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

Die Typen und weiteres Material der in der Holarktis vorkommenden Arten der Gattung Amarochara Thomson werden revidiert. Von 25 als valid erkannten Arten werden 24 beschrieben bzw. redeskribiert: 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. und A. splendens Jarrige. Die Identitat von A. tingitana Jarrige, deren Holotypus verschollen ist, bleibt ungeklart. Die Revision ergab folgende Synonymien, Neukombinationen und Ersatznamen: 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 (zuvor Synonym von Amarochara forticornis); Alevonota japonica (Cameron, 1933), comb. n. (ursprunglich Amarochara); Calodera caseyi nom. n., comb. n. (ursprunglich als Nasirema humilis Casey, 1893 beschrieben und spater Amarochara zugeordnet, sekundares jungeres Synonym von Calodera humilis Erichson); Calodera parviceps (Casey, 1893), comb. n. (ursprunglich als Nasirema beschrieben, spater als Synonym von Amarochara umbrosa vermutet); Pseudocalea korbi (Bernhauer, 1902), comb. n. (ursprunglich Amarochara); Aleochara (Ceranota) ocaleoides (Bernhauer, 1902), comb. n. (ursprunglich Amarochara); Aleochara (Ceranota) subtumida (Hochhuth, 1849) = Calodera brunnea Motschulsky, 1860 (vorher Synonym von Amarochara forticornis), = Ocalea reitteri Bernhauer, 1900, syn. n. Amarochara flavicornis Bernhauer, 1907 wird in die Tribus Athetini transferiert, ihre Gattungszugehorigkeit ist jedoch zweifelhaft. Fur Calodera bonnairei Fauvel wird ein Neotypus designiert. Lectotypen werden designiert fur Calodera umbrosa Erichson, 1837, Amarochara heterogaster Cameron, 1939, Nasirema inquilina Casey, 1906, Oxypoda glabriventris Rye, 1865, Ilyobates cribripennis Mulsant & Rey, 1875 und Ocalea reitteri Bernhauer, 1900. In einem historischen Uberblick werden die taxonomischen Entwicklungen zur Gattung Amarochara zusammengefasst. Auf der Grundlage phylogenetischer Untersuchungen und Schlussfolgerungen wird die bestehende Untergattungssystematik nicht ubernommen. Da aber die Arten anderer zoogeographischer Regionen bisher nicht revidiert wurden, werden die Subgenera nicht formal synonymisiert. Statt dessen werden die holarktischen Vertreter der Gattung 5 Artengruppen zugeordnet. Die Beschreibungen der Gattung, der Artengruppen sowie der Arten werden durch eine Bestimmungstabelle und durch Abbildungen der Mundteile, der primaren und sekundaren Geschlechtsmerkmale sowie weiterer Unterscheidungsmerkmale erganzt. Die verfugbaren biogeographischen Daten werden zusammengefasst; fur eine Reihe von Arten werden Verbreitungskarten vorgelegt. Ergebnisse von Freilanduntersuchungen und Sammlungsdaten deuten darauf hin, dass die holarktischen Amarochara-Arten univoltin sind und sich im Fruhjahr und Sommer fortpflanzen. Reproduktion und Uberwinterung finden offenbar in einem bisher unbekannten unterirdischen Habitat statt. Die verfugbaren okologischen Daten werden zusammengefasst und diskutiert. Die Phanologien einiger besser bekannter Arten werden in Form von Diagrammen illustriert. Die Sexualmerkmale zweier in andere Gattungen kombinierter Arten, Pseudocalea korbi (Bernhauer) und Aleochara ocaleoides (Bernhauer), werden abgebildet.StichworterColeoptera, 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.Nomenklatorische Handlungenocaleoides (Bernhauer, 1902) (Aleochara (Ceranota)), comb. n. hitherto Amarochara ocaleoidesjaponica (Cameron, 1933) (Alevonota), comb. n. hitherto Amarochara (Amarochara) japonicaarmata Assing, 2002 (Amarochara), spec. n.bonnairei (Fauvel, 1865) (Amarochara), Neotype; nom. protectum hitherto Calodera (Ilyobates) bonnaireibrevios 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 cribripennisdensepunctata Assing, 2002 (Amarochara), spec. n.formosana Assing, 2002 (Amarochara), spec. n.heterogaster Cameron, 1939 (Amarochara), Lectotypeinermis Assing, 2002 (Amarochara), spec. n.inquilina (Casey, 1906) (Amarochara), Lectotype described as Nasirema inquilinamegalops 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 umbrosawrasei Assing, 2002 (Amarochara), spec. n.simlaensis Cameron, 1939 (Amarochara (Lasiochara)), syn. n. of Amarochara heterogaster Cameron, 1939smetanai Pace, 1992 (Amarochara (Lasiochara)), syn. n. of Amarochara heterogaster Cameron, 1939caseyi Assing, 2002 (Calodera), nom. n. pro Nasirema humilis Casey, 1893, nec Erichson, 1837parviceps (Casey, 1893) (Calodera), comb. n. hitherto Nasirema parvicepsreitteri 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. protectumkorbi (Bernhauer, 1902) (Pseudocalea), comb. n. hitherto Amarochara korbi

In

Introduction and taxonomic history
When TH O M SO N (1858) described Amarochara in a key to the Swedish subfamilies, tribes and genera of Staphylinidae, the genus included only the type species A. umbrosa (E RIC H -SON). M U LSAN T & R E Y (1875) gave a detailed redescription of both the genus and the species and described Mniobates, a subgenus of llyobates K RAATZ, 1856, to which they assigned Bolitochara forticornis LACORDAIRE (later to be designated type species of the subgenus), Calodera bonnairei FAUVEL, and llyobates cribripennis. The latter name was made available in the same work, although the authors, apparently unaware that it was an inlitteris name, attributed it to Fauvel. FO W LER (1888) synonymized A marochara by trans ferring C. umbrosa ERICHSO N to Calodera again and, with some doubt regarding the prio rity, regarded llyobates bonnairei (FA U V E L, 1865) as a junior synonym of llyobates glabriventris (R Y E , 1865), which had originally been described in the genus Oxypoda M A N -NERH EIM , 1830. Ó an G L B A U E R (1895) reversed this synonymy, reinstated A marochara as a distinct genus, transferred M niobates M U LSAN T & R E Y with the sole representative A. forticornis (LACORDAIRE) to A marochara as a subgenus, and established the new subgenus Casiochara for A. bonnairei (FA U V E L), which had previously been in llyobates. He distin guished the subgenera based on the presence of an anterior impression on the abdomi nal tergum VI and a carina on the mesosternum (.Mniobates), as well as on the presence and absence of genal carinae (Casiochara vend Amarochara s. str., respectively). BERNHAUER (1902) maintained this concept and presented a key to the three subgenera and seven Palaearctic species known at that time, with one species (A. umbrosa) in the nominal subgenus, four species (A. bonnairei, A. cribripennis, A. crassicornis (QUED EN FELD T), and the new A. ocaleoides) in Casiochara, and two species {A. forticornis and the n ew ti. korbi) in Mniobates. BERNHAUER & SCHEERPELTZ (1926) listed 14 valid species, 5 synonyms, and 4 subge nera of Amarochara, 1 species from the Ethiopian, 2 species and a subgenus from the Neotropical, 3 species and a synonym from the Nearctic, and the remainder from the Palaearctic region. Three of the Nearctic names (2 valid species and the synonym) had been attributed to the genus Nasirema C A SE Y , 1893 by CASEY (1893CASEY ( , 1906) (see also FENYES (1920)), which was regarded as a junior synonym of A marochara by BERN HAU ER & SCHEERPELTZ (1926). Since the publication of this catalogue the subgeneric con cept has remained the same and 19 further species have been described, two from the Western Palaearctic (A. tingitana] AWAGE, 1952 and A. plendens ] AK$AGA 1952), six from the Eastern Palaearctic (A. simlaensis CAM ERO N , 1939, A. heterogaster CAM ERO N , 1939, A. sororcula CAMERON, 1939, A. japónica CAM ERO N , 1933, A. loebliVACE, 1992, and smetanai PA C E , 1992), eight from the Ethiopian, and three species from the Oriental region. The present revision wT as initiated especially by the difficulties encountered when at tempting to identify A marochara material from the south of the Western Palaearctic region. With the exception of some recently described species from the Eastern Pala earctic, the Oriental, and the Ethiopian region, the sexual characters have not been stu died and the systematic status (including the prevailing subgeneric concept) has not been reexamined. Since one of the North America names in the genus has been treated as a junior synonym of a species described from the Western Palaearctic region, this first revisionary step covers both the Palaearctic and the Nearctic region.
In the course of the revision, a loan of the types o f A. tlngltana JAKRIGE and A. splendens JA R R IG E was attempted, but repeated requests remained unanswered. As an agreed dead line had to be met, the revision was submitted and the two taxa were treated as doubtful species. It was by chance that several weeks after submission of the manuscript Alexey Solodovnikov, St. Petersburg, informed me that he was planning to visit the MNHNP and kindly offered to use this opportunity to try and initiate a loan of the holotypes in question. His efforts were successful, so that it was possible to clarify at least the identity of M. splendens. Unfortunately, however, the holotype of A. tingitana was lost, so that an interpretation of that species will have to wait until material matching the description becomes available and a neotype can be designated. In order to avoid a separate supple ment on the identity of A. splendens and because the date for the publication of volume 51 was postponed due to a reorganisation of the publishing procedures, the editorial staff of the journal kindly offered to incorporate not only the changes and additions resulting from the unexpected availability of the type of that species, but also some further material that had been studied in the meantime. In view of deadlines to be met, however, A. plendens could not be considered in chapter 3, its redescription (including the figures) is added as a separate section at the end of chapter 4 (rather than in section 4.2), and the additional material (marked with an asterisk) is accounted for neither in the maps nor in the diagrams.

. M aterial and measurements
Types and additional material from the following institutions and private collections were examined:

The genus A m arochara THOMSON, 1858
Amarochara T H O M S O N , 1858: 32. Type species: Calodera umbrosa ERICHSON

