Passalinae

Proculus goryi, copyright Archivo Gráfico.

Belongs within: Scarabaeoidea.

The Passalinae are a pantropical group of bess beetles in which the procoxae do not project much below the prosternum, and the the head usually bears paired tubercles.

<==Passalinae CW92
    |--Didimus [Solenocyclini] BB16
    |    `--D. ealaensis BB16
    |--Serrulus [Macrolinini] BB16
    |    `--S. sinicus BB16
    |--Leptaulax Kaup 1868 CW92 [Leptaulacini BB16]
    |    |--*L. dentatus (Fabricius 1792) CW92 (see below for synonymy)
    |    |--L. bicolor (Fabricius 1801) [=Passalus bicolor] CW92
    |    `--L. timoriensis Mas86
    |--Proculini BB16
    |    |--Veturius BB16
    |    `--Proculus B14
    |         |--P. burmeisteri B14
    |         |--P. goryi (Melly 1833) B14
    |         |--P. jicaquei B14
    |         |--P. mniszechi B14
    |         |--P. opacipennis B14
    |         `--P. opacus B14
    `--Passalini BB16
         |--Spasalus BB16
         |    |--S. kaupi BB16
         |    `--S. seag BB16
         `--Passalus L02
              |--P. compergus Boisduval 1835 B35
              |--P. cornutus RD77 [=Nigidius cornutus B35]
              |--P. grandis B35
              |--P. indormitus BB16
              `--P. interruptus L02

Passalinae incertae sedis:
  Plesthenus quadricornis Mas86
  Eriocnemis lottinii Mas86
  Austropassalus Mjöberg 1917 [incl. Notocetius Hincks & Dibb 1935] CW92
    `--*A. hultgreni Mjöberg 1917 [incl. *Notocetius cornutus Hincks & Dibb 1935] CW92
  Protomocoelus Zang 1905 [=Pelops Kaup 1871 nec Gistl 1834 nec Koch 1836] CW92
    |--*P. australis (Boisduval 1835) [=Passalus australis, *Pelops australis] CW92
    `--‘Pelops’ gestroi Mac86
  Analaches Kuwert 1891 CW92
    |--*A. puberilis Kuwert 1891 CW92
    `--A. australiensis (Stoliczka 1873) [=Cetejus australiensis] CW92
  Labienus Kaup 1871 CW92
    |--*L. ptox (Kaup 1871) [=Eriocnemis ptox] CW92
    `--L. inaequalis Gravely 1918 [incl. L. trigonophorus proximus Hincks 1933] CW92
  Gonatas Kaup 1871 CW92
    |--*G. naviculator (Percheron 1844) [=Passalus naviculator] CW92
    |--G. germari (Kaup 1868) [=Aceraius germari] CW92
    |--G. minimus Kuwert 1891 CW92
    |--G. pumilio (Kaup 1868) [=Aceraius pumilio] CW92
    `--G. schellongi Kuwert 1891 [incl. G. albertisi Kuwert 1891] CW92
  Pharochilus Kaup 1868 CW92
    |--*P. dilatatus (Dalman 1817) (see below for synonymy) CW92
    |--P. nitidulus (Macleay 1871) [=Mastochilus nitidulus] CW92
    |--P. puncticollis (Macleay 1871) [=Mastochilus puncticollis] CW92
    |--P. punctiger (Gravely 1918) [=Mastochilus (Pharochilus) punctiger] CW92
    |--P. rugiceps (Hope & Westwood 1845) CW92 (see below for synonymy)
    `--P. scutellonotus Kuwert 1898 [incl. P. pararius Kuwert 1898] CW92
  Mastachilus Kaup 1868 [=Mastochilus Kaup 1868; incl. Episphenoides Kuwert 1891] CW92
    |--*M. polyphyllus (Macleay 1826) CW92 (see below for synonymy)
    |--M. australasicus (Percheron 1841) (see below for synonymy) CW92
    |--M. dilatatus [incl. M. chevrolati, M. crenistrius Boisd. 1835] Mas86
    |--M. lottinii (Boisduval 1835) [=Passalus lottinii] CW92
    |--M. parvifrons (Kuwert 1891) [=Episphenoides parvifrons] CW92
    |--M. politus Mas86
    |--M. quaestionis (Kuwert 1891) (see below for synonymy) CW92
    `--M. rigiceps Hope 1845 Mas86

*Leptaulax dentatus (Fabricius 1792) CW92 [=Passalus dentatus CW92; incl. L. quadridentatus Sturm 1826 Mac86]

Mastachilus australasicus (Percheron 1841) [=Passalus australasicus, *Episphenoides australasicus; incl. E. distans Kuwert 1898, Mastachilus macleayi Kaup 1868, E. mulleri Kuwert 1891, E. perinvitus Kuwert 1891] CW92

*Mastachilus polyphyllus (Macleay 1826) CW92 [=Passalus polyphyllus CW92; incl. Mastochilus abdominipunctatus Kuwert 1898 CW92, P. hexaphyllus Boisduval 1835 CW92, P. impressicollis Boheman 1858 CW92, Mastochilus impressicollis Mas86, P. sexdentatus Eschscholtz 1829 CW92]

Mastachilus quaestionis (Kuwert 1891) [=Episphenoides quaestionis; incl. Mastochilus capitalis Blackburn 1900] CW92

*Pharochilus dilatatus (Dalman 1817) [=Passalus dilatatus; incl. Tiberius caffer Kuwert 1891, Pa. cancrus Percheron 1844, Pa. chevrolati Percheron 1835, Pa. crenistrius Boisduval 1835] CW92

Pharochilus rugiceps (Hope & Westwood 1845) CW92 [=Passalus rugiceps CW92; incl. Ph. bilineatopunctatus Kuwert 1891 CW92, Ph. brevidentatus Kuwert 1898 CW92, Ph. compar Kuwert 1898 CW92, Passalus dilatatus Percheron 1835 non Dalman 1817 CW92, Ph. politus languidilabris Kuwert 1891 CW92, Pa. politus Burmeister 1847 CW92, Ph. politus B70]

*Type species of generic name indicated

REFERENCES

[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.

[BB16] Boucher, S., M. Bai, B. Wang, W. Zhang & X. Yang. 2016. Passalopalpidae, a new family from the Cretaceous Burmese amber, as the possible sister group of Passalidae Leach (Coleoptera: Scarabaeoidea). Cretaceous Research 64: 67–78.

[B70] Britton, E. B. 1970. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 495–621. Melbourne University Press.

[CW92] Cassis, G., & T. A. Weir. 1992. Passalidae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 20–29. AGPS Press: Canberra.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[Mac86] 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.

[Mas86] 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.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.