophorus testaceipennis, *Comophus testaceipennis] HW92
  Plectris Saint-Fargeau & Serville 1828 [Macrodactylini] HW92
    |--*P. tomentosa Saint-Fargeau & Serville 1828 HW92
    `--P. aliena Chapin 1934 HW92
  Phyllotocidium Blackburn 1898 [Phyllotocidiini] HW92
    |--*P. macleayi (Blackburn 1892) [=Cheiragra macleayi] HW92
    |--P. bimaculiflavum Lea 1917 HW92
    |--P. pictum Lea 1916 HW92
    `--P. viridis Britton 1957 HW92
  Pachytricha Hope 1841 [=Pachytrichia (l. c.); Pachytrichini] HW92
    |--*P. castanea Hope 1841 [incl. P. munda Sharp 1874, P. pallens Sharp 1874] HW92
    |--P. demarzi Frey 1966 HW92
    |--P. minor Sharp 1874 HW92
    |--P. robusta Sharp 1874 HW92
    `--P. tecta Sharp 1874 HW92
  Automoliini [Automolini] HW92
    |--Automolius HW92
    |--Aplopsis HW92
    |--Bryantella Britton 1957 HW92
    |    `--*B. castanea Britton 1957 HW92
    |--Maechidinus Lea 1919 HW92
    |    |--*M. latericollis Lea 1919 HW92
    |    |--M. marginalis Lea 1919 HW92
    |    `--M. sculptilis Britton 1957 HW92
    `--Phyllochlaenia Blanchard 1846 (see below for synonymy) HW92
         |--*P. rufescens Blanchard 1846 HW92 (see below for synonymy)
         |--‘Caulobius’ modestus Blanch. 1850 M86
         `--P. villosus (Guillou 1844) (see below for synonymy) HW92
  Maechidiini [Maechidiides] HW92
    |--Maechidius HW92
    |--Microcoenus Britton 1957 HW92
    |    `--*M. nanus Britton 1957 HW92
    |--Harpechys Britton 1957 HW92
    |    `--*H. chilo Britton 1957 HW92
    |--Microthopus Burmeister 1855 HW92
    |    `--*M. castanopterus Burmeister 1855 [incl. Macleayia hybrida Blackburn 1888] HW92
    |--Termitophilus Britton 1957 HW92
    |    `--*T. spadix Britton 1957 HW92
    `--Epholcis Waterhouse 1875 HW92
         |--*E. divergens Waterhouse 1875 [incl. Maechidius albertisi Fairmaire 1877] HW92
         |--E. bilobiceps (Fairmaire 1877) [=Maechidius bilobiceps] HW92
         |--E. gracilis (Waterhouse 1875) [=Maechidius gracilis] HW92
         |--E. longior Blackburn 1898 HW92
         `--E. uniformis Britton 1957 HW92
  Systellopini HW92
  Liparetrini [Colpochilini] HW92
    |  i. s.: ‘Melolontha’ astrolabei Boisduval 1835 [=Haplonycha astrolabei] HW92
    |--Liparetrus HW92
    |--Colpochila HW92
    |--Plectrodes S38
    |--Dysphanochila Blackburn 1898 HW92
    |    `--*D. pilosipennis Blackburn 1898 HW92
    |--Parasciton Britton 1990 HW92
    |    `--*P. inermis Britton 1990 HW92
    |--Allara Britton 1955 HW92
    |    `--*A. insularis Britton 1955 HW92
    |--Anacanthodes Britton 1990 [=Anacanthopus Blackburn 1898 non Montandon 1894] HW92
    |    `--*A. inermis (Blackburn 1898) [=*Anacanthopus inermis] HW92
    |--Macleayella Britton 1990 [=Macleayia Blackburn 1888 non Haswell 1880] HW92
    |    `--*M. singularis (Blackburn 1888) [=*Macleayia singularis] HW92
    |--Aneucomides Blackburn 1898 HW92
    |    `--*A. coloratus Blackburn 1898 [incl. Haplonycha mauricei Blackburn 1906] HW92
    |--Aphanesia Britton 1990 HW92
    |    `--*A. greyi Britton 1990 HW92
    |--Astibicola Britton 1990 HW92
    |    `--*A. bicolor Britton 1990 HW92
    |--Cheilo Britton 1990 HW92
    |    `--*C. liparetroides Britton 1990 HW92
    |--Dikellites Britton 1990 HW92
    |    `--*D. abditus Britton 1990 HW92
    |--Engyopsina Britton 1990 [=Engyops Blackburn 1898 non Bauer & Bergenstamm 1889] HW92
    |    `--*E. spectans (Blackburn 1898) [=Engyops spectans] HW92
    |--Hadropechys Britton 1990 HW92
    |    `--*H. victoriae (Blackburn 1897) [=Pachygastra victoriae] HW92
    |--Ictigaster Britton 1986 HW92
    |    `--*I. ruficollis (Lea 1917) [=Haplonycha ruficollis] HW92
    |--Leonotus Britton 1986 HW92
    |    `--*L. pilosicollis (Lea 1930) [=Haplonycha pilosicollis] HW92
    |--Pachygastra Germar 1848 HW92
    |    `--*P. tasmanica Germar 1848 [incl. Prochelyna rubella Schaufuss 1882] HW92
    |--Paronyx Britton 1990 HW92
    |    `--*P. setifera Britton 1990 HW92
    |--Petinopus Blackburn 1898 HW92
    |    `--*P. aegrotus Blackburn 1898 HW92
    |--Scitaloides Britton 1990 HW92
    |    `--*S. malanda Britton 1990 HW92
    |--Stenochelyne Britton 1990 HW92
    |    `--*S. noctis Britton 1990 HW92
    |--Xyridea Britton 1990 HW92
    |    `--*X. hirticollis (Blackburn 1912) [=Aneucomides hirticollis] HW92
    |--Nosphisthis Blackburn 1898 HW92
    |    |--*N. parvicornis Blackburn 1898 HW92
    |    `--N. perkinsi Blackburn 1907 HW92
    |--Biphyllocera White 1841 [=Diphyllocera Agassiz 1847] HW92
    |    |--*B. kirbyana White 1841 HW92 [=Diphyllocera kirbyana M86]
    |    `--B. fabriciana White 1841 HW92
    |--Glossocheilifer Blackburn 1898 HW92
    |    |--*G. labialis Blackburn 1898 [incl. G. addendus Blackburn 1907] HW92
    |    `--G. bidentatus Lea 1919 HW92
    |--Teluroides Britton 1990 HW92
    |    |--*T. suturalis (Lea 1920) [=Telura suturalis] HW92
    |    `--T. clypealis (Lea 1919) [=Telura clypealis] HW92
    |--Sciton Blackburn 1892 HW92
    |    |--*S. ruber Blackburn 1892 HW92
    |    |--S. flavocastaneus Lea 1917 HW92
    |    |--S. paullus Blackburn 1898 HW92
    |    `--S. variicollis Blackburn 1907 HW92
    |--Neso Blackburn 1898 HW92
    |    |--*N. usta Blackburn 1898 HW92
    |    |--N. castaneus (Lea 1920) [=Engyops castaneus] HW92
    |    |--N. ducalis Blackburn 1907 HW92
    |    |--N. flavipennis (Macleay 1887) (see below for synonymy) HW92
    |    |--N. minuta (Lea 1926) HW92 [=Haplonycha minuta HW92, Pteroplatydesmus minutus B86]
    |    `--N. yorkensis Blackburn 1898 HW92
    |--Frenchella Blackburn 1892 HW92
    |    |--*F. lubrica Blackburn 1892 HW92
    |    |--F. cribriceps Lea 1919 HW92
    |    |--F. fimbriata Lea 1919 HW92
    |    |--F. gagatina Lea 1919 HW92
    |    |--F. hirticollis Blackburn 1898 HW92
    |    |--F. iridescens (Blanchard 1850) [=Haplonycha iridescens] HW92
    |    `--F. russula Britton 1959 HW92
    `--Colobostoma Blanchard 1850 (see below for synonymy) HW92
         |--*C. rufipennis (Boisduval 1835) (see below for synonymy) HW92
         |--C. castaneus (Lea 1919) [=Platydesmus castaneus] HW92
         |--C. hirsuta (Frey 1966) [=Frenchella hirsuta] HW92
         |--C. inamoenus (Blackburn 1907) [=Platydesmus inamoenus] HW92
         |--C. inusitatus (Blackburn 1907) [=Platydesmus inusitatus] HW92
         |--C. major (Blackburn 1907) [=Platydesmus major] HW92
         |--C. obscuricornis (Blanchard 1850) [=Haplonycha obscuricornis] HW92
         `--C. punctulaticeps (Blackburn 1908) [=Platydesmus punctulaticeps] HW92
  Lithanomala A06
  Cretoserica A06