Redescription
Species of small to intermediate size and of slender build. Microsculpture of head and pronotum absent or, if present, usually shallow and com posed of meshes of variable length and orientation, interstices of elytra in the vast majority of species without distinct microsculpture; abdomen very shining, anterior segments usually without microsculpture, posterior segments often with weakly pro nounced transverse microsculpture. Punctures on head and pronotum of variable den sity, moderately to extremely fine; puncturation on elytra and abdomen variable. Pube scence of forebody in most species relatively long and ± suberect, rarely short and/or decumbent. Head subquadrate, subcircular, or of + ovoid shape, slightly or distincdy narrower than pronotum; posteriorly with or without distinct neck; dorsal pubescence in lateral areas directed anteriad and in median area + transversely mediad or diagonally antero-mediad. Eyes moderately to very large, though in most species shorter than the postgenae in dorsal view, weakly to distinctly projecting from lateral outline of head. Genal carina present or reduced to various degrees. Gular sutures widely separated. Dorsal area of vertex not distinctly impressed. Antennae relatively long and massive; antennomere I dorso-apically distinctly emarginate, this emargination basad extending into a ± pronounced furrow (which accommoda tes the pedicel when the antenna is fully bent backwards); antennomeres I-III oblong, I DOI: 10.21248/contrib.entomol.52. 1.111-204 and II subequal in length; antennomere III coniform, somewhat shorter than I and II or of approximately equal length; IV and V strongly transverse, VI-X more or less trans verse (but less so than IV and V), at most only indistinctly increasing in width apically; XI approximately as long as the combined length of IX and X; pubescence of I-III relatively sparse and moderately short, that of IV and V dense and short, and that of VI-XI extremely short, fine, and very dense (Figs. 5,37,50,57,66,69,77,90,110). Penultimate joint of maxillary palpi very long and slender, apical joint long, slender, and weakly coniform (Figs. 1,18,34,49,61,76,89,109); mandibles apically distinctly bent and acute, right mandible without distinct molar tooth, faintly serrate or smooth in dorsal molar region (Fig. 4); ligula weakly to distincdy bifid, apically often with a fine and short, sometimes barely noticeable sensillum (Figs. 2,17,25,33,48,75,88,108,113); mentum with few long setae (Fig. 2); labrum anteriorly broadly membranous and with two obtuse median projections (3,16). Pronotum weakly to moderately transverse; posterior angles obtuse, usually weakly mar ked, sometimes more or less obsolete; maximal width in, or a short distance anterior to middle; surface distincdy convex in cross-section; pubescence along midline directed caudad, in lateral areas ± diagonally latero-caudad, and near posterior margin ± transversely laterad. Hypomera distinctly visible in lateral view. Prosternum with median carina. Elytra as wide as or slighdy wider than pronotum and at suture slightly to distincdy shorter than pronotum; hind margin near posterior angles strongly or indistincdy sinu ate; hind wings fully developed; mesosternum with (± reduced) or without median cari na; mesosternal process long, reaching halfway between mesocoxae, apically acute, roun ded, or distinctly truncate; mesocoxal cavities delimited from metasternum by carina. Legs long and slender; first metatarsomere very long and slender, at least as long as the combined length of the two following tarsomeres (Figs. 35,67,70). Abdominal terga III-V with ± deep, tergum VI with weaker or without anterior impres sions; sterna III-V or III-VI anteriorly constricted; tergum VIII posteriorly convex and with relatively long thin setae (Figs. 11,13); tergum X anteriorly often with ± long and slender extension, with numerous stout setae, which are longer posteriorly than an teriorly, in posterior median area in most species without setae, posterior margin with membranous fringe (Figs. 6,20,27,36,43,56,79,91,111,114). d : sternum VIII posteriorly weakly to moderately convex with a row of moderately long and a row of longer thin setae (Fig. 12); median lobe of aedeagus with almost straight ventral process (lateral view), at base of ventral process often with lateral and/ or median folds, and with weakly pronounced crista apicalis; internal sac with moderate ly to very long internal tube, and with weakly to distinctly sclerotized additional struc tures; apical lobe of paramere conspicuously small and short, usually ± coniform (e. g. Fig. 7), sometimes modified (flattened, with torsion, or even transverse) (Figs. 84,85,95,98), with two moderately long subapical and two short apical setae. For illustrations of the parameres of A. umbrosa (plesiomorphic condition) and A.. armata sp. n. (derived condition) see SEEVERS (1978) and Fig. 94, respectively. 9: sternum VIII similar to that of d , posteriorly usually less strongly convex, its po sterior margin with slender, weakly modified or unmodified setae, and without micropu bescence (Fig. 14); spermatheca of distinctive morphology, capsule large and bulbous, duct relatively short and ± straight (e. g. Figs. 10,24,28,32,42,47); general shape of spermatheca somewhat resembling that of Piochardia H EYDEN of the Aleocharini.

S ystem atics and comparative notes
Based on morphological characters, A marochara is correctly placed in the Oxypodini. From other genera of this tribe, it is distinguished especially by the morphology of the antennae, by the morphology of the median lobe of the aedeagus, of the apical lobe of the paramere, and of the spermatheca, and from many genera also by the absence of micropubescence and of distinctly modified setae on the posterior margin of the 9 sternum VIII. A closer relationship to Calodera MAN N ERH EIM , 1 830 and Ilyobates KRAATZ, 1856, as presumed by previous authors because of the similar external appearance (slen der body, weakly transverse pronotum, relatively long and slender legs) and especially the stout antennae, seems rather unlikely. Amarochara is separated from these genera by numerous significant characters, especially by the morphology of the mouthparts (ligula, chaetotaxy of the labrum, slender maxillary palpi) and of the antennae (dorso-apical furrow on antennomere I, strongly transverse antennomere IV and V, shape and pube scence of antennomeres VI-X), by the shape and chaetotaxy of tergum X, the shape and internal structures of the median lobe of the aedeagus, the short apical lobe of the paramere with relatively short setae, and by the characteristic morphology of the sper matheca. In addition, A marochara is of almost global distribution, whereas Ilyobates is Western Palaearctic and Calodera probably Holarctic. (Several species of Calodera have been described also from other biogeographic region, but those species that I have examined refer to other genera; the same is probably true of those taxa that have not been reexamined.) The adelphotaxon of A marochara is unknown. Among the betterknown genera of Palaearctic Oxypodini, there is no likely candidate.

Phylogenetic analysis
In order to assess the intrageneric phylogenetic affiliations, morphological and biogeo graphic data were compiled and transformed into a data matrix. As the adelphotaxon of Amarochara is unknown, several representatives of presumably more closely related oxypodine genera were selected to determine character polarity: Calodera nigrita M A N N ER HEIM, C. rufescens KRA ATZ , Euryalea decumana (E R IC H SO N ), E. murina (E RICH SO N ), Ilyoba tes bennetti DO N ISTH O RPE, and O caleapicata (STEPHEN S). As more distant outgroup, a species of Zoosetha was chosen: Z rufescens (K RAA TZ ). Whenever in doubt regarding character polarity', the corresponding character states in other aleocharine tribes (Falagriini, Athetini) were considered in the character discussion, but are not listed in the data ma trix below. For illustrations and descriptions of the taxa here selected as outgroups see ASSING (1996,1998,1999) and A ssiN G & WUNDERLE (1997A ssiN G & WUNDERLE ( , 1998. 3.3.1.1. Discussion of character states and polarities 1 Eyes: large, in dorsal view approxim ately as long as postgenae or nearly so (0); distinctly shorter than postgenae (1).

7
Antenna: not massive and antennomeres IV-X not transverse (0); massive and antennomeres IV-X transverse (1). The massive antennae in A m arochara, llyobates, Calodera, and Zoosetha are very unlikely to be homologous. A t least in llyobates and Zoosetha, they are o f different m orphology in many other respects. 8 Antenna: antennomeres VI-X distinctly transverse, but m oderately w ide, <1.5 times as w ide as antennomere I (0); antennomeres VI-X massive, >1.5 times as w ide as antennomere I (1).

9
Antenna: massive and relatively short, when turned backwards only slightly extending beyond pronotal hind margin (0); massive and long, distinctly extending beyond pronotal hind margin, rea ching posterior margin o f elytra or nearly so (1). The long antennae in most o f the outgroup taxa are unlikely to be hom ologous to those in A m arochara and prim arily due to the long and slender antennomeres IV-X; this is particularly true o f the antennae in E uryalea and Ocalea. Long antennae are evidently hom oplastic in several species groups of A marochara.

11
Antenna: dorso-apical furrow o f antennomere I weakly pronounced, usually about h alf the length o f antennomere (0); dorso-apical furrow pronounced, almost extending over w hole length of antennomere I (1). This character is subject to considerable intraspecific variation and consequent ly attributed only little significance.

12
Antenna: antennom ere III relatively long and slender, approxim ately as long as antennomere II or nearly so (0); antennomere III distinctly coniform and clearly shorter than antennom ere II. The polarity o f this characters is doubtful.
14 M axillary palpus: third joint m oderately slender, 2 -3 times as long as broad (0); third joint very slender, more than 3 times as long as broad (1). The derived condition o f this character in some species o f A m arochara on the one hand and in Ocalea and Fmryalea on the other hand is very unlikely to be homologous.

20
Pronotum: puncturation moderately sparse (0); puncturation sparse (1); puncturation conspicuously dense (2). The similar character code for some A marochara, llyobates, E. m urina, Calodera spp., and Z. rufescens is without doubt hom oplastic and at least in part a m orphological adaptation to humid habitats; see also character 17. The interm ediate condition is hypothesized to be plesiom orphic, the two extreme conditions are likely to be derived.

21
Pronotum : posterior angles well-defined (0); posterior angles obsolete or nearly so (1). This charac ter is subject to some intraspecific variation and attributed little phylogenetic significance.

22
Pronotum sm all in relation to head, less than 1.3 times as wide as head (0); pronotum large, at least 1.3 time as w ide as head (1). This character is o f lim ited significance for assessing phylogenetic relationships, since it is subject to some intraspecific variation and several species are o f interm e diate condition.

25
Mesosternum: m eso stern al carin a absent (0); p resen t (1). T h e p o larity o f this ch aracter is un cer tain. It is kn ow n to be sub ject to in tragen eric variatio n also in o th er aleo ch arin e gen era (e. g. A kochara GRAVENHORST and A crotona THOMSON).

27
M etatarsus: metatarsomere I relatively short, distinctly shorter than the com bined length o f the three following tarsomeres (0); metatarsomere I approxim ately as long as the com bined length of the three following tarsomeres or longer (1). This character is subject to considerable intraspecific and in-group variation and therefore attributed little phylogenetic significance.

29
Abdom inal tergum VI: anterior im pression shallow or obsolete (0); impression deep (1). The oc currence o f species with the derived condition in A m arochara, Calodera, and E uryalea reflects a parallel evolutionary trend in these genera. There is, however, some doubt if the polarity' here assumed also applies to Calodera.

30
Abdom inal terga III-VI: puncturation anteriorly m oderately coarse (0); puncturation anteriorly conspicuously coarse and dense (1); puncturation anteriorly fine (2). The derived character state 1 in some species o f the umbrosa group and in other species groups o f A m arochara is apparently the result o f parallel evolution.

33
Abdom inal terga III-V: without transverse series o f coarse punctures in posterior h alf (0); with transverse series o f punctures in posterior h alf (1). 34 9 sternum VIII: posteriorly w ith micropubescence (0); without micropubescence at posterior margin (1). The polarity assumed here is based on the observation that the presence o f m icropubescence is the usual condition found in m any Aleocharinae, not only in Oxypodini, but also in Athetini, Falagriini, etc. 35 9 sternum VIII: marginal setae at posterior margin distinctly modified (i. e. very stout) (0); weakly m odified (1). It can be inferred from the presence o f distinctly modified setae in m ost Oxypodini and in several other aleocharine tribes that the weakly modified setae are apparently the result o f a reduction and consequently represent the derived condition.

39
Aedeagus: ventral process fused with dorsal part o f m ain body (0); ventral process separated for most o f its length (1). The derived condition in some A m arochara on the one hand and in E uryalea on the other hand is clearly not homologous.

49
Aedeagus: apical lobe o f paramere long and slender, >3 times as long as w ide and w ith long setae (0); apical lobe o f reduced length and w ith shorter setae (1).

51
Aedeagus: apical lobe o f paramere of ± coniform shape and oval in cross-section (0); apical lobe of paramere not coniform , distinctly flattened in cross-section (1).    For several reasons it is difficult to determine the intrageneric phylogenetic affiliations unequivocally. The main weakness of the character matrix (Tab. 1) is obviously the fact that the male sexual characters, which are of particular phylogenetic significance in Ama rochara, of almost one third of the species are unknown. In addition, many external character are subject to considerable in-group and even intraspecific variation, so that it was sometimes difficult to determine the respective character state or to distinguish between homoplasies and secondary reductions. A transformation of the character matrix into a phylogenetic tree is here refrained from, not only because many of the basal ramifications and lineages are weakly supported, but also because this would require an examination of the species from other zoogeographic regions. Instead, some basic con clusions are here discussed in more detail.

The subgenera o f A m a ro c h a ra
The species of Amarochara occurring in the Holarctic region have been attributed to three subgenera (see also

Distribution and bionomics
Species described in or transferred to Amarochara are known from the Holarctic, the Neotropical, the Oriental, and the Ethiopian region (see also section 1). However, apart from those occurring in the Holarctic region and some Ethiopian representatives, the species have not been reexamined.
Remarkably, Amarochara ecologically resembles the aleocharinae genera Callicerus GRA-VENHORST (Athetini), Pseudosemiris MACHULKA (Athetini), and Ilyobates KRAATZ (Oxypodini) in many ways, all of these taxa additionally sharing some morphological charac ters, such as a slender body and remarkably long and stout antennae (see ASSING, 1999ASSING, , 2001. The collection data suggest that Amarochara species are univoltine and that repro duction and larval development take place in an unknown subterranean habitat in spring and early summer. Most, if not all the species are capable of flight, which explains why many species are apparently relatively widespread. Adult beetles have been collected from litter, moss, decaying organic matter, and other substrates in a wide variety of forested and open habitats. Amarochara larvae are unknown. For further details see the species sections.

