A taxonomic and phylogenetic revision of Amarochara Thomson. I. The species of the Holarctic region (Coleoptera: Staphylinidae, Aleocharinae, Oxypodini).

Abstract

The types and additional material of the Amarochara species occurring in the Holarctic region are revised. 25 valid species are recognized, 24 of them (re-)described: A. umbrosa (Erichson), A. heterogaster Cameron, A. sororcula Cameron, A. inquilina (Casey), A. fenyesi Blatchley, A. brevios sp. n., A. bonnairei (Fauvel), A. cribripennis (Mulsant & Rey), A. siculifera sp. n., A. inermis sp. n., A. caeca sp. n., A. crassicornis (Quedenfeldt), A. forticornis (Lacordaire), A. carinata sp. n., A. loebli Pace, A. seriepunctata sp. n., A. armata sp. n., A. wrasei sp. n., A. megalops sp. n., A. formosana sp. n., A. densepunctata sp. n., and A. splendens Jarrige. The identity of A. tingitana Jarrige, whose holotype is lost, remains doubtful. The following synonymies, new combinations, and replacement name are established or confirmed, respectively: Calodera Mannerheim, 1830 = Nasirema Casey, 1893, syn. n.; Aleochara Gravenhorst, 1802 = Sorecocephala Bernhauer, 1902, syn. n.; Amarochara heterogaster Cameron, 1939 = A. simlaensis Cameron, 1939, syn. n., = A. smetanai Pace, 1992, syn. n.; Calodera bonnairei Fauvel, 1865, nomen protectum = Oxypoda glabriventris Rye, 1865, nomen oblitum; Dasygnypeta velata (Erichson, 1837) = Calodera flavipes Motschulsky, 1858 (previously a synonym of Amarochara forticornis); Alevonota japonica (Cameron, 1933), comb. n. (from Amarochara); Calodera caseyi nom. n., comb. n. (originally described as Nasirema humilis Casey, 1893, previously attributed to Amarochara, and a secondary junior homonym of Calodera humilis Erichson); Calodera parviceps (Casey, 1893), comb. n. (originally in Nasirema, previously a presumed synonym of Amarochara umbrosa); Pseudocalea korbi (Bernhauer, 1902), comb. n. (from Amarochara); Aleochara (Ceranota) ocaleoides (Bernhauer, 1902), comb. n. (from Amarochara); Aleochara (Ceranota) subtumida (Hochhuth, 1849) = Calodera brunnea Motschulsky, 1860 (previously a synonym of Amarochara forticornis), = Ocalea reitteri Bernhauer, 1900, syn. n. Amarochara flavicornis Bernhauer, 1907 is excluded from Amarochara and attributed to the Athetini, but of doubtful generic affiliations. A neotype is designated for Calodera bonnairei Fauvel. Lectotypes are designated for Calodera umbrosa Erichson, 1837, Amarochara heterogaster Cameron, 1939, Nasirema inquilina Casey, 1906, Oxypoda glabriventris Rye, 1865, Ilyobates cribripennis Mulsant & Rey, 1875, and Ocalea reitteri Bernhauer, 1900. The taxonomic history of Amarochara is outlined. Based on a phylogenetic analysis, the prevailing subgeneric concept is not adopted. As the species of zoogeographic regions other than the Holarctic have not yet been revised, however, the subgenera are not formally synonymized. Instead, the Holarctic species of the genus are attributed to five species groups. The (re-)descriptions of the genus, the species groups, and the species are complemented by a key and by figures of the mouthparts, primary and secondary sexual characters, and other distinguishing characters. The biogeographic data available are compiled and the distributions of several species are mapped. As can be inferred from field studies and collection data, species of Amarochara are univoltine, with reproduction taking place in spring and summer. Apparently, reproduction and hibernation take place in an unknown subterranean habitat. Ecological data are compiled and discussed, and the phenologies of the better-known species are figured. The genitalia of Pseudocalea korbi (Bernhauer) and Aleochara ocaleoides (Bernhauer) are illustrated.

Keywords

Coleoptera, Staphylinidae, Aleocharinae, Oxypodini, Amarochara, Nasirema, Calodera, Ocalea, Pseudocalea, Aleochara, Alevonota, Dasygnypeta, Sorecocephala, Holarctic region, Palaearctic region, Nearctic region, taxonomy, biogeography, ecology, life history, new species, new synonymy, new combination, neotype designation, lectotype designation.

Nomenclatural Acts

ocaleoides (Bernhauer, 1902) (Aleochara (Ceranota)), comb. n. hitherto Amarochara ocaleoides
japonica (Cameron, 1933) (Alevonota), comb. n. hitherto Amarochara (Amarochara) japonica
armata Assing, 2002 (Amarochara), spec. n.
bonnairei (Fauvel, 1865) (Amarochara), Neotype; nom. protectum hitherto Calodera (Ilyobates) bonnairei
brevios Assing, 2002 (Amarochara), spec. n.
caeca Assing, 2002 (Amarochara), spec. n.
carinata Assing, 2002 (Amarochara), spec. n.
cribripennis (Mulsant & Rey, 1875) (Amarochara), Lectotype described as Ilyobates cribripennis
densepunctata Assing, 2002 (Amarochara), spec. n.
formosana Assing, 2002 (Amarochara), spec. n.
heterogaster Cameron, 1939 (Amarochara), Lectotype
inermis Assing, 2002 (Amarochara), spec. n.
inquilina (Casey, 1906) (Amarochara), Lectotype described as Nasirema inquilina
megalops Assing, 2002 (Amarochara), spec. n.
seriepunctata Assing, 2002 (Amarochara), spec. n.
siculifera Assing, 2002 (Amarochara), spec. n.
umbrosa (Erichson, 1837) (Amarochara), Lectotype described as Calodera umbrosa
wrasei Assing, 2002 (Amarochara), spec. n.
simlaensis Cameron, 1939 (Amarochara (Lasiochara)), syn. n. of Amarochara heterogaster Cameron, 1939
smetanai Pace, 1992 (Amarochara (Lasiochara)), syn. n. of Amarochara heterogaster Cameron, 1939
caseyi Assing, 2002 (Calodera), nom. n. pro Nasirema humilis Casey, 1893, nec Erichson, 1837
parviceps (Casey, 1893) (Calodera), comb. n. hitherto Nasirema parviceps
reitteri Bernhauer, 1900 (Ocalea), Lectotype; syn. n. of Aleochara (Ceranota) subtumida (Hochhuth, 1849)
glabriventris Rye, 1865 (Oxypoda), Lectotype; nom. oblitum now a synonym of Amarochara bonnairei (Fauvel, 1865): nom. protectum
korbi (Bernhauer, 1902) (Pseudocalea), comb. n. hitherto Amarochara korbi