*Anthotocus luridus (Macleay 1864) HW92 [=Cheiragra lurida HW92, Phyllotocus lurida B57; incl. P. truncatidens Lea 1920 HW92]

Colobostoma Blanchard 1850 [incl. Platydesmus Macleay 1887 non Lucas 1843, Pteroplatydesmus Dalla Torre 1912] HW92

*Colobostoma rufipennis (Boisduval 1835) [=Sericesthis rufipennis; incl. Frenchella sparsiceps Blackburn 1898, *Platydesmus sulcipennis Macleay 1887, *Pteroplatydesmus sulcipennis] HW92

Comophorina Strand 1928 [=Comophorus Blanchard 1850 non Agassiz 1846, Comophus Britton 1978; Comophini, Comophorini, Comophorinini] HW92

*Cubidens unguiculatus (Burmeister 1855) [=Heteronyx unguiculata, *Paraheteronyx unguiculata] HW92

Neso flavipennis (Macleay 1887) [=Platydesmus flavipennis; incl. N. planicollis Blackburn 1898] HW92

Ophropyx ciliata (Boisduval 1835) HW92 [=Melolontha ciliata HW92, Aplonycha ciliata B86, Haplonycha ciliata B86, Heteronyx ciliata B86; incl. Frenchella aspericollis Blackburn 1898 HW92, Ha. rugosa Burmeister 1855 HW92]

Phyllochlaenia Blanchard 1846 [=Philochlaenia (l. c.); incl. Caulobius Guillou 1844 non Duponchel 1838, Deuterocaulobius Dalla Torre 1912] HW92

*Phyllochlaenia rufescens Blanchard 1846 HW92 [=Caulobius rufescens M86, Deuterocaulobius rufescens B57, Heteronyx rufescens B57, Philochlaenia rufescens B57; incl. Caulobius advena Blackburn 1898 HW92]

Phyllochlaenia villosus (Guillou 1844) HW92 [=*Caulobius villosus HW92, *Deuterocaulobius villosus HW92; incl. Hymenoplia villigera Blanchard 1846 HW92, Caulobius villiger M86, Omaloplia villigera B57]

Webbella labralis (Blackburn 1908) HW92 [=Heteronyx labralis HW92; incl. H. anomalus Blackburn 1908 B00, HW92, H. leai Blackburn 1909 B00, HW92]

*Type species of generic name indicated

REFERENCES

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[B28] Betrem, J. G. 1928. Monographie der Indo-Australischen Scoliiden mit zoogeographischen Betrachtungen. H. Veenman & Zonen: Wageningen.

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d'Urville. Faune entomologique de l'océan Pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

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[B86] Britton, E. B. 1986. A revision of the Australian chafers (Coleoptera: Scarabaeidae: Melolonthinae) vol. 4. Tribe Liparetrini: genus Colpochila. Australian Journal of Zoology, Supplementary Series 118: 1–135.

[B00] Britton, E. B. 2000. A review of Heteronyx Guérin-Méneville (Coleoptera: Scarabaeidae: Melolonthinae). Invertebrate Taxonomy 14: 465–589.

[C01] Csiki, E. 1901. Bogarak [Coleopteren]. 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. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[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.

[F05] Fernández, J. 2005. Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (IX). Graellsia 61 (2): 261–282.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[HW92] Houston, W. W. K., & T. A. Weir. 1992. Melolonthinae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 174–358. AGPS Press: Canberra.

[L19] Lea, A. M. 1919. Notes on some miscellaneous Coleoptera with descriptions of new species.—Part V. Transactions of the Royal Society of South Australia 43: 166–261, pls 25–27.

[M86] Masters, G. 1886. Catalogue of the described Coleoptera of Australia. Part III. Proceedings of the Linnean Society of New South Wales, series 2, 1 (1): 21–126.