The u m b ro sa group
The umbrosa group comprises six rather small species of Palaearctic (1 species), Eastern Palaearctic (2 species), and Nearctic distribution (3 species). They share the following derived characters: a more or less subrectangular head, usually with weakly marked po sterior angles, pardy to completely reduced genal carinae, a relatively short antennomere III, usually straight (i. e. not sinuate) posterior elytral margins, and an aedeagus with a moderately to very long apical part and a moderately long basal part of the internal tube, and with a usually reduced crista apicalis (see also section 3.3).

A m a ro c h a ra u m b ro sa (E r ic h s o n , 1837)
Figs.

Comments
The original description of Calodera umbrosa is based on an unspecified number of syntypes collected near Berlin by Schüppel. In order to fix a single name-bearing type and to secure the present interpretation of the species, the male syntype is here designated as lectotype.
According to the respective curators in charge, the type(s) of Homalota tenuisAsSA (type locality: "Geneva") were looked for, but not found, in the Heer collection in Zürich and in the collections of the B M N H , where, according to HORN et al. (1990), part of the types of the species described by HEER are kept (Brendell,   Slovakia: 23 exx., Vysocany, 6.VIII.1919 (NMP, cAss); 11 exx., sam e locality, 12.VIII.1918 (NMP); 9 exx., same locality, 30.VII.1919 (NMP); 19 exx., same locality, 9.VII.1919 (NMP); 1 ex., Zliechov, 25.IV.1917 (NMP); 4 exx., Kosice, 1926Kosice, , 1928Kosice, , 1929  Redescription 2.2 -3.2 mm (abdomen fully extended). Colour of body dark brown to blackish, often with the elytra lighter, yellowish brown to brown, and the posterior margins of the abdominal segments dark ferrugineous; antenna dark brown to blackish brown, legs light brown to brown. Integument shining; forebody usually without appreciable micro sculpture (sometimes traces of microsculpture may be present on head and pronotum); abdomen at most with barely noticeable microsculpture. Head of subquadrate shape, approximately as -wide (across eyes) as long; eyes very large and prominent, distinctly projecting from lateral outline of head and only slightly shor ter than postgenae in dorsal view, the latter almost parallel for about half their length. Puncturation very fine and sparse. Genal carina reduced. Antenna as in Fig. 5, antennomere I with pronounced dorso-apical furrow. Mouthparts as in Figs. 1-4. Pronotum approximately as wide as long and 1.2 -1.3 times as wide as head; maximal width usually approximately halfway between anterior angles and middle; puncturation highly variable, very fine to distinct, usually denser and more distinct than that of head. Elytra at suture 0.90 -0.95 times the length of pronotum and approximately 1.20 -1.25 times as wide as pronotum; posterior margin near posterior angles not or only indistincdy sinuate; puncturation weakly granulóse and somewhat denser than that of pro notum. Hind wings fully developed. First metatarsomere approximately as long as the combined length of the two following tarsomeres. Mesosternum with microsculpture composed of meshes of various lengths, without median carina; mesosternal process apically ± narrowly truncate or rounded. Abdominal terga III-V with moderately deep, tergum VI without appreciable anterior impression; sterna III-V anteriorly constricted; puncturation fine, but distinct, modera tely dense (though the density1 is subject to some intraspecific variation), and more or less evenly distributed on dorsal surface; tergum X as in Fig. 6. d : posterior margin of tergum VIII slightly' convex, with a row of moderately long and a row of long thin marginal setae (Fig. 11); posterior margin of sternum VIII distincdy convex, with two rows of long thin setae (Fig. 12); aedeagus with ventral process of median lobe in ventral view relatively broad, in lateral view weakly' curved, and with basal fold; crista apicalis and crista proximalis reduced; internal sac with very long flagellum (Figs. 8,9); apical lobe of paramere relatively short and broad (Fig. 7). 9: posterior margin of tergum VIII of similar chaetotaxy as in d , but slightly more convex (Fig. 13); posterior margin of sternum VIII weakly convex, row of marginal setae very weakly modified, relatively thin, and shorter than in d (Fig. 14); spermatheca as in Fig. 10.

Comparative notes
From other Western Palaearctic congeners, A. umbrosa is distinguished by' the combina tion of dark colour, small body size, more slender antennae, an almost subrectangular head shape, large eyres, the absence of genal carinae, and especially by the morphology of the aedeagus. For separation from similar species occurring in the Nearctic and the Eastern Palaearctic region see the comparative notes below those species and the key' in section 5. In the collections examined, A. umbrosa was most frequently7 confused with Calodera aethiops (GRAVENHORST), with which it shares a similar general appearance (size, proportions, slender pronotum) and the massive antennae, from which it is, however, distinguished by the much sparser puncturation and more distinct shine of forebody' and abdomen, the different shape of antennomere I, and by the completely different morphology of the sexual characters.

Distribution and bionomics
XL umbrosa is widespread in the southern and central parts of the Palaearctic region from Spain in the West to China in the east (Maps 1,2). In the north and northwest it reaches the south of Norway, Sweden, and Finland, almost all of Denmark, and southern Eng land (see also FOW LER (1988), H ANSEN (1964), HAN SEN (1996) 1995), and SlLFVERBERG (1992)). In the west it is known from various regions in France (material examined; H O RIO N , 1967). There is only a single record from Spain (north of Madrid) (G AM ARRA, 1987), which, however, requires veri fication and is not included in Map 1. It occurs in all of Italy from the Alps in the north to Sicily in the south (LU IG IO N I, 1929, W Ö R N D LE , 1950, ZAN ETTI, 1995, and material examined). It is known from some localities in Switzerland and all Austrian regions (H O RIO N , 1967, HUGEN TO BLER, 1966, SCHEERPELTZ, 1968. In Germany, the species is widespread, but has not been reported from Niederelbe ( Poland, the Czech Republic, Slovakia, Slovenia, Hungary, Romania, and the northern Balkans, but has not been reported from Greece (H O R IO N , 1967, RAM BO USEK, 1910, and material examined). The easternmost records are from Novosibirsk, Russian Feder ation (first record) and from China, from where it was first reported by PACE (1998).
For additional records see BO H ÄC et al. (1993), BÜ CH E (1994), FRAN Z (1938), KO CH (1968), K ö h l e r & S t u m p f (1992), L i n k e (1907,1927) LO H SE, 1967,1984 have assumed thatM. umbrosa is associated with subterranean burrows and nests of small mammals. However, this hypothesis has not been confirmed by systematic studies of such habitats (especially of mole nests). Morphological (slender body, massive antennae) and bionomic evidence (see below) suggests that the ecology of A. umbrosa and other congeners may be similar to that of lljobates K RA A TZ , Callicerus GRAVENHORST, and Pseudosemiris M ACH ULKA, whose reproduction and hibernation habitats are unknown, too (A SSIN G , 1999(A SSIN G , , 2001. The specimens examined were collected from February through August and in October ( Fig. 15), with a maximum from April through August; PEEZ AND K A H LE N (1977) also observed adult beetles in September and November. Records from December and Janu ary are unknown. According to H O RIO N (1967), teneral specimens appear in July and August. These observations suggest that the species is univoltine with spring reproduc tion and pre-imaginal development taking place in spring and early summer. The mature eggs of A. umbrosa are very large (approximately 0.6 mm long), so that the number of eggs laid per female is probably relatively low.

Comments
The original description of Amarochara heterogaster CAMERON is based on an unspecified number of syntypes. As the possibility that more syntypes exist cannot be excluded, the male syntype in the Cameron collection is here designated as lectotype to preserve the present interpretation of the species. The coloration indicated in the original descripti on may be misleading, since the type is evidently a teneral specimen. The original description of A. simlaensis CAMERON is based on an unspecified number of syntypes, two of which were located in the BMNH. A lectotype is not designated, since both of these syntypes are females and the possibility that a male symtype exists cannot be excluded. A comparison of the types of A. heterogaster, A. simlaensis, and A. smetanai yielded no convincing evidence that they should represent distinct species. The lectotype of A. heterogaster is apparently a nanistic male with a relatively small pronotum, but the aedeagus is identical with that of the holotype of A. smetanai. According to PACE (1992), A. smetanai is distinguished from A. simlaensis and A. heterogaster by the different coloration of antennomere I, by the relatively shorter antennomere III, and by the wider elytra. A comparative examination of the tyrpes, however, revealed that these differences do not exist; in the lectotype of A. heterogaster, the elytra are even wider in relation to the prono tum than in the types of A. smetanai. Consequently, both A. simlaensis CAMERON  Pubescence of forebody relatively short and dense (especially on pronotum). Head of similar shape and morphology as in A. umbrosa, but less distinctly subquadrate, slightly (approximately 1.05 times) wider than long, not dilated behind eyes; postgenae relatively shorter, slightly longer than eyes; eyes large and convex, weakly projecting from lateral outline of head; postgenae weakly and smoothly convex, posterior angles obsolete. Puncturation extremely fine, barely noticeable. Genal carina rudimentary, usually visible only below eyes and/or near posterior constriction. Antenna more massive than in A. umbrosa, otherwise of similar morphology; antennomere I with shorter dorsoapical furrow than in average A. umbrosa-, antennomere III shorter than II (Fig. 19).
Mouthparts as in A. umbrosa, but ligula relatively larger (Fig. 17), labrum with median pair of sensilla at anterior margin stouter, and lateral margins of labrum with some very short setae (Fig 16); maxillary palpus as in Fig. 18. Pronotum of similar relative size and proportions as in A. umbrosa, weakly transverse, approximately 1.05 -1.10 times as wide as long and 1.13 -1.25 times as wide as head; maximal width in anterior half; puncturation dense, but shallow, fine, and ill-defined. Mesosternum without median carina; mesosternal process apically relatively broadly trun cate. Elytra at suture approximately 0.9 times the length of pronotum and approximately 1.3 times as wide as pronotum; posterior margin near posterior angles not distinctly sinuate; puncturation fine (but more distinct than that of head and pronotum), indi stinctly granulose, and relatively dense. Legs as in A. umbrosa, first metatarsomere of variable length, approximately as long as the combined length of the two following tarsomeres or slightly longer. Abdomen with anterior impressions of terga III-V slightly deeper and broader than in A. umbrosa, tergum VI without anterior impression; sterna III-V anteriorly constricted; puncturation in anterior impressions of terga III-V and near anterior margin of tergum VI very dense and coarse; on remainder of tergal surface, too, with numerous coarse (though somewhat less dense) punctures, which are decreasing in size and density to wards the posterior tergal margins. Tergum X anteriorly broadly truncate, not distinctly elongated (Fig. 20); practically all of its surface, except for a small median area near hind margin, with very dense stout and long setae. c?: posterior margin of tergum VIII weakly convex, with a row of long and rather stout marginal setae; posterior margin of sternum VIII strongly convex, with a dense row of relatively long marginal setae and with interspersed long submarginal setae; aedeagus with long and slender ventral process, which is distinctly separated from the dorsal part of the median lobe; crista apicalis small, crista proximalis long, but not very prominent; internal sac with very long tube and with additional pair of long, weakly sclerotized structures ; apical lobe of paramere with two long and two short setae (Fig. 23). 9: tergum VIII posteriorly almost truncate and with sparse and moderately long margi nal setae; posterior margin of sternum VIII convex, row of marginal setae not dis tinctly modified, moderately short, denser and somewhat shorter in the middle than in lateral parts of posterior margin; spermatheca smaller than in A. umbrosa and with shor ter duct (Fig. 24).

