Aphodiinae

European dung beetle Aphodius fimetarius, copyright James K. Lindsey.

Belongs within: Scarabaeidae.
Contains: Ataenius, SaprositesRhyssemus, Aphodius (Podotenus).

The Aphodiinae are a cosmopolitan group of mostly relatively small, elongate scarabaeids. Species of the tribe Aphodiini are mostly dung feeders and some have become very widespread in association with farmed livestock. Members of other tribes are commonly saprophagous (Cassis & Weir 1992).

Characters (from Cassis & Weir 1992): Size from 2 to 10 mm; tarsi of equal length; elytra covering abdomen, without narrow margin; abdominal spiracles located in pleural membrane, covered by elytra; midcoxae strongly oblique, separated by less than their width; scutellum present (occasionally minute to not visible); labrum concealed or only slightly exposed; hind tibia usually with two apical spurs (rarely reduced in size or only one). Larva with body not hump-backed; head capsule dark brown; antennae four- or apparently five-segmented with reduced apical segment, at most half as wide as penultimate; galea and lacinia distinctly separated; anal opening usually surrounded by fleshy lobes; legs five-segmeted with well-developed claw.

Aphodiinae [Aphodiidae, Aphodiides]
    |  i. s.: Saprus Blackburn 1904CW92
    |           `--*S. griffithi Blackburn 1904CW92
    |--Odochilus [Odochilini]CW92
    |--Didactylia d’Orbigny 1896 [Didactyliina, Didactyliini]RJS02
    |    |--D. knausiiRJS02
    |    `--D. parcusRJS02
    |--EupariiniCW92
    |    |--AtaeniusCW92
    |    |--SaprositesCW92
    |    |--Annegialia Howden 1971RJS02
    |    |    `--A. ataeniformisRJS02
    |    |--Aphotaenius Cartwright 1952RJS02
    |    |    `--A. carolinusRJS02
    |    |--Martineziella Chalumeau 1986 [incl. Martinezia Chalumeau 1983]RJS02
    |    |    `--M. dutertreiRJS02
    |    |--Euparixia Brown 1927RJS02
    |    |    |--E. duncaniRJS02
    |    |    `--E. moseriRJS02
    |    |--Parataenius Balthasar 1961CW92
    |    |    |--*P. mirabilis Balthasar 1961CW92
    |    |    |--P. granuliceps Petrovitz 1971CW92
    |    |    `--P. simulator (Harold 1868)RJS02, CW92 [=Ataenius simulatorRJS02]
    |    |--Euparia LePeletier & Serville 1828RJS02
    |    |    |--*E. castanea Saint-Fargeau & Serville 1828CW92
    |    |    |--E. olliffi Blackburn 1904CW92
    |    |    `--E. squamosa Lea 1923CW92
    |    `--Pseudataenius Brown 1927RJS02
    |         |--P. contortusRJS02
    |         |--P. socialisRJS02
    |         `--P. waltherhorniRJS02
    |--Psammodiini [Psammodiina, Rhyssemina]CW92
    |    |--RhyssemusRJS02
    |    |--Phycocus Broun 1886CW92
    |    |    `--*P. graniceps Broun 1886CW92
    |    |--Pleurophorus Mulsant 1842RJS02
    |    |    `--P. caesusRJS02
    |    |--Geopsammodius Gordon & Pittino 1992RJS02
    |    |    |--G. hydropicusRJS02
    |    |    |--G. relictillusRJS02
    |    |    `--G. sabinaeRJS02
    |    |--Odontopsammodius Gordon & Pittino 1992RJS02
    |    |    |--O. armaticepsRJS02
    |    |    |--O. bidensRJS02
    |    |    `--O. cruentusRJS02
    |    |--Trichiorhyssemus Clouët 1901CW92
    |    |    |--*T. riparius (Horn 1871) [=Rhyssemus riparius]CW92
    |    |    |--T. alternatusRJS02
    |    |    |--T. cristatellusRJS02
    |    |    `--T. hirsutus Clouët 1901CW92
    |    |--Diastictus Mulsant 1842CW92
    |    |    |--*D. vulneratus (Sturm 1805) [=Aphodius vulneratus; incl. D. sabuleti Mulsant 1842]CW92
    |    |    |--D. obscurior (Blackburn 1904) [=Psammodius obscurior]CW92
    |    |    |--D. pachypus (Lea 1923) [=Psammodius pachypus]CW92
    |    |    `--D. tibialis (Fabricius 1798) [=Aphodius tibialis]CW92
    |    |--Psammodius Fallén 1807 [incl. Psammobius Heer 1841]RJS02
    |    |    |--P. basalisRJS02
    |    |    |--P. laevipennisRJS02
    |    |    |--P. pierottiRJS02
    |    |    `--‘Psammobius’ sabulosus [incl. Pleurophorus ovipennis]B89
    |    |--Tesarius Rakovič 1981CW92
    |    |    |--*T. sulcipennis (Lea 1904) [=Phycocus sulcipennis]CW92
    |    |    |--T. caelatusRJS02
    |    |    |--T. doyeniRJS02
    |    |    |--T. mcclayiRJS02
    |    |    `--T. oregonensisRJS02