[MW15] McKenna, D. D., A. L. Wild, K. Kanda, C. L. Bellamy, R. G. Beutel, M. S. Caterino, C. W. Farnum, D. C. Hawks, M. A. Ivie, M. L. Jameson, R. A. B. Leschen, A. E. Marvaldi, J. V. McHugh, A. F. Newton, J. A. Robertson, M. K. Thayer, M. F. Whiting, J. F. Lawrence, A. Ślipiński, D. R. Maddison & B. D. Farrell. 2015. The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology 40 (4): 835–880.

[N02] Nikolajev, G. V. 2002. To possible assignment of a new genus of lamellicorn beetles from the Lower Cretaceous of Mongolia to the subfamily Pleocominae (Coleoptera, Scarabaeidae). Paleontologicheskii Zhurnal 2002 (3): 51–55 (transl. Paleontological Journal 36 (3): 279–282).

[S38] Saylor, L. W. 1938. Revision of the subfamily Oncerinae with description of a new genus (Coleoptera: Scarabaeidae). Proceedings of the Entomological Society of Washington 40 (4): 99–103.

[T27] Thomson, G. M. 1927. The pollination of New Zealand flowers by birds and insects. Transactions and Proceedings of the New Zealand Institute 57: 106–125.

Last updated: 28 December 2021.

Palpatores

Acropsopilio neozelandiae, copyright Stephen Thorpe.

Belongs within: Opiliones.
Contains: Ischyropsalidoidea, Troguloidea, Phalangioidea.

The Palpatores, the long-legged harvestmen, are a diverse group of harvestmen including many of the most familiar members of the order. Members of this clade have ozopores flush with the body surface (not raised on ozophores) and simple, single claws on the legs; most (but not all) also have particularly elongate legs. Basal members of the Palpatores with small bodies and greatly enlarged eye mounds were classified until recently in a family Caddidae, but phylogenetic analyses have recently indicated that this group is para- or polyphyletic, and may represent the plesiomorphic appearance of the clade as a whole (Groh & Giribet 2015).

Palpatores [Apagosterni, Caddidae, Caddoidea, Plagiostethi]
    |--Dyspnoi [Nemastomini, Nemastomoidae, Trogulini] GG15
    |    |  i. s.: Echinopustulus samuelnelsoni Dunlop 2004 D07
    |    |--Acropsopilionidae [Acropsopilioninae, Acropsopilionoidea] GFB14
    |    |    |--Caddella Hirst 1925 SC06 [=Cadella (l. c.) K61; incl. Oonopsopilio Lawrence 1931 SC03]
    |    |    |    |--*C. capensis Hirst 1925 S75 [=C. (Oonopsopilio) capensis K61]
    |    |    |    |--C. africana (Lawrence 1931) [=*Oonopsopilio africanus] S75
    |    |    |    |--C. croeseri Staręga 1988 S92
    |    |    |    |--C. haddadi Lotz 2011 L11
    |    |    |    |--C. jocquei Staręga 2008 L11
    |    |    |    `--C. spatulipilis Lawrence 1934 S92 (see below for synonymy)
    |    |    |--Austropsopilio Forster 1955 [incl. Tasmanopilio Hickman 1957] GG15
    |    |    |    |--*A. novaehollandiae Forster 1955 SC03
    |    |    |    |--A. altus Cantrell 1980 C80
    |    |    |    |--A. cygneus Hickman 1957 H57
    |    |    |    |--A. fuscus (Hickman 1957) GG15 [=*Tasmanopilio fuscus H57]
    |    |    |    |--A. inermis Cantrell 1980 C80
    |    |    |    |--A. megalops (Hickman 1957) [=Tasmanopilio megalops] GG15
    |    |    |    `--A. sudamericanus Shultz & Cekalovic 2003 SC03
    |    |    `--Acropsopilio Silvestri 1904 SC06 [incl. Zeopsopilio Forster 1948 SC03]
    |    |         |--*A. chilensis Silvestri 1904 S75 (see below for synonymy)
    |    |         |--A. australicus Cantrell 1980 C80
    |    |         |--A. boopis (Crosby 1904) [=Caddo boopis] S75
    |    |         |--A. chomulae (Goodnight & Goodnight 1948) [=Caddo chomulae, A. chommulae (l. c.)] S75
    |    |         |--A. neozelandiae (Forster 1948) GFB14 [=*Zeopsopilio neozelandiae F48]
    |    |         |--A. talishensis Morin 1937 S04
    |    |         `--A. venezuelensis González-Sponga 1992 GG15
    |    `--+--Ischyropsalidoidea GG15
    |       `--Troguloidea GG15
    `--Eupnoi [Phalangioidae] GG15
         |  i. s.: Brigantibunum Dunlop & Anderson 2005 DA05
         |           `--*B. listoni Dunlop & Anderson 2005 DA05
         |         Daohugopilio Huang, Selden & Dunlop 2009 HSD09
         |           `--*D. sheari Huang, Selden & Dunlop 2009 HSD09
         |--Mesobunus Huang, Selden & Dunlop 2009 GS14, HSD09
         |    |--*M. martensi Huang, Selden & Dunlop 2009 HSD09
         |    `--M. dunlopi Giribet, Tourinho et al. 2012 GT12
         `--+--Phalangioidea GS14
            `--+--Macrogyion Garwood, Dunlop et al. 2011 GS14, GD11
               |    `--*M. cronus Garwood, Dunlop et al. 2011 GD11
               `--Caddo Banks 1892 GS14, S75 [Caddinae, Caddini, Caddoinae]
                    |--*C. agilis Banks 1892 [incl. C. glaucopis Crosby 1904] S75
                    |--C. dentipalpus (Koch & Berendt 1854) [=Platybunus dentipalpus] S02
                    `--C. pepperella Shear 1975 S75

Nomen nudum: Caddo medama Suzuki 1958 GG15

*Acropsopilio chilensis Silvestri 1904 S75 [incl. A. normae Cekalovic 1974 GG15, A. chilensis var. ogloblini Canals 1932 S75]

Caddella spatulipilis Lawrence 1934 S92 [=Cadella spatulipilis L34, Caddella spatulipalpis (l. c.) S75; incl. Cadella spatulipilis var. caledonica Lawrence 1934 L34]

*Type species of generic name indicated

REFERENCES

[C80] Cantrell, B. K. 1980. Additional Australian harvestmen (Arachnida: Opiliones). Journal of the Australian Entomological Society 19: 241–253.