Comparative and systematic notes
For distinction from the similar A. umbrosa see description above. From other Himalay an species, except for A. sororcula, A. heterogaster differs in the absence of a distinct neck. Apparently based on the presence of fine (but rudimentary) genal carinae, CAMERON (1939) attributed the species to the subgenus luisiochara. However, as can be inferred from other characters, the species is more closely related to A. umbrosa, the type species of the nominal subgenus, than to A. bonnairei, the type species of l^asiochara (see section 3.3).

Distribution and bionomics
A. heterogaster is known from northern India (Himachal Pradesh and Ghum district near the border to Sikkim) and from two localities in Nepal; the type locality of A. smetanai is in the Khandbari district in eastern Nepal, and PACE (1984) recorded the species (as A. simlaensis) also from Jaljale Himal. A. heterogaster was collected at altitudes between 2100 and 2950m in spring (April through June) and in autumn (September, November).

A m a r o c h a r a s o r o r c u l a Cam e r o n , 1939
Figs. 25-28 A.marochara (Lasiochara) sororcula CAMERON, 1939: 574f. Type examined Syntype 9: Dehra Dun. Dr. Cameron. 7-10-1922

Comments
The original description is based on an unspecified number of syntypes. The possibility that a male syntype exists cannot be ruled out, so that a lectotype is not designated.

Redescription
Of similar size as A. umbrosa-, 2.1 mm. Head and abdomen (except for the lighter tergal margins) dark brown; pronotum brown; elytra yellowish brown; antennomeres I-II light brown and III-XI dark brown; legs and maxillary palpi testaceous. Microsculpture as in A. umbrosa. Pubescence of forebody distinctly shorter than in A. umbrosa and ± decum bent.
Head of similar shape and size as in A. umbrosa, but postgenae relatively shorter and posterior angles only weakly marked; eyes large, in dorsal view almost as long as postge nae; puncturation similar to that in A. umbrosa. Genal carinae present, but fine. Antenna as slender as in A. umbrosa and of similar morphology, but antennomere III slightly stouter. Mouthparts as in Figs. 25-26. Pronotum relatively large, 1.3 times as wide as head and 1.15 times as wide as long; maximal width near the middle, posterior angles weakly marked; puncturation slightly denser than in A. umbrosa. Elytra only slightly (1.15 times) wider than and at suture 0.85 times as long as pronotum; posterior margins near posterior angles not sinuate; puncturation fine and dense, dis tinctly denser than in A. umbrosa. Mesosternum apically acute. Legs slightly shorter than in A. umbrosa, metatarsus almost as long as metatibia; first metatarsomere almost as long as the combined length of the three following tarsomeres. Abdomen with anterior impressions of terga III-V shallow and posteriorly ill-defined; sterna III-V with weak anterior constriction; puncturation distinctive: on terga III-VI weakly granulose; on terga III-V dense and well-defined, but not very coarse, not coarser and denser in anterior impressions than on remainder of tergal surface; on tergum VI slightly coarser and denser than on terga III-V, and on tergum VII finer and sparser than on terga III-VI. Tergum X apically elongated and broadly truncate (Fig. 27); setae testa ceous and stout, in anterior half very dense (also in the middle); posterior median area without setae.
cT: unknown. 9: tergum VIII posteriorly weakly convex and with row of relatively sparse and long marginal setae; posterior margin of sternum VIII weakly convex and with moderately dense row of weakly modified marginal setae; spermatheca of similar size and morpho logy as in A. umbrosa, but capsule somewhat less globulous (Fig. 28).

Comparative and systematic notes
From A. umbrosa, A. sororcula is readily separated especially by the presence of genal carinae, the less marked posterior angles of the head, the relatively larger pronotum, the short, dense, and decumbent pubescence of the forebody, the shallower anterior im pressions of the abdominal terga III-V, and by the characteristic puncturation of the abdomen. From other Palaearctic congeners, it differs in the slender antennae, the short, dense, and decumbent pubescence, the shallow anterior impressions of the abdominal terga III-V, and in the puncturation of the abdomen. Apparently based on the presence fine genal carinae, CAMERON (1939) attributed the species to the subgenus Fasiochara. However, as can be inferred from other characters, the species is more closely related to A, umbrosa, the type species of the nominal subgenus, than to A. bonnairei, the type species of Lasiochara (see section 3.3).

Distribution and bionomics
Only the holotype of this species has become known, which was collected from dung in Dehra Dun (Northern India: Uttar Pradesh).

Comments
The original description is based on an unspecified number of type specimens. In order to fix a single name-bearing type and to secure the present interpretation of the species, the syntype in better condition is here designated as lectotype.

Redescription
In size and general appearance similar to A. umbrosa, but distinguished as follows: Coloration lighter, whole body ± uniformly ferrugineous, with the anterior 3/4 of ab dominal'tergum VI infúscate, and with the legs and the basal antennomeres testaceous. (In the paralectotype, the posterior part of the head is somewhat darker and the elytra are lighter than the pronotum.) Pubescence ± decumbent. Head with extremely weak, but (at higher magnifications) noticeable microsculpture. Head weakly (approximately 1.05 times) wider than long, larger in relation to pronotum than in A. umbrosa (see ratio below); hind angles very weakly marked, almost obsolete; eyes slightly smaller and less convex, only weakly protruding from lateral outline of head. Genal carina absent. Antenna of similar length as in A. umbrosa, but distinctly more massive; antennomere with distinct and rather long dorso-apical furrow, antennomere IV longer and less transverse, antennomeres VT-X much wider and more transver se than in A. umbrosa, more than twice as wide as long. Mouthparts of similar morpho logy as in A. umbrosa. Pronotum 1.10 -1.15 times as wide as long and only 1.20 times as wide as head; maximal width approximately halfway between middle and anterior angles; posterior angles obtu se, moderately marked; puncturation extremely fine; microsculpture similar to that of head or barelv noticeable. Elytra at suture approximately 0.9 times the length of pronotum and approximately 1.25 times as wide as pronotum, distinctly shorter in relation to pronotum than in A. umbrosas, posterior margin near posterior angles not or very indistinctly sinuate. Legs slightly lon ger than in A. umbrosa, first metatarsomere slightly or distinctly longer than the combi ned length of the two following tarsomeres. Mesosternum without median carina; mesosternal process apically acute. Abdomen as in A. umbrosa, but puncturation on the whole finer, in anterior impressions of terga III-V moderately coarse and on remainder of dorsal surface very fine; tergum X of similar chaetotaxy as in A. umbrosa, but anteriorly less elongate (Fig. 43). d1: unknown. 9: posterior margin of tergum VIII truncate, with row of sparse marginal setae; posterior margin of sternum VIII weakly convex, marginal setae weakly modified, distinctly denser in the middle than laterally; spermatheca larger than in A. umbrosa (Fig. 38).

Comparative notes
FromH. umbrosa, A. inquilina differs as indicated in the redescription above. For distinction from other North American congeners see comparative notes and descriptions below.

Distribution and bionomics
The species is known only from the type locality in Iowa, where it was collected in ant nests (Formica sp.).

Description
In general appearance similar to A. umbrosa, but distinguished as follows: Body larger, 3.0 -3.8 mm (abdomen fully extended). Coloration on average lighter than in A. umbrosa-, head, antennae, and abdomen dark brown to blackish, with the tergal margins, the abdominal apex and sometimes the basal antennomeres lighter, ferrugineous to castaneous; pronotum castaneous to dark brown; elytra light brown; legs testaceous; maxillary palpi testaceous or infúscate. Microsculpture similar to that in A. umbrosa. Head of similar shape and morphology as in A. umbrosa, but eyes relatively larger, in dorsal view approximately as long as postgenae or nearly so. Puncturation as in A. um brosa. Genal carina absent. Antenna more massive than in A. umbrosa, antennomeres IV and V more transverse, and VI-X wider; dorso-apical furrow of antennomere I variable, usually pronounced (Fig. 37). Mouthparts as in A. umbrosa, but third joint of maxillary palpus longer and more slender (Fig. 34); labium as in Fig. 33. Pronotum larger (especially in relation to head) than in A. umbrosa, approximately as wide as long and 1.25 -1.3 times as wide as head; maximal width near the middle; puncturation very fine and sparser than in average A. umbrosa. Elytra at suture 0.75 times the length of pronotum and approximately 1.25 times as wide as pronotum, distinctly shorter in relation to pronotum than in A. umbrosa-, posterior mar gin near posterior angles weakly sinuate; puncturation similar to that in A. umbrosa. Legs longer and more slender than in A. umbrosa, first metatarsomere approximately as long as the combined length of the three following tarsomeres (Fig. 35). Mesosternum wit hout median carina; mesosternal process apically ± narrowly truncate or rounded, al most acute. Abdomen as in A. umbrosa, but puncturation in anterior impressions of terga III-V and on remainder of dorsal surface coarser and sparser, integument with more shine; tergum X of similar chaetotaxy as in A. umbrosa, but anteriorly more strongly elongated ( Fig. 36), at least in 9 occasionally with 2-4 setae near posterior membranous fringe. d : posterior margin of tergum VIII + truncate, that of sternum VIII weakly convex to almost truncate and with row of rather dense and long marginal setae; median lobe of aedeagus of similar general morphology as in A. umbrosa, but larger, more slender, with much longer ventral process, and with distinct crista apicalis and crista proximalis ; apical lobe of paramere short, broadly triangular, and with rather long setae (Fig. 31). 9: posterior margin of sternum VIII moderately convex, row of marginal setae very weakly modified, relatively thin; spermatheca of similar absolute, but of lower relative size than in A. umbrosa (Fig. 32).

Comments and comparative notes
The original description is based on material from "Lawrence and Clark counties", so that it can be assumed that Blatchley had at least two syntypes before him. Since the type specimen in the Blatchley collection is a female, it is not designated as lectotype; a male syntype may yet be discovered. Apparently, either the date indicated in the original des cription ("May 6-May 12") or that given on the label (June 6) is erroneous. Nevertheless, there is little doubt that the examined specimen from the Blatchley collection in fact represents a syntype. The specimen from Georgia is of somewhat lighter coloration and its pronotum is even larger than in the material from Kansas. Since the examined syntype is a female, it is not absolutely certain that it is conspecific with the specimens listed as additional material examined. However, significant distinguishing characters were found neither in external morphology nor in the female primary and secondary sexual characters, so that they are here attributed to the same species. From the similar A. umbrosa, A. fenyesi is readily distinguished especially by its larger size, the more massive antennae, the larger pronotum, the relatively shorter elytra, the longer first metatarsomere, and by the coarser puncturation of the abdomen (see description above). The North American A. inquilina is smaller, of lighter coloration, and has much shorter antennae, a more transverse and distinctly smaller pronotum (especially in relation to head), longer elytra, a much finer abdominal puncturation, and a larger spermatheca.

Distribution and bionomics
The species is known from Indiana, Kansas, and Georgia. Most of the specimens were collected with flight intercept traps between 21 May and 25 July (maximum in July), suggesting that A. fenyesi is an active flyer and more widespread than is currently known.