    |    |--Neopsammodius Rakovic 1986RJS02
    |    |    |--N. blandusRJS02
    |    |    |--N. interruptusRJS02
    |    |    |--N. mimeticusRJS02
    |    |    |--N. quinqueplicatusRJS02
    |    |    `--N. werneriRJS02
    |    |--Aphodopsammobius Endrödi 1964CW92
    |    |    |--*A. mirabilis (Endrödi 1964) [=Aphodius mirabilis]CW92
    |    |    |--A. australicus (Blackburn 1904) [=Psammodius australicus; incl. Psammobius densepunctatus Petrovitz 1964]CW92
    |    |    |--A. demarzi (Petrovitz 1964) [=Psammobius demarzi]CW92
    |    |    |--A. matthewsi Rakovič 1981CW92
    |    |    |--A. rugicollis (Macleay 1888) [=Ammaecius rugicollis]CW92
    |    |    `--A. zietzi (Blackburn 1895) [=Ataenius zietzi]CW92
    |    |--Platytomus Mulsant 1842RJS02
    |    |    |--P. atlanticusRJS02
    |    |    |--P. caelicollisRJS02
    |    |    |--P. longulusRJS02
    |    |    |--P. microsRJS02
    |    |    |--P. notialisRJS02
    |    |    `--P. tibialisRJS02
    |    `--Leiopsammodius Rakovič 1981CW92
    |         |--*L. laevicollis (Klug 1845) [=Psammodius laevicollis]CW92
    |         |--L. aceiRJS02
    |         |--L. deyrupiRJS02
    |         |--L. freyi (Petrovitz 1961) [=Psammobius freyi]CW92
    |         |--L. implicatus (Schmidt 1925) [=Psammobius implicatus]CW92
    |         |--L. litoralis (Lea 1923) [=Psammodius litoralis]CW92
    |         |--L. malkiniRJS02
    |         |--L. ocmulgeeiRJS02
    |         `--L. rufus (Rakovič 1981) [=Psammodius (Leiopsammodius) rufus]CW92
    `--AphodiiniCW92
         |--Hornietus Stebnicka 2000RJS02
         |    `--H. ventralis [=Saprosites ventralis]RJS02
         |--Dialytellus Brown 1929RJS02
         |    |--D. dialytoidesRJS02
         |    `--D. humeralisRJS02
         |--Oxyomus Laporte 1840 [incl. Xenoheptaulacus Hinton 1934]RJS02
         |    |--O. silvestrisRJS02
         |    `--O. striatocrenatus Fairmaire 1889F89
         |--Xeropsamobeus Saylor 1937RJS02
         |    |--X. ambiguusRJS02
         |    `--X. desertusRJS02
         |--Proctophanes Harold 1861 [=Proctammodes Blackburn 1892]CW92
         |    |--*P. sculptus (Hope 1847) [=Aphodius sculptus, *Proctammodes sculptus]CW92
         |    |--P. metasternalis (Lea 1923) [=Proctammodes metasternalis]CW92
         |    `--P. minor (Blackburn 1897) [=Proctammodes minor]CW92
         |--Dialytes Harold 1869RJS02
         |    |--*D. striatulus (Say 1825) [=Trox striatulus]CW92
         |    |--D. criddleiRJS02
         |    |--D. granifer Schmidt 1909CW92
         |    |--D. truncatusRJS02
         |    `--D. ulkeiRJS02
         |--Australammoecius Petrovitz 1958CW92
         |    |--*A. demarzi Petrovitz 1958CW92
         |    |--A. australis Petrovitz 1961CW92
         |    |--A. bicolor Petrovitz 1961CW92
         |    |--A. brunneus Petrovitz 1961CW92
         |    |--A. flavipennis Petrovitz 1963CW92
         |    `--A. granuliceps Petrovitz 1961CW92
         `--Aphodius Illiger 1798CW92
              |  i. s.: A. affinis Lucas 1846E12
              |         A. aterC01
              |         A. badipesRJS02
              |         A. bicrenulatus Lea 1923CW92
              |         A. bimaculatusR26
              |         A. bipustulatusR26
              |         A. brunneus Klug 1845G89
              |         A. concavusRJS02
              |         A. conspurcatusR26
              |         A. cribricollis Lucas 1846E12
              |         A. cynomysi Brown 1927S69
              |         A. desertusG89
              |         A. erraticusC01
              |         A. fossorC01
              |         A. haemorrhoidalisC01
              |         A. hamatus Say 1824A06
              |         A. hepaticus Roth 1851G89
              |         A. heterodoxus Lea 1923CW92
              |         A. hieroglyphicusG89
              |         A. hirtipennis Lucas 1846E12
              |         A. holdereriGE05
              |         A. immundusC01
              |         A. integrifrons Lea 1923CW92
              |         A. laevigatusF89
              |         A. luridipennis Muls 1851G89
              |         A. luridus [incl. A. luridus var. nigripes]C01