[D07] Dunlop, J. A. 2007. Paleontology. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 247–265. Harvard University Press: Cambridge (Massachusetts).

[DA05] Dunlop, J. A., & L. I. Anderson. 2005. A fossil harvestman (Arachnida, Opiliones) from the Mississippian of East Kirkton, Scotland. Journal of Arachnology 33: 482–489.

[F48] Forster, R. R. 1948. A new genus and species of the family Acropsopilionidae (Opiliones) from New Zealand. Transactions of the Royal Society of New Zealand 77 (1): 139–141.

[GD11] Garwood, R. J., J. A. Dunlop, G. Giribet & M. D. Sutton. 2011. Anatomically modern Carboniferous harvestmen demonstrate early cladogenesis and stasis in Opiliones. Nature Communications 2 (444): 1–7.

[GS14] Garwood, R. J., P. P. Sharma, J. A. Dunlop & G. Giribet. 2014. A Paleozoic stem group to mite harvestmen revealed through integration of phylogenetics and development. Current Biology 24 (9): 1017–1023.

[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.

[GT12] Giribet, G., A. L. Tourinho, C. Shih & D. Ren. 2012. An exquisitely preserved harvestman (Arthropoda, Arachnida, Opiliones) from the Middle Jurassic of China. Organisms, Diversity & Evolution 12: 51–66.

[GG15] Groh, S., & G. Giribet. 2015. Polyphyly of Caddoidea, reinstatement of the family Acropsopilionidae in Dyspnoi, and a revised classification system of Palpatores (Arachnida, Opiliones). Cladistics 31 (3): 277–290.

[H57] Hickman, V. V. 1957. Some Tasmanian harvestmen of the sub-order Palpatores. Papers and Proceedings of the Royal Society of Tasmania 91: 65–79.

[HSD09] Huang, D., P. A. Selden & J. A. Dunlop. 2009. Harvestmen (Arachnida: Opiliones) from the Middle Jurassic of China. Naturwissenschaften 96 (8): 955–962.

[K61] Kauri, H. 1961. Opiliones. In: Hanström, B., P. Brinck & G. Rudebeck. South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 9–197. Almqvist & Wiksell: Uppsala.

[L34] Lawrence, R. F. 1934. New South African Opiliones. Annals of the South African Museum 30 (4): 549–586.

[L11] Lotz, L. N. 2011. Three new harvestmen species from southern Africa (Arachnida: Opiliones: Caddidae, Neopilionidae, Assamiidae). Journal of Afrotropical Zoology 7: 3–8.

[S75] Shear, W. A. 1975. The opilionid family Caddidae in North America, with notes on species from other regions (Opiliones, Palpatores, Caddoidea). Journal of Arachnology 2: 65–88.

[SC03] Shultz, J. W., & T. Cekalovic. 2003. First species of Austropsopilio (Opiliones, Caddoidea, Caddidae) from South America. Journal of Arachnology 31: 20–27.

[SC06] Shultz, J. W., & T. Cekalovic. 2006. First species of Hesperopilio (Opiliones, Caddoidea) from South America. Journal of Arachnology 34 (1): 46–50.

[S04] Snegovaya, N. Y. 2004. Preliminary notes on the harvestman fauna (Opiliones) of Azerbaijan. Arthropoda Selecta Special Issue 1: 307–318.

[S92] Staręga, W. 1992. An annotated check-list of Afrotropical harvestmen, excluding the Phalangiidae (Opiliones). Annals of the Natal Museum 33 (2): 271–336.

[S02] Staręga, W. 2002. Baltic amber harvestmen (Opiliones) from Polish collections. Annales Zoologici 52 (4): 601–604.

Lepidiota

Lepidiota stigma, copyright U. Schmidt.

Belongs within: Melolonthini.

Lepidiota is a genus of chafers found in southern Asia and northern Australasia, many of which have a dorsal covering of white scales. Some species are notable pests with their larvae feeding on crops such as sugarcane (Britton 1978).

Characters (from Britton 1978): Anterior face of clypeus shallow (ratio of greatest width:mid >8.0:1), usually smooth and unpunctured in middle; anterior edge of clypeus as seen from above usually broadly bilobed; antenna 10- (rarely 9-)segmented, with a 3-lamellate club; lamellae usually shorter than antennal segments 1–7 together; surface of body usually bearing few or many broad adpressed white scales, scales sometimes minute and wholly contained within their punctures; tarsal claws each with strong tooth in middle of concave side; anterior edge of posterior femora not concave near base; mandibles not curved downwards at apices.