Description
In size and general appearance highly similar to A. umbrosa, but distinguished as follows. Length of holotype: 2.8-3.2 mm. Coloration lighter than in A. umbrosa-, head and abdomen dark brown, with margins of the abdominal terga and the abdominal apex lighter; pro notum brown, elytra light brown; legs and maxillary palpi testaceous; antennae bicoloured, dark brown, with the basal antennomeres yellowish to ferrugineous. Microsculpture similar to that in A. umbrosa. Pubescence relatively short and ± decumbent. Head of similar shape and morphology as in A. umbrosa, but posterior angles less mar ked. Antenna of similar length as in A. umbrosa, but distinctly more massive; antennomere I in the holotype with very short and weak dorso-apical furrow, antennomere III stouter and apically more strongly dilated than in A. umbrosa-, antennomeres VI-IX wider and more transverse (distinctly more than twice as wide as long, antennomere VI almost three times as wide as long) than in A. umbrosa. Genal carina absent. Pronotum and elytra of similar size, shape, and proportions as in A. umbrosa-, puncturation of pronotum very fine and rather dense. Legs of similar morphology as in A, umbrosa, first metatarsomere longer than the combined length of the two following tarsomeres, but shorter than the combined length of the three following tarsomeres; length of metatarsus approximately 0.3 mm. Mesosternum without median carina; mesosternal process apically narrowly truncate. Abdomen with dense and relatively coarse puncturation, on the whole similar to A. umbro sa, but coarser, especially on tergum VI; punctures finer near posterior than near anterior tergal margins. Tergum X of similar shape and chaetotaxy as in A. umbrosa. 6 : sternum VIII posteriorly weakly convex and with dense long marginal setae; aedeagus with median lobe very similar to that in A. umbrosa, but ventral process with pro nounced lateral folds and crista proximalis not reduced ; apical lobe of paramere with longer setae (Fig. 41).
? : posterior margin of tergum VIII truncate, that of sternum VIII weakly convex, row of marginal setae very weakly modified; spermatheca with larger capsule and longer duct than in A. umbrosa (Fig. 42). Derivatio nominis: The name is a noun in apposition composed of the Latin adjective brevis (short) and the Latin noun os (leg) and refers to the relatively short legs, which distinguish this species from other North American congeners.
Comparative notes From the similar A. umbrosa, A. brevios is best distinguished especially by the lighter colour, the stouter antennae, and the different genitalia. The two other North American species, A. inquilina and A .fenyesi, differ in the longer legs, the different coloration, the distinctly sparser abdominal puncturation, the former in addition by the less convex eyes, the much finer abdominal puncturation, and the latter by larger size, a much larger pronotum, longer antennae, and a smaller spermatheca.

Distribution and bionomics
The species is known only from the type locality in Kansas, where it was collected with a flight intercept trap in July, together with A. fenyesi.

T he b o n n a i r e i group
The bonnairei group includes at least 7 species, all of them occurring in the Western Palaearctic region (see also section 4.6). A. tingitana JARRIGE may belong to this group, too, but as the holotype is lost, its phylogenetic affiliations must remain doubtful. The position of A. crassicornis is somewhat uncertain, because the apical abdominal segments of the holotype were missing and no additional material was available for examination, so that the conditions of many significant characters are unknown. The bonnairei group is relatively weakly defined, because most of the synapomorphies are either not realized in all the species of the group or because they also occur in other groups of Amarochara. In addition, the fact that the male sexual characters of two spe cies are unknown renders a definition difficult. The species of the bonnairei group share the following derived characters: relatively small eyes (exceptions: A. caeca, A. splendens, A. crassicornis), fine genal carinae (exception: A. splendens, A. crassicornis), long antennae, a slender third joint of the maxillary palpus, a large pronotum (exception: A. bonnairei), a long and slender first metatarsomere, a fine and sparse puncturation of the posterior halves of the abdominal terga III-VI (relatively coarse and dense in A. splendens), an anteriorly distinctly elongated tergum X, an aedeagus with the ventral process distinctly separated from the main body of the aedeagus, a moderately or very long basal part of the internal tube of the aedeagus, and a relatively large spermatheca (see section 3.3.1).  (1996), Fr a n z (1970), G e is e r (1998) In conclusion, based on A rticle 23.9 o f the Code, the follow ing synonym y is established: Redescription 2.8 -4.0 mm (abdomen fully extended). Colour of body usually yellowish brown to light brown, with the head, antennomeres IV-XI, the anterior central areas of the abdominal terga VI-VII, sometimes also of terga IV-V, often slightly darker brown; legs and mouthparts testaceous. Microsculpture of head and pronotum shallow, but usually distinct, composed of meshes of variable length and orientation; abdominal terga VI-VIII or V-VIII usually with extremely weak transverse microsculpture or traces of such micros culpture. Pubescence of forebody relatively long and suberect. Head of ovoid shape, slightly (approximately 1.10 times) wider than long; noticeably dilated behind eyes, maximal width therefore not across eyes, but at some distance be hind eyes; eyes moderately large, weakly convex, not distincdy projecting from lateral outline of head, only little more than half the length of postgenae in dorsal view, the latter smoothly curved towards posterior constriction. Puncturation very fine and sparse. Genal carina present, but fine. Antenna as in Fig. 50, antennomere I with dorso-apical furrow less pronounced than in A. umbrosa. Third joint of maxillary palpus very long and slender, more so than in A. umbrosa (Fig, 49); labium as in Fig. 48. Pronotum approximately 1.1 times as wide as long and 1.2 -1.3 times as wide as head; maximal width in or near the middle; puncturation similar to that of head or slightly denser and more distinct. Elytra relatively shorter than in A. umbrosa, at suture 0.74 -0.80 times the length of pronotum and approximately 1.20 times as wide as pronotum; posterior margin near posterior angles weakly sinuate; puncturation weakly granulöse, more distinct and some what denser than that of pronotum. Hind wings fully developed. Legs long and slender, first metatarsomere approximately as long as the combined length of the three follo wing tarsomeres. Mesosternum with microsculpture similar to that in A. umbrosa, without median carina; mesosternal process acute.
Abdominal terga III-V with moderately deep, tergum VI without appreciable anterior impression; sterna III-V anteriorly constricted; puncturation fine, but distinct, denser and slightly coarser near anterior margin than on posterior part of terga; tergum X similar to that in the following species (cf. Fig. 56). c?: posterior margin of tergum VIII slightly convex, with a row of moderately long and a sparse row of long thin setae; posterior margin of sternum VIII distinctly convex, with a dense row of relatively short marginal and a sparse row of long submarginal thin setae; aedeagus with ventral process of median lobe distinctly separated, in ventral view slender, in lateral view almost straight, with small basal fold, weak crista apicalis, and with pronounced crista proximalis; internal tube with very long basal part (Figs. 44-45); apical lobe of paramere as in Fig. 46. 9: tergum VIII of similar shape and chaetotaxy as in <?; posterior margin of sternum VIII weakly convex, row of marginal setae very weakly modified, relatively thin, and very short; spermatheca as in Fig. 47.

Comparative notes
A. bonnairei is readily distinguished from A. umbrosa especially by larger size, lighter colour, the different shape of the head, the smaller and less prominent eyes, the more transverse antennomere VI, the more slender maxillary palpi, the presence of genal carinae, the microsculpture on head and pronotum, the more transverse and differently shaped pronotum, the shorter elytra, and the different primary and secondary sexual characters. For distinction from other species of Amarochara, especially the similar A. cribripennis (MULSANT & R ey), see the following species sections and the key in section 5. For systematic comments see remarks below the following species and section 3.3.

Distribution and bionomics
The area of distribution of A. bonnairei, evidently a very rare and local Ponto-Mediterranean element sensu LATTIN (1967), ranges from France and southern England in the west and northwest to southern Italy, the Balkans, and northern Anatolia in the south and southeast (Map 3). It seems to have been collected only once in Denmark and on very rare occasions in England. In France, it is known from several localities in the northern, eastern, central, and southeastern parts. In Germany, it has been recorded from the following regions: Bayern, Baden-Württemberg, Hessen, Rheinland-Pfalz, .. in the 'runs' leading from a n est... of Formica fuliginosa, at the root of an old beech tree". One of the beetles from Schles wig-Holstein was taken from a rotting alder trunk; see also LO H SE (1967). LIN K E (1962) found four specimens in black humus under grass roots. A considerable proportion of the specimens examined were collected by sweeping (May), at windows (July), and espe- daily in flight-intercept traps (May through July) (material examined and K Ö H LE R , pers. comm.). These observations suggest that .A bonnairei reproduces (and hibernates) in an unknown subterranean habitat and that it disperses by flight in late spring and early summer. The vast majority of the material examined was collected during the period from May through July; only one specimen was found in August (Fig. 51). According to HO RIO N (1967), the species has been found also April and September. Records from late autumn, winter, and early spring are unknown.

Comments
The publication year indicated by most previous authors is 1874 (see e. g. BERNHAUER & SCHEERPELTZ (1926)). According to BLACKW ELDER (1952), however, the work con taining the description of M. cribripennis was published in 1875. The original description is based on an unspecified number of syntypes from the "envi rons de Corte en Corse (collection Reveliere)", two of which were located in the collec tions of the MHNL. J. Vogel, Görlitz, had examined these syntypes and attached a lectotype label to one of them, but a lectotype designation has not been published. In order to fix a single name-bearing type and preserve the present interpretation of the species, the 9 in better condition is here designated as the lectotype.

Redescription
In size and many other external characters very similar to A. bonnairei. Colour of body variable; usual coloration: whole body brown to dark brown, with the hind margins of the abdominal segments and often the elytra and the antennae, at least partly, lighter brown; legs light brown; palpi testaceous. Occasionally, the body may be yellowish brown and of similar colour as in M. bonnairei, with only the head and the preapical abdominal segments somewhat darker. Microsculpture on head, pronotum, and abdomen absent or nearly so. Pubescence somewhat shorter and ± decumbent, distinctly less erect than in A. bonnairei. Head of ± ovoid shape, indistinctly (approximately 1.05 times) wider than long, not distinctly dilated behind eyes; width across eyes equal to width across postgenae; eyes moderately convex, larger than in A. bonnairei, distinctly more than half the length of postgenae in dorsal view. Puncturation very fine and sparse, similar to that in A. bonnai rei. Genal carina present. Antenna of similar morphology as in A. bonnairei, but longer and more massive (Fig. 57) Elytra on average relatively longer than in A. bonnairei, at suture 0.8 -0.9 times the length of pronotum and usually 1.10 -1.15 times as wide as pronotum; posterior margin near posterior angles weakly sinuate; puncturation distinctly denser and more granulose than in At. bonnairei. Hind wings fully developed. Legs long and slender, first metatarsomere very long, longer than in average v4. bonnairei, at least as long as the three following tarsomeres. Mesosternum with microsculpture similar to that in A. umbrosa, without median carina; mesosternal process apically acute. Abdominal terga III-V with moderately deep, tergum VI without appreciable anterior impression; sterna III-V anteriorly constricted; puncturation as in A. bonnairei sparser in posterior than in anterior parts of terga and sparser on posterior than on anterior terga, but on the whole much denser and more distinct, particularly so on terga III and IV; tergum X with long anterior extension (Fig. 56). cf: tergum and sternum VIII as in A. bonnairei, but marginal setae of sternum VIII longer; aedeagus of similar general morphology as in A. bonnairei, but ventral process of median lobe in ventral view broader, in lateral view subapically more distinctly curved, and with more distinct basal folds (Fig. 52-53); apical lobe of paramere similar to that in A. bonnairei (Fig. 54). ? : tergumVIII, sternum VIII, and spermatheca ( Fig. 55) similar to those in A. bonnairei.

Comparative and systematic notes
From the similar A. bonnairei, A. cribripennis is readily distinguished above all by the (mostly!) darker colour, the absence of microsculpture on head and pronotum, the longer and more massive antennae, the denser and coarser puncturation of the elytra and the abdo men (especially on terga III and IV), and by the morphology of the aedeagus. For di stinction from other West Palaearctic congeners see the species sections below and the key in section 5.
A. cribripennis is apparently the adelphotaxon of A. bonnairei. Both species are highly similar in external and sexual characters and share the following synopomorphies: abdo minal terga III-VI anteriorly with fine puncturation, ventral process of aedeagus basally with lateral carinae, and internal tube of aedeagus with very long basal part (see section 3.3).

Distribution and bionomics
A. cribripennis has a Tyrrhenian distribution sensu LATTIN (1967); its range is confined to Corsica and Sardinia (Map 4). It has been found by sifting leaf litter, under stones, and with pitfall traps at elevations of 900-1500m in February and during the period from April through June (material examined).