              |         A. melanostictusC01
              |         A. merdariusC01
              |         A. nigerC01
              |         A. nitidulusC01
              |         A. pardalis LeConte 1857A06
              |         A. parvoniger Lea 1923CW92
              |         A. plagiatus [incl. A. plagiatus var. concolor]C01
              |         A. prodromusC01
              |         A. pyriformisRJS02
              |         A. quadriguttatusC01
              |         A. quadrimaculatusR26
              |         A. rufipes (Linnaeus 1758)A06
              |         A. scutator (Herbst 1789)A06
              |         A. solskyiC01
              |         A. sordescensC01
              |         A. sordidus [incl. A. sordidus var. quadripunctatus]C01
              |         A. subterraneus [incl. A. subterraneus var. fuscipennis]C01
              |         A. sulcatusC01
              |         A. suturalis Lucas 1847E12
              |         A. thomomysi Brown 1928S69
              |         A. tricolor Lea 1923CW92
              |         A. tristisC01
              |         A. tuberifrons Fairmaire 1889F89
              |         A. unicolor Lucas 1847E12
              |--A. (Aphodius)RF04
              |    |--*A. (A.) fimetarius (Linnaeus 1758) [=Scarabaeus fimetarius]CW92
              |    `--A. (A.) conjugatus (Panzer 1785)RF04
              |--A. (Acrossidius Schmidt 1914)CW92
              |    |--A. (*A.) tasmaniae Hope 1847 (see below for synonymy)CW92
              |    |--A. (A.) pallidihirtus Balthasar 1941CW92
              |    `--A. (A.) pseudotasmaniae Given 1950CW92
              |--A. (Acrossoides Schmidt 1914)CW92
              |    `--A. (*A.) yorkensis Blackburn 1892CW92
              |--A. (Australaphodius Balthasar 1942)CW92
              |    `--A. (*A.) melbournicus Balthasar 1942CW92
              |--A. (Blackburneus Schmidt 1914)CW92
              |    |--A. (*B.) furcatus Schmidt 1909CW92
              |    `--A. (B.) albertisi Harold 1877CW92
              |--A. (Calamosternus Motschulsky 1860)CW92
              |    `--A. (*C.) granarius (Linnaeus 1767)CW92 (see below for synonymy)
              |--A. (Drepanocanthoides Schmidt 1914)CW92
              |    |--A. (D.) demarzi Petrovitz 1958CW92
              |    |--A. (D.) neglectus Schmidt 1912CW92
              |    `--A. (D.) walshi Horn 1870CW92
              |--A. (Liothorax) wilsonae Maté & Angus 2005F05
              |--A. (Melinopterus)C01
              |    |--A. (M.) mehelyi Csiki 1901C01
              |    |--A. (M.) roschlapili Csiki 1901C01
              |    `--A. (M.) semiluteusC01
              |--A. (Mendidius Harold 1868) [=Cnemargus Harold 1866 non Schönherr 1837]CW92
              |    |--A. (*M.) fimbriolatus Mannerheim 1849 [=*Cnemargus fimbriolatus]CW92
              |    |--A. (M.) burgaltaicus Csiki 1901C01
              |    |--A. (M.) diffidensC01
              |    `--A. (M.) distinguendus Schmidt 1910CW92
              |--A. (Nialus Mulsant & Rey 1870)CW92
              |    |--A. (*N.) varians Duftschmid 1805CW92
              |    |--A. (N.) lividus (Olivier 1789) (see below for synonymy)CW92
              |    `--A. (N.) pseudolividus Balthasar 1941CW92
              |--A. (Parammoecius Seidlitz 1890) [incl. Pseudammoecius Schmidt 1914]CW92
              |    |--A. (*P.) gibbus Germar 1817CW92
              |    `--A. (P.) castanicolor Motschulsky 1858 (see below for synonymy)CW92
              |--A. (Phaeaphodius Reitter 1892)CW92
              |    |--A. (*P.) rectus Motschulsky 1866CW92
              |    |--A. (P.) frenchi Blackburn 1892 [=A. ambiguus Boheman 1858 non Mulsant 1842]CW92
              |    `--A. (P.) zichyi Csiki 1901C01
              |--A. (Podotenus)CW92
              |--A. (Teuchestes) analisCW92
              |--A. (Trichaphodius Schmidt 1914)CW92
              |    |--A. (*T.) humilis Roth 1851CW92
              |    `--A. (T.) reichei Harold 1859CW92
              `--A. (Volinus Mulsant & Rey 1870)CW92
                   |--A. (*V.) lineolatus Illiger 1803CW92
                   `--A. (V.) candezei Harold 1868CW92