Lepidiota Kirby in Kirby & Spence 1828 HW92
    |--*L. stigma (Fabricius 1798) [=Melolontha stigma] HW92
    |--L. aenigma Britton 1978 HW92
    |--L. alticalceus Allsopp 1989 HW92
    |--L. amitina Britton 1978 HW92
    |--L. argentea Britton 1978 HW92
    |--L. arnhemensis Britton 1978 HW92
    |--L. aspera Britton 1978 HW92
    |--L. bidentata Allsopp 1989 HW92
    |--L. cabala Britton 1978 HW92
    |--L. caudata Blackburn 1890 HW92
    |--L. centralis Britton 1978 HW92
    |--L. ciliata Britton 1978 HW92
    |--L. clypealis Moser 1913 HW92
    |--L. comes Britton 1978 HW92
    |--L. condita Britton 1978 HW92
    |--L. consobrina Girault 1918 HW92
    |--L. contigua Britton 1978 HW92
    |--L. crinita Brenske 1900 [incl. L. trichosterna Lea 1924] HW92
    |--L. degener Blackburn 1888 HW92
    |--L. delicatula Blackburn 1888 [incl. L. suavior Blackburn 1912] HW92
    |--L. draconis Britton 1978 HW92
    |--L. flavipennis Lea 1926 HW92
    |--L. frenchi Blackburn 1912 HW92
    |--L. froggatti Macleay 1887 HW92
    |--L. gibbifrons Britton 1978 HW92
    |--L. gilesi Blackburn 1912 HW92
    |--L. grata Blackburn 1890 HW92
    |--L. grisea Britton 1959 HW92
    |--L. helleri Moser 1913 HW92
    |--L. hilli Britton 1978 HW92
    |--L. hylaea Britton 1978 HW92
    |--L. laciniata Britton 1978 HW92
    |--L. laevis Arrow 1932 HW92
    |--L. lepidosterna Lea 1926 HW92
    |--L. lewisae Britton 1978 HW92
    |--L. minuta Moser 1913 HW92
    |--L. negatoria Blackburn 1912 [incl. L. deceptrix Blackburn 1912, L. mungomeryi Britton 1963] HW92
    |--L. noxia Britton 1985 HW92
    |--L. oblonga Brenske 1900 HW92
    |--L. oryx Britton 1959 HW92
    |--L. perkinsi Blackburn 1912 HW92
    |--L. picticollis Lea 1916 HW92
    |--L. podicalis Moser 1913 [incl. L. platyura Lea 1924] HW92
    |--L. quinaria Britton 1978 HW92
    |--L. quinquelineata M86
    |--L. rothei Blackburn 1888 [incl. L. bovilli Blackburn 1912, L. koebelei Blackburn 1912, L. parva Moser 1913] HW92
    |--L. rubrior Blackburn 1912 [incl. L. rugicollis Moser 1913] HW92
    |--L. rufa Blackburn 1888 HW92
    |--L. salax Britton 1978 HW92
    |--L. sororia Moser 1913 [incl. L. wilsoni Britton 1959] HW92
    |--L. sparsa Britton 1978 HW92
    |--L. squalida Macleay 1886 M86
    |--L. squamulata Waterhouse 1875 (see below for synonymy) HW92
    |--L. sticta Britton 1978 HW92
    |--L. stradbrokensis Lea 1919 [=L. froggatti stradbrokensis] HW92
    |--L. townsvillensis Blackburn 1912 HW92
    |--L. trihomines Allsopp 1990 HW92
    |--L. ultima Britton 1978 HW92
    |--L. uptoni Britton 1978 HW92
    |--L. verna Britton 1978 HW92
    |--L. weiri Britton 1978 HW92
    `--L. yorkensis Britton 1978 HW92

Lepidiota squamulata Waterhouse 1875 [incl. L. darwini Blackburn 1888, L. leai Blackburn 1912, L. rugosipennis Lea 1924] HW92

*Type species of generic name indicated

REFERENCES

Britton, E. B. 1978. A revision of the Australian chafers (Coleoptera: Scarabaeidae: Melolonthinae) vol. 2. Tribe Melolonthini. Australian Journal of Zoology, Supplementary Series 60: 1–150.

[HW92] Houston, W. W. K., & T. A. Weir. 1992. Melolonthinae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 174–358. AGPS Press: Canberra.

[M86] Macleay, W. 1886. The insects of the Fly River, New Guinea, "Coleoptera". Proceedings of the Linnean Society of New South Wales, series 2, 1 (1): 136–157.

Last updated: 27 April 2021.

Antitrogus

Childers canegrub Antitrogus parvulus, from PaDIL.

Belongs within: Melolonthini.

Antitrogus is a genus of chafers endemic to eastern Australia. They are rarely collected, seemingly not feeding as adults and only emerging for very brief periods on just a few days of the year (Britton 1978).

Characters (from Britton 1978): Anterior face of the clypeus deep (ratio greatest width: mid depth 3.5–4.8:1), bearing scattered setiferous punctures in the middle as well as towards the sides, anterior edge of the clypeus as seen from above not broadly bilobed; antennae 10-segmented, with a 3–7-lamellate club; head, pronotum, elytra and abdomen not densely and uniformly clothed with short setae, bare or with minute setae which lie largely within their punctures, or with long, sparse, irregularly distributed setae but without scales or flattened adpressed setae; sutures separating abdominal sternites 3–5 fainter in the middle than at the sides.

Antitrogus Burmeister 1855 HW92
    |--*A. tasmanicus Burmeister 1855 [incl. A. nigricornis Blackburn 1911] HW92
    |--A. adamsi Britton 1978 HW92
    |--A. adustus Britton 1978 HW92
    |--A. brooksi Britton 1978 HW92
    |--A. burmeisteri Burmeister 1911 HW92
    |--A. carnei Britton 1978 HW92
    |--A. circulifrons Britton 1978 HW92
    |--A. morbillosus (Blackburn 1898) [=Rhopaea morbillosa; incl. R. incognita Blackburn 1911] HW92
    |--A. mussoni (Blackburn 1892) (see below for synonymy) HW92
    |--A. nox Britton 1978 HW92
    |--A. parilis Britton 1978 HW92
    |--A. parvulus Britton 1978 HW92
    |--A. planiceps (Blackburn 1911) [=Rhopaea planiceps] HW92
    |--A. politus (Lea 1920) [=Rhopaea polita] W92
    |--A. robertsi Britton 1978 HW92
    |--A. setifer Britton 1978 HW92
    |--A. setifrons Britton 1979 HW92
    `--A. tumidifrons Britton 1978 HW92

Antitrogus mussoni (Blackburn 1892) [=Rhopaea mussoni; incl. R. consanguinea Blackburn 1911, R. dubitans Blackburn 1911, R. rugulosa Blackburn 1911] HW92

*Type species of generic name indicated

REFERENCES

Britton, E. B. 1978. A revision of the Australian chafers (Coleoptera: Scarabaeidae: Melolonthinae) vol. 2. Tribe Melolonthini. Australian Journal of Zoology, Supplementary Series 60: 1–150.

[HW92] Houston, W. W. K., & T. A. Weir. 1992. Melolonthinae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 174–358. AGPS Press: Canberra.

Last updated: 27 April 2021.

Abyssocottidae

Female Asprocottus korjakovi, copyright V. G. Sideleva.

Belongs within: Cottoidea.

The Abyssocottidae is a group of sculpins endemic to Lake Baikal, many of which live in deep water and exhibit associated adaptations such as reduced scalation and a lateral line of exposed neuromasts.

See also: The sculpins of Baikal.