Description
Relatively large species, 3.8 -4.6 mm (abdomen fully extended). Colour of body variable: forebody uniformly rufous or bicoloured; head and pronotum rufous to dark brown, elytra rufous to light brown; abdomen dark brown to blackish, with the tergal margins and the abdominal apex lighter; legs and antennae testaceous to light brown; mouthparts testaceous. Forebody and most of abdomen without appreciable microsculpture, apical abdominal segments occasionally with traces of shallow transverse microsculpture. Pubescence of forebody relatively long and suberect to decumbent. Head of subcircular shape, approximately as wide as long; not or only weakly dilated behind eyes; eyes moderately large, weakly convex, not distinctly projecting from lateral outline of head, distinctly more than half the length of postgenae in dorsal view, the latter smoothly curved towards posterior constriction. Puncturation sparse and extre mely fine. Genal carina present, but fine. Antenna very long and massive, similar to that in the following species (cf. Fig. 66), antennomere I with dorso-apical furrow reduced, very short and inconspicuous; antennomere III as long as II. Third joint of maxillary palpus very long and slender (Fig. 61), similar to that in A. bonnairei. Pronotum large in relation to head, 1.15 -1.25 times as wide as long and approximately 1.4 times as wide as head or even wider; maximal width in or near the middle; posterior angles + ill-defined; puncturation similar to that of head. Elytra of similar relative length as in A. bonnairei, at suture usually 0.75 -0.80 times the length of pronotum and approximately 1.15-1.20 times as wide as pronotum; posterior margin near posterior angles noticeably sinuate, somewhat more so than in A. bonnairei (but distinctly less than in A. forticornis)\ puncturation weakly granulose, more distinct than in A. bonnairei and much more conspicuous than that of pronotum. Hind wings fully developed. Legs long and slender, first metatarsomere approximately as long as the combined length of the three following tarsomeres or slightly longer. Abdominal terga III-V with deeper anterior impressions than in A. bonnairei, tergum VI without or with very weak anterior impression; sterna III-V anteriorly constricted; punctu ration in impressions of terga III-V and near anterior margin of tergum VI very coarse and dense, on remainder of dorsal abdominal surface distinctly finer and sparser, punctures on abdomen on the whole coarser, deeper and larger than in A. bonnairei. d : posterior margin of tergum VIII moderately convex, with a row of moderately long and a sparse row of long thin setae; posterior margin of sternum VIII strongly convex, with a dense row of relatively long marginal and a sparse row of very long submarginal thin setae; aedeagus of similar general morphology as in A. bonnairei, with ventral pro cess distinctly separated from median lobe; at base of ventral process with remarkable median process (similar to the process of the crista apicalis in species of Geostiba T H O M SON s. str.), crista apicalis relatively small, crista proximalis pronounced; internal sac with long tube (Figs. 58-59). 9: tergum VIII of similar shape and chaetotaxy as in d , but with denser marginal setae; posterior margin of sternum VIII weakly convex, row of marginal setae very weakly modified, relatively thin, and short; spermatheca as in Fig. 60.
Derivatio nominis: The name (Lat., adj.: siculus = small dagger) refers to the distinctive median process at the base of the ventral process of the aedeagus.
Comparative notes and comments From A. bonnairei and A. cribripennis, this species is separated by larger size, a distinctly larger and more transverse pronotum (especially in relation to head), by more massive and longer antennae, by the much coarser puncturation of the anterior impressions of the abdominal terga III-V, and by the morphology of the aedeagus. A. siculifera is extre mely similar to the following species, from which it is distinguished with absolute cer tainty only by the shape of the aedeagus. Due to relatively high intraspecific variation in external characters and some overlap in the distinguishing characters indicated in the description of A. inermis sp. n., it may be difficult to attribute the 9 9 to either of the species.

Distribution and bionomics
The known distribution of the species, evidently a Syrian element sensu LATTIN (1967), ranges from central southern Anatolia to Lebanon (Map 5). Two of the types were collected in April and May. Apart from the altitude (1000-1200m) indicated on the label attached to the specimen from Mersin, no further bionomic data are available.

Description
Externally almost identical to A. siculifera, distinguished only by the following characters: Body on average more distinctly bicoloured, with pronotum and elytra yellowish brown to rufous and with head and most of abdomen dark brown to blackish. Antenna in most specimens with less transverse antennomeres III-V and with antennomeres IV and V less distinctly differing in length (Fig. 66). First metatarsomere longer than in average A. siculifera, usually longer than the combi ned length of the three following metatarsomeres (Fig. 67). c?: aedeagus with ventral process longer and less distinctly tapering apically, at base without long process (Figs. 62-63); apical lobe of paramere as in A. siculifera (Fig. 64). 9: shape of spermatheca very similar to that of A. siculifera (Fig. 65).

Derivatio nominis
The name (Lat., adj.: unarmed) refers to the fact that, in contrast to the otherwise extre mely similar A. siculifera, the aedeagus of A. inermis lacks the conspicuous median pro cess at the base of the ventral process.

Comparative and systematic notes
For distinction from A. siculifera, evidently the sister species of A. inermis, the descripti on above is referred to. For separation from other species of Amarochara see the compa rative notes below A. siculifera.

Distribution and bionomics
The species is known only from Israel, where the types were collected in February and April at low elevations of up to 300m (Map 5). The specimen from Yodefat was collec ted in Quercus macchia (SCHAWALLER, pers. comm.). Head of subcircular shape, indistinctly wider than long; not dilated behind eyes; eyes large (much larger than in A. bonnairei), moderately convex, slightly projecting from lateral outline of head (maximal width across eyes), almost as long as postgenae in dor sal view, the latter smoothly curved towards posterior constriction. Puncturation sparse and extremely fine. Genal carina present, but fine. Antenna longer and more massive than in A. bonnairei; antennomere III as long as II, IV and V short and strongly transverse, IV -X not gradually increasing in length (as is the case in At. bonnairei), but V distincdy (approxi mately 1.5 times) longer and less transverse than IV, and V -X almost subequal in length; antennomere I with dorso-apical furrow reduced, very short and inconspicuous. Third joint of maxillary palpus very long and slender, similar to that in A. bonnairei. Pronotum larger in relation to head than in A. bonnairei, 1.14 times as wide as long and approximately 1.3 times as wide as head; maximal width approximately in the middle; posterior angles weakly marked; puncturation very fine and sparse, barely noticeable in the microsculpture. Elytra relatively longer than in A. bonnairei, at suture usually 0.84 times the length of pronotum and 1.17 times as wide as pronotum; posterior margin near posterior angles as in A. bonnairei weakly sinuate; puncturation indistinctly granulose, somewhat denser and slightly coarser than in A. bonnairei, and much more conspicuous than that of pronot um. Hind wings fully developed. Legs long and slender, first metatarsomere approximately as long as the combined length of the three following tarsomeres. Abdominal terga III-V with impressions of similar depth as in A, bonnairei, tergum VI without anterior impression; sterna III-V anteriorly constricted; puncturation in im pressions of terga III-V and near anterior margin of tergum VI rather coarse and dense, much coarser and denser than in A. bonnairei, on remainder of dorsal abdominal surface distinctly finer and sparser, but slightly more conspicuous than in A. bonnairei. <S: unknown. ¥: tergum VIII posteriorly moderately convex and with row of rather long thin margi nal setae; posterior margin of sternum VIII weakly convex, row of marginal setae al most unmodified, relatively thin, and moderately short; spermatheca as in Fig. 68.

A m a r o c h a r a c a e c a
Derivatio nom inis: The name (Lat., adj.: dark) refers to the distinctive uniformly dark colour of the species.

Comparative notes and comments
From other Western Palaearctic congeners, A. caeca is distinguished especially by its uniformly dark colour and by the distinct micro sculpture of the head and pronotum, which it only shares with A. bonnairei. For additional characters separating it from the latter species and from the similarly dark A. splendens see description above and section 4.6, respectively.

Distribution and bionomics
The species is currently known only from Tunisia. Data other than those indicated on the labels attached to the two type specimens are not available.

Redescription
Of similar size as A. forticornis (see section 4.3.1). Head dark brown, pronotum brown, elytra yellowish brown; abdomen dark brown with lighter tergal margins; antennae brown, with slightly lighter basal antennomeres; legs and mouthparts testaceous. Head and pro notum with very shallow, but noticeable microsculpture. Pubescence of forebody simi lar to that in A. forticornis. Head of subcircular shape, indistinctly transverse; eyes large, though not strongly con vex, slightly more distinctly projecting from lateral outline of head than in A. forticornis, their diameter almost as long as postgenae in dorsal view. Puncturation finer, shallower, and less distinct than in A. forticornis. Genal carina present. Antenna long and massive, of similar morphology as in A. forticornis, but antennomeres IV and V less transverse (Fig. 69). Mouthparts similar to those in A. forticornis. Pronotum relatively large and broad (width: 0.74 mm), 1.18 times as wide as long and 1.4 times as wide as head; maximal width approximately in the middle; posterior angles weakly marked; puncturation similar to that of head, but denser. Elytra relatively longer and broader than in A. forticornis-, at suture 0.85 times the length of pronotum and approximately 1.1 times as wide as pronotum; posterior margin near posterior angles weakly sinuate; puncturation much finer than in A. forticornis and not granulöse. Legs very long and slender; length of metatibia and metatarsus (without claws) 0.72 mm and 0.63 mm, respectively; first metatarsomere slightly longer than the combi ned length of the three following tarsomeres (Fig. 70). Mesosternum without median carina; mesosternal process apically rounded, almost acute. Abdominal terga III-V with anterior impressions of similar depth as in A. carinata (see section 4.3.2), distinctly less deep than in A. forticornis', puncturation of terga III-V simi lar to that in A. carinata (much denser than in A. forticornis), only slightly coarser and denser in anterior impressions than on remainder of dorsal surface. Sexual characters missing in the holotype.

Comparative notes
From A. forticornis, the species is readily distinguished especially by the larger eyes, the larger and broader pronotum, the presence of shallow microsculpture on head and pro notum, the weakly sinuate posterior margins of the elytra, the much finer elytral punctu ration, the longer legs, the shallower transverse impressions on the abdominal terga III-V, and the puncturation of the abdomen, which is finer in the anterior impressions of terga III-V and much denser on the posterior parts of these terga. A. carinata, which is similar in coloration and many other external characters, differs from A. crassicornis in the morphology of the antennae (shorter antennomere III, much more transverse an tennomeres IV-X, antennomere V approximately 3 times as wide as long), in the much smaller eyes, the absence of microsculpture on head and pronotum, the smaller and less transverse pronotum, the relatively shorter elytra, the distinctly shorter legs, the presence of a carina on the mesosternum, and the slightly less dense abdominal puncturation.

Distribution and bionomics
A. crassicornis has become known only from Morocco, Algeria, and Tunisia (CO IFFAIT, 1973, N o r m a n d , 1934, Qued e n fe ld t , 1882). Literature records from the eastern Mediterranean refer to either A. inermis or A. siculifera. The record from Sicily (PO RTA, 1926, Z a n e t t i , 1995) should be considered doubtful until the corresponding specimen has been revised. Neither the original description nor the labels attached to the holotype reveal any data regarding the circumstances of collection.

The f o r t i c o r n i s group
Primarily based on the presence of a median carina on the mesosternum, two species are here tentatively attributed to this group. However, as is the case with the bonnairei group, the monophyly of A. forticornis and A. carinata is only weakly supported, especial ly because the remaining synapomorphies also occur in other Amarochara and because the male sexual characters of A. carinata as well as of some species of the bonnairei group are unknown (for further details see section 3.3). In addition, A. forticornis is characterized by several distinct autapomorphies, especially the presence of a distinct anterior impres sion on abdominal tergum VI and the posteriorly strongly sinuate elytra.