Aphodius (*Aphodius) tasmaniae Hope 1847 [incl. A. andersoni Blackburn 1904, A. australasiae Blanchard 1846 (nom. rej.), A. howitti Hope 1847, A. longitarsus Redtenbacher 1867]CW92

Aphodius (*Calamosternus) granarius (Linnaeus 1767)CW92 [=Scarabaeus granariusCW92; incl. A. adelaidae Hope 1847CW92, A. granarius var. guadeloupensis Fleutiaux & Sallé 1890FS90]

Aphodius (Nialus) lividus (Olivier 1789) [=Scarabaeus lividus; incl. A. cincticulus Hope 1847, A. spilopterus Germar 1848]CW92

Aphodius (Parammoecius) castanicolor Motschulsky 1858 [incl. A. australasiae Boheman 1858 non Blanchard 1846, A. (*Pseudammoecius) australasiae]CW92

*Type species of generic name indicated

References

[A06] Ahrens, D. 2006. The phylogeny of Sericini and their position within the Scarabaeidae based on morphological characters (Coleoptera: Scarabaeidae). Systematic Entomology 31: 113–144.

[B89] Bedel, L. 1889. Un lapsus. Annales de la Société Entomologique de France, 6e série 9: xxxiii.

[CW92] Cassis, G., & T. A. Weir. 1992. Aphodiinae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 81–105. AGPS Press: Canberra.

[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.

[F89] Fairmaire, L. 1889. Coléoptères de l’intérieur de la Chine. 5e partie. Annales de la Société Entomologique de France, 6e série 9: 5–84.

[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.

[FS90] Fleutiaux, E., & A. Sallé. 1890. Liste des coléoptères de la Guadeloupe et descriptions d’espèces nouvelles. Annales de la Société Entomologique de France, 6e série 9: 351–484.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso “Esploratore” dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 5–72.

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

[RF04] Rahola Fabra, P. 2004. Nuevos datos sobre Aphodius (Aphodius) conjugatus (Panzer, 1795) (Coleoptera, Aphodiidae) en España. Graellsia 60 (1): 125.

[RJS02] Ratcliffe, B. C., M. L. Jameson & A. B. T. Smith. 2002. Scarabaeidae Latreille 1802. In: Arnett, R. H., Jr, M. C. Thomas, P. E. Skelley & J. H. Frank (eds) American Beetles vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea pp. 39–81. CRC Press.

[R26] Risso, A. 1826. Histoire naturelle des principales productions de l’Europe méridionale et particulièrement de celles des environs de Nice et des Alpes maritimes vol. 5. F.-G. Levrault: Paris.

[S69] Steyskal, G. C. 1969. The mistreatment of the Latin genitive case in forming names of parasites. Systematic Zoology 18 (3): 339–342.

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