Abyssocottidae [Cottocomephoridae, Cottocomephorinae]
     |--+--Leocottus kesslerii [=Cottus kesslerii] KKV03
     |  `--Cottocomephorus KKV03
     |       |--C. alexandrae KKV03
     |       |--C. grewingkii KKV03
     |       `--C. inermis KKV03
     |--+--Paracottus knerii KKV03
     |  `--Procottus KKV03
     |       |--P. gotoi KKV03
     |       |--P. gurwici KKV03
     |       |--P. jeittelesii KKV03
     |       `--P. major KKV03
     `--+--Comephorus Lacepède 1800 KKV03, B78 [=Comophorus Agassiz 1846 B78; Comephoridae]
        |    |--C. baicalensis KKV03
        |    `--C. dybowski KKV03
        `--Abyssocottinae KKV03
             |  i. s.: Neocottus KKV03
             |           |--N. thermalis KKV03
             |           `--N. werestschagini KKV03
             |--Cyphocottus KKV03
             |    |--C. eurystomus KKV03
             |    `--C. megalops KKV03
             `--+--+--‘Abyssocottus’ gibbosus KKV03
                |  `--+--Batrachocottus KKV03
                |     |    |--B. baicalensis KKV03
                |     |    |--B. multiradiatus KKV03
                |     |    |--B. nikolskii KKV03
                |     |    `--B. talievi KKV03
                |     `--Limnocottus KKV03
                |          |--+--L. bergianus KKV03
                |          |  `--L. pallidus KKV03
                |          `--+--L. godlewskii KKV03
                |             `--L. griseus KKV03
                `--+--+--Cottinella boulengeri KKV03
                   |  `--Abyssocottus KKV03
                   |       |--A. elochini KKV03
                   |       `--A. korotneffi KKV03
                   `--Asprocottus KKV03
                        |  i. s.: A. abyssalis KKV03
                        |         A. parmiferus KKV03
                        |--+--A. herzensteini KKV03
                        |  `--A. platycephalus KKV03
                        `--+--A. korjakovi KKV03
                           |    |--A. k. korjakovi KKV03
                           |    `--A. k. minor KKV03
                           `--A. pulcher KKV03

*Type species of generic name indicated

REFERENCES

[B78] Britton, E. B. 1978. A revision of the Australian chafers (Coleoptera: Scarabaeidae: Melolonthinae) vol. 2. Tribe Melolonthini. Australian Journal of Zoology, Supplementary Series 60: 1–150.

[KKV03] Kontula, T., S. V. Kirilchik & R. Väinölä. 2003. Endemic diversification of the monophyletic cottoid fish species flock in Lake Baikal explored with mtDNA sequencing. Molecular Phylogenetics and Evolution 27: 143–155.

Nemastoma

Nemastoma sp., from Axel L. Schönhofer.

Belongs within: Nemastomatidae.

Nemastoma is a widespread genus of small harvestmen in Europe.

Characters (from Martens 1978): Male genital morphology with truncus penis long and narrow, usually almost at right angles to the dilated base; Glans in a straight extension of trunk, scarcely wider than latter; dorsoventrally flattened. Glans symmetrical or slightly asymmetrical, laterally with one to eight strong, basad-directed hooks, exceptionally missing. Scutum primarily a scutum magnum, old and strongly sclerotized individuals with scutum compositum (abdominal tergites VI, VII and rarely VIII fused with scutum magnum). Smooth-shiny to roughly granulated, without dorsal spines; deep black, carapace often with gold or silver markings.

<==Nemastoma Koch 1836 [incl. Lugubrostoma Kratochvíl 1958, Stridulostoma Hadži 1973] S13
    |--*N. bimaculatum (Fabricius 1775) M06 (see below for synonymy)
    |--N. bidentatum Roewer 1914 M06 [=N. (Lugubrostoma) bidentatum N05]
    |    |--N. b. bidentatum M06
    |    |--N. b. pluridentatum Hadži 1973 (see below for synonymy) S13
    |    |--N. b. relictum Gruber & Martens 1968 M06
    |    `--N. b. sparsum Gruber & Martens 1968 M06
    |--N. daciscum Koch 1869 (n. d.) S13
    |--N. dentigerum Canestrini 1873 M06
    |--‘Crosbycus’ goodnighti Roewer 1951 (n. d.) S10, S13 [=C. goodnightorum (l. c.) S13]
    |--N. incertum Koch & Berendt 1854 D07
    |--N. lilliputanum (Lucas 1849) [=Goniosoma lilliputanum] S13
    |--N. lugubre (Müller 1776) M06 (see below for synonymy)
    |--N. rude Simon 1881 D04
    |--N. schuelleri Gruber & Martens 1968 M06
    |--N. superbum Koch 1869 M06
    |--N. transsylvanicum Gruber & Martens 1968 M06
    `--N. triste (Koch 1835) M06 (see below for synonymy)

Nemastoma bidentatum pluridentatum Hadži 1973 [=N. (Lugubrostoma) triste pluridentatum; incl. N. (*Stridulostoma) seliskari Hadži 1973] S13

*Nemastoma bimaculatum (Fabricius 1775) M06 [=Phalangium bimaculatum M06, N. lugubre bimaculatum D04, Opilio bimaculatus D04]

Nemastoma lugubre (Müller 1776) M06 [=Phalangium lugubre D04, Histricostoma lugubre D04, Nemastoma (*Lugubrostoma) lugubre S13; incl. N. lugubre helvetica Lessert 1917 S13, N. lugubre var. striatum Morin 1934 S13]

Nemastoma triste (Koch 1835) M06 [=Phalangium triste R23, N. (Lugubrostoma) triste S13; incl. N. kollari Doleschall 1852 R23, N. lugubre unicolor Roewer 1914 S13]

*Type species of generic name indicated

REFERENCES

[D04] Delfosse, E. 2004. Catalogue préliminaire des Opilions de France métropolitaine (Arachnida Opiliones). Bulletin de Phyllie 20: 34–58.

[D07] Dunlop, J. A. 2007. Paleontology. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 247–265. Harvard University Press: Cambridge (Massachusetts).

Martens, J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. Gustav Fischer Verlag: Jena.

[M06] Martens, J. 2006. Weberknechte aus dem Kaukasus: Nemastomatidae (Arachnida, Opiliones, Nemastomatidae). Senckenbergiana Biologica 86 (2): 145–210.

[N05] Novak, T. 2005. An overview of harvestmen (Arachnida: Opiliones) in Bosnia and Herzegovina. Natura Croatica 14 (4): 301–350.

[R23] Roewer, C.-F. 1923. Die Weberknechte der Erde: Systematische Bearbeitung der bisher bekannnten Opiliones. Gustav Fischer: Jena.

[S13] Schönhofer, A. L. 2013. A taxonomic catalogue of the Dyspnoi Hansen and Sørensen, 1904 (Arachnida: Opiliones). Zootaxa 3679 (1): 1–68.