A m a r o c h a r a f o r t i c o r n i s ( L a c o r d a i r e , 1835)
Figs. 71-77, 80-81, Map 6 Bolitochara forticornis LACORDAIRE, 1835: 543.

Material examined
Germany: Schleswig-Holstein: 1 e x , H usum, leg. Varendorff (NHMW). Niedersachsen: 3 d c?, 1 9 , Hannover, Langenhagen, ditch, pitfall, 17.-31.V.1991, leg. Sprick (cAss); 3c? d , 2 9 9 , same data, but 1.-30.V I.1991 (cAss, cWun)  Redescription 3.0 -5.0 mm (abdomen folly extended). Size and size-related characters subject to considerable intraspecific variation. Colour of body highly variable, too; usual coloration: head and abdominal segments III-VII, except for the hind margins, dark brown to blackish; elytra, hind margins of abdominal segments III-VII, and abdominal apex ferrugineous; pronotum ferrugineous to dark brown, in most specimens ± castaneous; antennae (except for the lighter basal antennomeres) ± castaneous; legs and mouthparts yellowish to yellowish brown, sometimes with the maxillary palpi partly infúscate; occasionally whole body of similar colour as infoL umbrosa, apart from the legs and mouthparts ± completely dark brown to blackish, or of similar colour as A. bonnairei, with the whole body ± uniformly dark testaceous, ferrugineous, or yellowish brown. Head, pronotum, elytra, and most of abdomen usually without appreciable microsculpture; abdominal terga VII and VIII with very shallow transverse microsculpture. Pubescence of forebody relatively long and suberect. Head of subcircular shape, approximately as wide as or slightly wider than long; not or only indistinctly dilated behind eyes, eyes moderately large and convex, weakly projec ting from lateral outline of head, their diameter approximately half the length of postgenae in dorsal view or slightly larger, the latter smoothly curved towards posterior constriction or ± subparallel. Puncturation moderately fine and sparse, more distinct than in A. umbrosa. Genal carina distinct. Antenna long and massive, antennomere I with dorso-apical furrow of somewhat variable length and depth; antennomere III approxi mately as long as antennomere II (Fig. 77). Third joint of maxillary palpus very long and slender, similar to that in A. bonnairei (Fig. 76); labrum of similar shape of chaetotaxy as in A. bonnairei', labium as in Fig. 75. Pronotum approximately 1.1 times as wide as long and 1.25 -1.35 times as wide as head; maximal width in or a short distance anterior to middle; puncturation shallow, ± similar to that of head or finer. Elytra at suture in most specimens 0.80 -0.85 times the length of pronotum and approxi mately 1.20 times as wide as pronotum; posterior margin near posterior angles strongly sinuate; puncturation granulóse, relatively dense and coarse, near scutellum often ± rugose. Hind wings fully developed. Legs long and slender, first metatarsomere approximately as long as the combined length of the three following tarsomeres or only slightly shorter. Mesosternum with median carina of variable length; sometimes only a short rudiment of such a carina present near anterior margin; mesosternal process apically acute. Abdominal terga III-V with very deep, tergum VI with moderately deep anterior im pression; sterna III-VI anteriorly constricted, this constriction more pronounced on sterna III-V than on sternum VI; puncturation in anterior impressions of terga III-VI dense and coarse, on remainder of dorsal surface fine and sparse; tergum X as in Fig. 79. <$: posterior margin of tergum VIII weakly convex, with a row of rather long thin marginal setae; posterior margin of sternum VIII distincdy convex, with a dense row of moderately long marginal and a sparse row of very long submarginal thin setae; aedeagus with ventral process of median lobe not distinctly separated, in lateral view weakly bent, and with distinct basal fold; crista apicalis of reduced, crista proximalis of moderate size; internal sac with relatively short tube (Figs. 71-72); apical lobe of paramere relatively slender and with external fold-like structure (Fig. 73). ? : tergum VIII of similar shape and chaetotaxy as in c? ; posterior margin of sternum VIII weakly convex, row of marginal setae very weakly modified, relatively thin, and very short; spermatheca as in Fig. 74.

In trasp ecific variation
Coloration and all size-related parameters may vary considerably. Nanistic specimens were seen from various regions, especially from the Balkans. The shape of the posterior margin of the elytra and the depth of the anterior impression of the abdominal tergum VI, too, are subject to some intraspecific variation.

C om m ents
The original description is based on an unspecified number of syntypes ("Rare. Fontaine bleau"). Repeated requests for a loan of the types, which are deposited in the Muséum National d'Histoire Naturelle Paris, have remained unanswered.
C om parative notes AE forticornis is distinguished from all congeners occurring in the Holarctic region espe cially by the presence of a distinct anterior impression on the abdominal tergum VI, by the strongly sinuate posterior margin of the elytra, and by the male primary sexual cha racters (morphology of the median lobe of the aedeagus and of the apical lobe of the paramere. From all species, except A. carinata, it is additionally separated by the presence of a median carina on the mesosternum.
Distribution and bionom ics LikeM. bonnairei, A. forticornis is apparently a Ponto-Mediterranean element (see LATTIN, 1967), its distribution ranging from (northern?) Spain, France, southern England, southern Scandinavia, and Estonia in the west, northwest, and north, to southern Italy, southern Greece, and Ukraine in the south, southeast, and east (Map 6) (material examined and F o w le r , 1888, H a b e rm a n , 1983, H o r io n , 1938, J o y , 1932, L u ig io n i, 1929, L u n d -BERG, 1995, SlLFVERBERG, 1992, ZANETTI, 1995. The species is also indicated from the Caucasus region and from Turkey (BerNHAUER, 1902, HORION, 1967, but these records require confirmation. O nly two specim ens w ere seen from Spain; the precise locality, however, is not specified on the labels. OlJTERELO (1980) reports the species from southern Spain (Sierra de Cazorla, under pine bark), but this record is based on a m isidentification and in fact refers to a species o f Phloeopora ERICHSON (specim en examined). In France, the species is apparently w idespread, but rare (m aterial exam ined; HORION, 1967); it has also been reported from Corsica (LUIGIONI, 1929, SAINTE-CLAIRE DEVILLE, 1914. In E ngland and Scandinavia, its known distribution is confined to the south (FOYTER, 1888, HORION, 1967, JOY, 1932, Lundberg, 1995, PALM, 1972. It has becom e known from various regions in D enm ark (HANSEN, 1964, HANSEN, 1996 and was rather recently discovered also in E stonia (HABERMAN, 1983).
In Germany, it is present in almost all regions (see material examined) and here also recorded from Mecklenburg-Vorpommern, from where it was previously unknown (KÔH- A. forticornis is here -apparently for the first time -recorded from Switzerland; neither HO RION (1967) nor LUCHT (1987) indicate any Swiss records. Material from eastern and southeastern Austria (Niederösterreich, Steiermark, Burgenland) was examined; H O R I-  and SCHEERPELTZ (1968) also report the species from Vorarlberg, Ober österreich, and Kärnten, but the corresponding specimens were not found in the Scheerpeltz collection, where they are supposedly kept (H O RIO N , 1967). In Italy, it is distribu ted from the north to the extreme south, including Sardinia and Sicily (material exami ned; L u i g i o n i , 1929, PO RTA , 1926, Z a n e t t i , 1995).
A. forticornis has repeatedly been recorded from the Czech Republic, Slovakia, Hungary, and Romania; for additional records see ÄDÄM (1987), BöHÄC et al. (1993), HORION (1967), KOCIAN (1993), VÄSÄRHELYI (1985). Records from the Balkans are relatively scarce (see m aterial exam ined and COIFFAIT (1976), NOVAK (1952), and SCHEERPELTZ (1959).  As in the case of A. umbrosa and A. bonnairei, the species was observed in a wide range of biotopes, but reproduction and hibernation take place in an unknown subterranean habitat. Based on the material examined (additional literature data given in brackets), the species was found in various forest types (flood-plain, mixed, and deciduous woodland), on or near banks of ponds, lakes, and rivers (HORION, 1967), in ditches, in xerothermous grassland (ASSING 1994a, HORION, 1967, on one occasion in the alpine region near a snowfield, several times in arable land (HORION, 1967), vineyards, in sand and gravel pits (FOWLER, 1888, KÖHLER, 1997, and often also in urban habitats (fallows, gardens, etc.) (ASSING, 1988, KACHE, 1998, STRZELCZYK, 1988. The vast majority7 of records are from lower elevations, but one Greek specimen was taken at an altitude of almost 1800 m. A. forticornis has been observed throughout the year, with a maximum in spring (Fig. 80).
In spring and summer, the species is usually collected on the ground surface by pitfall trapping or by manual sampling methods, whereas nearly all the late autumn and winter records (November through March) are from flood debris, suggesting that the beetles were washed out of their cryptic hibernation habitat. Some winter records are also from ant nests (Lasiusplatythorax, E. brunneus, L. fuliginosus) and from mole nests; for additional data see HO RION (1967) and O SELLA & ZAN ETTI (1975). Remarkably, one specimen was found in December using a floating technique (soil-washing) used for collecting endogean beetle species (see material examined). Pitfall trap studies showed that epigeic activity7 begins in March and ends in summer, with a maximum from April through June (Fig. 81). From the observations that the species may be caught in greater numbers by pitfall trapping in spring, but is rarely found by sifting or other manual samp-ling tech niques or during other seasons of the year, it can be inferred that the beetles are epigeically active only to search for suitable (subterranean) reproduction habitats and/or for mates. Dispersal activity by flight was recorded with malaise traps and car-nets in May and August (material examined; KÖHLER, pers. comm.). However, no flight muscles were found in four dissected beetles collected by pitfall trapping, suggesting that A. forticornis may be dimorphic regarding its flight-muscle development. One female caught in April had a mature egg in the ovaries; teneral adults were observed in August. Description 4.2 mm (abdomen fully extended). Pronotum, elytra, legs, mouthparts, clypeus, anterior part of frons, antennomeres I and II, margins ot abdominal terga, and abdominal apex yellowish brown to rufous; apical part of antenna brown; posterior part of head dark brown; abdominal terga III-VII, except for the lighter margins, blackish. Head, pronotum, elytra, and most of abdomen without appreciable microsculpture; traces of shallow microsculpture present only on the preapical abdominal terga. Pubescence of forebody shorter, less erect, and less conspicuous than in A. forticornis.

A m a r o c h a r a c a r i n a t a
Head of subcircular shape, approximately as wide as long; weakly dilated behind eyes; eyes slightly smaller than in A. forticornis, weakly projecting from lateral outline of head, their diameter approximately half the length of postgenae in dorsal view, the latter more distincdy convex than in A. forticornis, smoothly curved towards posterior constriction. Puncturation finer, shallower, and less distinct than in A. forticornis. Genal carina present. Antenna long and massive, of similar morphology as in A. forticornis, but antennomere III wider and more strongly dilated apically, antennomere IV slightly wider and more transverse, V wider, more transverse, and only slightly longer than IV; antennomeres VI -X more transverse and wider than in A. forticornis, X approximately 1.5 times as wide as long, and XI longer than the combined length of IX and X. Mouthparts similar to those in A. forticornis. Pronotum 1.13 times as wide as long and 1.3 times as wide as head; maximal width a short distance anterior to middle; puncturation fine, more distinct than that of head, and more clearly defined than in average A. forticornis. Elytra shorter and narrower in relation to pronotum than in A. forticornis-, at suture 0.74 times the length of pronotum and approximately 1.1 times as wide as pronotum; posterior margin near posterior angles weakly sinuate; puncturation slightly granulose, somewhat denser than in A. forticornis, near scutellum denser, but not rugose. Legs long and slender; metatarsi longer and more slender than in A. forticornis, first metatarsomere distinctly longer than the combined length of the three following tarsomeres. Mesosternum with median carina; mesosternal process apically acute.
Abdominal terga III-V with anterior impressions of similar depth as in A. bonnairei, much less deep than in A. forticornis-, tergum VI with shallow anterior impression; sterna III-VI anteriorly constricted, this constriction more pronounced on sterna III-V than on ster num VI; punctures in anterior impressions of terga III-VI distinct and larger than in A. bonnairei, but much less coarse than in A. forticornis-, puncturation on posterior parts of terga III-VII, however, distinctly denser and coarser than in A. forticornis-, tergum X as in Fig. 79. dk unknown. 9: posterior margin of tergum VIII moderately convex and with row of relatively long setae; posterior margin of sternum VIII, with row of short and moderately modified marginal setae; spermatheca as in Fig. 78.