[S10] Shear, W. A. 2010. Hesperonemastoma smilax, n. sp., a remarkable new harvestman from a cave in West Virginia, with comments on other reported cave-dwelling Hesperonemastoma species (Opiliones, Ischyropsalidoidea, Sabaconidae). Journal of Cave and Karst Studies 72 (2): 105–110.

Paranemastoma

Paranemastoma sp., from Axel L. Schönhofer.

Belongs within: Nemastomatidae.

Paranemastoma is a genus including some of the largest nemastomatid harvestmen, found in central and eastern Europe.

Characters (from Martens 1978): Male genital morphology with truncus penis long and extremely narrow, base widened, markedly separated from the rest of the trunk, but not angled; deeply divided, the two muscular strands concentrated in the two halves, glans in a straight extension of truncus, scarcely wider than latter, differentiated only by stronger sclerotization and tiny sensory bristles. Tip of glans divided into two branches, the right branch (in ventral view) narrower and slightly overarching the shorter and wider left branch. Scutum with para-median spine pairs, most developed on abdominal area II, weaker in female. Body always deep black, usually with gold or silver spotting.

<==Paranemastoma Redikorzev 1936 [incl. Buresiolla Kratochvíl 1958, Dromedostoma Kratochvíl 1958] S13
    |--*P. superbum Redikorzev 1936 M11 (see below for synonymy)
    |--P. aeginum (Roewer 1951) [=Nemastoma aeginum] S13
    |--P. amseli (Roewer 1951) [=Nemastoma amseli] S13
    |--P. amuelleri (Roewer 1951) [=Nemastoma amuelleri] S13
    |--P. ancae Avram 1973 S13
    |--P. armatum (Kulczyński 1909) [=Nemastoma quadripunctatum armatum] N05
    |--P. aurigerum (Roewer 1951) [=Nemastoma aurigerum] S13
    |    |--P. a. aurigerum S13
    |    |--P. a. joannae Staręga 1976 S13
    |    `--P. a. ryla (Roewer 1951) [=Nemastoma ryla, N. (Dromedostoma) ryla] S13
    |--P. aurosum (Koch 1869) N05 [=Nemastoma quadripunctatum aurosum N05, N. quadripunctatum var. aurosa D04]
    |--P. beroni Mitov 2011 M11
    |--P. bicuspidatum (Koch 1835) (see below for synonymy) S13
    |--P. bolei (Hadži 1973) [=Nemastoma (Dromedostoma) bolei] S13
    |--P. brevipalpatum (Roewer 1951) [=Nemastoma brevipalpatum] S13
    |--P. bureschi (Roewer 1926) (see below for synonymy) S13
    |--P. caporiaccoi (Roewer 1951) [=Nemastoma caporiaccoi] S13
    |--P. carneluttii (Hadži 1973) [=Nemastoma carneluttii] S13
    |--P. corcyraeum (Roewer 1917) [=Nemastoma quadripunctatum corcyraeum] S13
    |--P. emigratum (Roewer 1959) [=Nemastoma emigratum] S13
    |--P. ferkeri (Roewer 1951) [=Nemastoma ferkeri] S13
    |--P. filipes (Roewer 1917) [=Nemastoma quadripunctatum var. filipes, N. (Dromedostoma) filipes] S13
    |--P. gostivarense (Hadži 1973) [=Nemastoma gostivarense] S13
    |--‘Crosbycus’ graecus Giltay 1932 (see below for synonymy) S13
    |--P. ikarium (Roewer 1951) [=Nemastoma ikarium] S13
    |--P. ios (Roewer 1917) [=Nemastoma quadripunctatum ios] S13
    |--P. iranicum Martens 2006 M06
    |--P. kaestneri (Roewer 1951) [=Nemastoma kaestneri] S13
    |--P. kalischevskyi (Roewer 1951) (see below for synonymy) S13
    |--P. kochii (Nowicki 1870) (see below for synonymy) S13
    |--P. longipalpatum (Roewer 1951) [=Nemastoma longipalpatum] S13
    |--P. longipes (Schenkel 1947) N05 (see below for synonymy)
    |--P. macedonicum (Hadži 1973) [=Nemastoma macedonicum] S13
    |--P. machadoi (Roewer 1951) [=Nemastoma machadoi] S13
    |--P. mackenseni (Roewer 1923) [=Nemastoma mackenseni] S13
    |--P. multisignatum (Hadži 1973) [=Nemastoma (Dromedostoma) multisignatum] S13
    |--P. ‘nigrum’ (Hadži 1973) [=Nemastoma nigrum non N. paspalevi var. nigrum Kratochvíl 1958] S13
    |--P. perfugium (Roewer 1951) [=Nemastoma perfugium] S13
    |--P. quadripunctatum (Perty 1833) M06 (see below for synonymy)
    |--P. radewi (Roewer 1926) (see below for synonymy) S13
    |--P. roeweri Starega 1978 [=Nemastoma redikorzevi Roewer 1951 non Charitonov 1946, P. redikorzevi] S13
    |--P. santorinum (Roewer 1951) [=Nemastoma santorinum] S13
    |--P. senussium (Roewer 1951) [=Nemastoma senussium] S13
    |--P. sillii (Herman 1871) S13
    |    |--P. s. sillii S13 (see below for synonymy)
    |    |--P. ‘gigas’ montenegrinum Nosek 1904 S13 [=Nemastoma gigas montenegrinum S13, N. g. montenegrina (l. c.) R23]
    |    `--P. s. monticola Babalean 2011 S13
    |--P. sketi (Hadži 1973) [=Nemastoma (Dromedostoma) sketi] S13
    |--P. spinosulum (Koch 1869) [=Nemastoma spinosulum, N. spinulosum (l. c.)] S13
    |--P. thessalum (Simon 1885) [=Nemastoma quadripunctatum thessalum, P. teshalum (l. c.)] S13
    |--P. titaniacum (Roewer 1914) S13 (see below for synonymy)
    |--P. umbo (Roewer 1951) [=Nemastoma umbo] M06
    `--P. werneri (Kulczyński 1903) [=Nemastoma quadripunctatum werneri, P. quadripunctatum werneri] S13

‘Crosbycus’ graecus Giltay 1932 [incl. Nemastoma simplex Giltay 1932 non N. bacilliferum simplex Simon 1913, Paranemastoma simplex] S13

Paranemastoma bicuspidatum (Koch 1835) [=Phalangium bicuspidatum, Nemastoma quadripunctatum bicuspidatum; incl. N. reimoseri Roewer 1951] S13

Paranemastoma bureschi (Roewer 1926) [=Nemastoma bureschi, Crosbycus bureschi, Paranemastoma (*Buresiolla) bureschi; incl. N. tunetanum Roewer 1951, B. tunetanum, B. tunetorum (l. c.)] S13

Paranemastoma kalischevskyi (Roewer 1951) [=Nemastoma kalischevskyi; incl. N. charitonovi Mcheidze 1952, Paranemastoma charitonovi, N. suchumium Roewer 1951, P. suchumium] S13

Paranemastoma kochii (Nowicki 1870) [=Nemastoma quadripunctatum kochii; incl. N. hungaricum Roewer 1951, N. nicaeum Roewer 1951] S13

Paranemastoma longipes (Schenkel 1947) N05 [=Nemastoma gigas longipes M00, N. (Dromedostoma) longipes N05; incl. N. tenue Hadži 1973 S13]

Paranemastoma quadripunctatum (Perty 1833) M06 [=Phalangium quadripunctatum D04, Nemastoma quadripunctatum N05; incl. N. alfkeni Roewer 1951 D04, Paranemastoma alfeki D04, N. armeniacum Roewer 1951 S13, P. armeniacum M06, N. austriacum Roewer 1951 D04, N. quadripunctatum carniolicum Hadži 1973 D04, N. chaucicum Roewer 1951 D04, Paranemastoma chaucicum D04, N. emonense Hadži 1973 D04, Phalangium flavimanum Koch 1835 K35, Nemastoma flavimanum D04, Opilio flavimanum D04, N. gertschi Roewer 1951 D04, Paranemastoma gertschi D04, Crosbycus lengersdorfi Roewer 1951 D04, Nemastoma quadripunctatum var. lessinensis Caporiacco 1940 S13, N. mediosignatum Hadži 1973 D04, N. quadripunctatum var. moesiacum Roewer 1917 S13, Paranemastoma moesiacum N05, N. nervosum Roewer 1923 S13, N. slovenicum Hadži 1973 D04, N. (Dromedostoma) strasseri Roewer 1951 N05, N. triglavense Hadži 1973 D04]

Paranemastoma radewi (Roewer 1926) [=Nemastoma radewi S13, N. (*Dromedostoma) radewi S13, N. radevi (l. c.), Paranemastoma radevi; incl. N. (D.) atanasovi Kratochvíl 1958, P. atanasovi, N. (D.) atanasovi balcanica Kratochvíl 1958, N. insulare Roewer 1951, N. (D.) markovi Kratochvíl 1958, N. (D.) megarae Hadži 1973, N. (D.) paspalevi var. nigrum Kratochvíl 1958, N. (D.) paspalevi Kratochvíl 1958, Paranemastoma paspalevi, N. polonicum Roewer 1951, N. riparium Roewer 1951, n. schenkeli Roewer 1951] S13

Paranemastoma sillii sillii (Herman 1871) S13 [=Nemastoma quadripunctatum sillii S13; incl. N. bishopi Roewer 1951 S13, N. bulgaricum Roewer 1951 S13, N. carpathicum Roewer 1951 S13, Crosbycus cavernae Roewer 1951 S13, N. gigas Sørensen in Lendl 1894 S13, N. sillii gigas, Paranemastoma gigas S13, N. lineatum Sørensen in Lendl 1894 S13, N. quadripunctatum lineatum, N. mohilevum Roewer 1951 S13, N. politum Roewer 1951 S13, N. romanium Roewer 1951 S13, N. rumanicum (l. c.) S13, N. rumelium Roewer 1951 S13, N. wiehlei Roewer 1951 S13, N. zilchi Roewer 1951 S13]

*Paranemastoma superbum Redikorzev 1936 non Nemastoma superbum Koch 1869 M11 [=P. supersum Roewer 1951 M06, Nemastoma supersum M06; incl. N. superbum bacuriana Mcheidze 1959 M06, N. superbum bacurianum M06]

Paranemastoma titaniacum (Roewer 1914) S13 [=Nemastoma quadripunctatum titaniacum N05, N. (Dromedostoma) titaniacum S13; incl. N. acrospinosum Roewer 1951 S13, N. bimaculosum Roewer 1951 S13, Paranemastoma bimaculosum N05, N. bipunctatum Hadži 1973 S13, N. (D.) grabrovicae Hadži 1973 N05, N. gabbrovicae (l. c.) N05, N. acrospinosum pretneri Hadži 1973 S13]

*Type species of generic name indicated

REFERENCES

[D04] Delfosse, E. 2004. Catalogue préliminaire des Opilions de France métropolitaine (Arachnida Opiliones). Bulletin de Phyllie 20: 34–58.

[K35] Koch, C. L. 1835. Arachniden. In: Herrich-Schäffer, G. A. W. Deutschlands Insecten. Band 34, Hefte 128–133. F. Pustet: Regensburg.

Martens, J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. Gustav Fischer Verlag: Jena.

[M06] Martens, J. 2006. Weberknechte aus dem Kaukasus: Nemastomatidae (Arachnida, Opiliones, Nemastomatidae). Senckenbergiana Biologica 86 (2): 145–210.

[M00] Mitov, P. 2000. Contribution to the knowledge of the harvestmen (Arachnida: Opiliones) of Albania. Ekológia 19 (Suppl. 3): 159–169.

[M11] Mitov, P. G. 2011. A new anophthalmous species of Paranemastoma from Bulgaria (Opiliones: Nemastomatidae). Journal of Arachnology 39 (2): 303–319.

[N05] Novak, T. 2005. An overview of harvestmen (Arachnida: Opiliones) in Bosnia and Herzegovina. Natura Croatica 14 (4): 301–350.

[R23] Roewer, C.-F. 1923. Die Weberknechte der Erde: Systematische Bearbeitung der bisher bekannnten Opiliones. Gustav Fischer: Jena.

[S13] Schönhofer, A. L. 2013. A taxonomic catalogue of the Dyspnoi Hansen and Sørensen, 1904 (Arachnida: Opiliones). Zootaxa 3679 (1): 1–68.