Derivatio nominis:
The name (Lat., adj.) refers to the presence of a median carina on the mesosternum, which distinguishes the species from all Palaearctic and Nearctic con geners, except for A. forticornis.

Comparative notes
From A. forticornis, the species is readily distinguished especially by the lighter coloration of the forebody, the wider and more transverse antennomeres, the relatively shorter and narrower elytra, the weakly sinuate elytra! hind margin, the longer metatarsomere I, the distinctly shallower anterior impressions of the abdominal terga III-VI, and by the com pletely different puncturation of the abdomen (see description above). From the simi larly coloured A. bonnairei, it differs in the larger size, the much more massive antennae, the presence of a mesosternal carina, the relatively larger and flatter pronotum, the much longer metatarsomere I, and by the much coarser abdominal puncturation.

D istribution and bionomics
The species is known only from one locality in northern Spain, where it was collected at an altitude of 1900 m.

The loebli group
Species of the loebli group occur in the Eastern Palaearctic region and have been recorded from the Himalayas, China, and Taiwan. The monophyly of this group is secured by several significant synapomorphies, i. e. a posteriorly constricted head with a distinct neck, broad connecting joints between the antennomeres IV-XI, a short ventral process of the median lobe of the aedeagus, and by the derived morphology of the apical lobe of the paramere. In addition, they share the following characters, some of them synapomorphic, but shared also with other species groups, and some plesiomorphic: genal carinae absent, eyes relatively large, antennomere III shorter than II, third joint of maxillary palpus long and slender, ligula only apically bifid, long and slender mandibles, pronotum slender and with relatively sparse puncturation, mesosternum without carina, mesosternal process apically broadly truncate, tarsi long and slender, anterior impressions of abdominal terga III-V deep and coarsely punctate, posterior parts of abdominal terga IIEVI sparsely punctate, abdominal tergum X with numerous stout setae also in the median area and with relatively short anterior extension, internal sac of aedeagus with distinctly sclerotized internal structures (see also section 3.3). Redescription 2.9 -3.3 mm. Head dark brown; pronotum brown, elytra slightly lighter, yellowish brown to light brown; abdomen brown, with the tergal margins and the abdominal apex lighter; antenna dark brown, with antennomeres I-II or I-III light brown to brown; legs yellowish brown to brown. Head with extremely shallow, barely noticeable microsculpture, apical abdominal terga with very fine and shallow transverse microstriae; remainder of dorsal surface without microsculpture and shining. Pubescence of forebody moderately long and + suberect. Head of subcircular shape, slightlv wider than long, not dilated behind eyes; posteriorly with distinct and slender neck of approximately half the width of head; eyes large and convex (but less so than in A. umbrosd), weakly projecting from lateral outline of head, their diameter slighdy shorter than postgenae in dorsal view, the latter smoothly curved towards posterior constriction, posterior angles ± obsolete. Puncturation extremely fine and sparse, barely noticeable. Genal carina absent. Antenna massive, antennomere I with dorso-apical furrow of somewhat variable length and depth, similar to that in A. umbrosa; antennomere III slightly shorter than II; connecting joints between antennomeres broad, more so than in W-Palaearctic congeners (Fig. 90). Maxillary palpi of simi lar morphology as in A. umbrosa, but fourth joint slightly longer (Fig. 89); ligula apically rounded, not distinctly bifid; apical joint of labial palpi very long and slender (Fig. 88); mandibles very long and slender, more so than in A. umbrosa. Pronotum approximately 1.10 -1.15 times as wide as long and 1.24 -1.31 times as wide as head; maximal width in or a short distance anterior to middle; puncturation shallow, fine, and ill-defined. Elytra at suture 0.85 -0.90 times the length of pronotum and approximately 1.25 -1.30 times as wide as pronotum; posterior margin near posterior angles not distinctly sinuate; puncturation fine (but more distinct than that of head and pronotum), indistinctly granulose, somewhat sparser than in A. umbrosa. Legs relatively longer than in A. umbrosa-, first metatarsomere at least slightly longer than the combined length of the two following tarsomeres, sometimes as long as the combined length of the three following tarsomeres. Mesosternum without median carina; mesosternal process apically broadly truncate (much broader than in A. umbrosa).

A m a ro c h a ra loebli P a c e , 1992
Abdominal terga III-V with deep (slightly deeper and more well-defined than in A. umbrosd), tergum VI without anterior impression; sterna III-V anteriorly constricted; puncturation in anterior impressions of terga III-V and near anterior margin of tergum VI very dense and coarse, on remainder of tergal surface very fine and sparse. Tergum X anteriorly not distinctly elongated, narrowly truncate (Fig. 91); with very dense stout setae. d : posterior margin of tergum VIII truncate, with a row of moderately long thin mar ginal setae; posterior margin of sternum VIII convex, with a dense row of moderately long and thin marginal setae and with interspersed long thin submarginal setae; aedeagus with short, slender, and, in lateral view, weakly bent ventral process; at base of ventral process with distinct median process; crista apicalis reduced, crista proximalis long, but not very prominent; internal sac with short tube and with additional pairs of long, mo derately sclerotized structures 86); paramere strongly modified, apical lobe of paramere separated from basal part of paramerite by membrane, strongly bent, flatte ned, with torsion, and apically broadly rounded (Figs. 84-85). 9: tergum VIII of similar shape and chaetotaxy as in d ; posterior margin of sternum VIII weakly convex, row of marginal setae unmodified, thin, and short; spermatheca as in Fig. 87.

A m a ro c h a ra se rie p u n cta ta sp. n.
Map 7 <$: unknown. 9: posterior margin of tergum VIII distinctly convex and with longer marginal setae; posterior margin of sternum VIII more strongly convex; a spermatheca was not found in the apparently teneral holotype.

Derivatio nom inis
The name (Lat., adj.) refers to the distinctive transverse series of dense and coarse punctures on the abdominal terga III-V

Comparative notes
A. seriepunctata is separated from all other species of the Holarctic region by the charac teristic puncturation of terga III -V For additional characters distinguishing it from A. loebli and other Eastern Palaearctic congeners see description above and the comparative notes in the following species sections, respectively. The phylogenetic affiliations to the other species of the loebli group are uncertain, particularly because the male sexual cha racters are unknown.

Distribution and bionomics
The species is known only from the type locality in eastern Nepal, where it was collected in April at an altitude of only 1100m.

Description
3.2-3.9 mm. In external appearance highly similar to A. loebli, but distinguished as fol lows: Coloration as in darkM. loebli. Antenna equally massive, but somewhat longer; antennomere I with dorso-apical furrow of variable length. Aleso-and metasternal process broadly truncate (holotype) or very narrowly truncate, almost acute (paratype). First metatarsomere shorter than the combined length of the three following tarsomeres. Anterior impressions of abdominal terga III-V in the holotype slightly more pronounced, wider, and with more and slightly coarser punctures. Tergum X with less dense and less stout setae.
3: posterior margin of tergum VIII almost truncate and with row of rather long margi nal setae; sternum VIII posteriorly strongly convex, almost pointed, and with dense row of relatively long thin setae; aedeagus of similar general morphology as in A. loebli, ventral process of median lobe short in relation to median lobe and in ventral view of triangular shape; at base of ventral process with weak and obtuse median process; crista apicalis reduced; crista proximalis very long and prominent; internal sac with several pairs of large and distinctly sclerotized structures; internal tube relatively short ; paramere as in A. loebli strongly modified (Fig. 94); apical lobe of paramere flattened, transverse, and apically truncate (Fig. 95). 9: unknown.

Derivatio nominis
The name (Lat., adj.) refers to the distinctive set of sclerotized structures in the internal sac.

Intraspecific variation, comparative notes, and phylogenetics
The holotype is much larger, somewhat darker, has longer first metatarsomeres, and, above all, has an apically broadly truncate meso-and metasternal process, whereas in the paratype, the meso-and metasternal process is almost acute. In both specimens, however, the -highly derived and very distinctive -morphology of the aedeagus is identical, so that they are here attributed to the same species. The paratype is apparently a nanistic speci men, the reduction in size presumably also involving changes in other external charac ters. Based on external characters alone, this species is difficult to distinguish from A. loebli (see description above). The morphology of the aedeagus, however, is highly distinctive and separates this species from all known congeners. The similarly highly derived mor phology of the median lobe of the aedeagus and of the paramera suggest that A. armata and A. loebli are adelphotaxa.

Distribution and bionomics
The species is known from Shaanxi and Sichuan (China) (Map 8), where it was collected at altitudes between 1300 and 2600m. The holotype was sifted in a damp degraded deciduous forest near a stream (SCHULKE, pers. comm.). Antenna conspicuously massive, more so than in A. loebli, antennomeres IV-X wider and more transverse, IV and V almost disc-like and approximately 4 times as wide as long; antennomere I with distinct dorso-apical furrow. First metatarsomere approxi mately as long as or slightly shorter than the combined length of the three following tarsomeres.

A m a r o c h a r a w r a s e i
Abdomen as in A. loebli. <5: posterior margin of tergum VIII weakly convex and with row of rather long margi nal setae; sternum VIII posteriorly convex and with moderately dense row of relatively long thin setae; aedeagus with ventral process longer in relation to median lobe than in A. loebli, at base of ventral process without distinct median process; crista apicalis reduced; crista proximalis long, but not very prominent; internal sac with distinctive apical pair of sclerotized structures; internal tube of similar length as in A. loebli ; paramere of similar general morphology, but less strongly modified than in A. loebli', apical lobe of paramere flattened, oblong, slightly curved, and apically rounded (Fig. 98). 9: tergum VIII posteriorly almost truncate and with row of sparse and rather long thin marginal setae; posterior margin of sternum VIII weakly convex, with row of dense and short, distinctly modified marginal setae and with interspersed long thin setae; spermatheca with longer duct than in A. loebli (Fig. 99).

Derivado nominis
The species is dedicated to David W Wrase (Berlin), specialist of Carabidae, who collected two of the types.

Comparative notes
From the similar A. loebli and Vt. armata, A. wrasei is distinguished by the more transverse head, the more massive antennae (more strongly transverse antennomeres IV-X), and especially by the morphology of the aedeagus.

Distribution and bionomics
The types were sifted in southern Sichuan (Map 8) and in southern Shaanxi at an altitude of 1250 and 1700m. Pronotum with maximal width clearly in anterior half; posterior angles obtuse, but welldefined; dorsal surface with or without very shallow traces of microsculpture; pubes cence somewhat longer and more erect than in A. loebli. Pubescence of elytra denser and more distinct. Legs longer and stouter than in A. loehli. First metatarsomere as long as the combined length of the three following tarsomeres or nearly so. Anterior impressions of terga III-V deeper and broader than in ML loebli and with more and coarser punctures (similar to A. armata); punctures in posterior areas of terga III-V coarse and very distinct, much coarser than in A. loebli, sometimes forming irregular transverse rows. c?: posterior margin of tergum VIII weakly convex, almost truncate, and with r o w o f rather long and dense marginal setae; sternum VIII posteriorly moderately convex and with dense row of relatively long thin setae; aedeagus with ventral process of median lobe longer than in A. loebli, but less slender, and at base without median process; crista apicalis and crista proximalis small; internal tube relatively shorter and other internal structures of different shape (Figs. 100-101); paramere less modified than in A. loebli, apical lobe of paramere less flattened, not distinctly bent, and more slender (Fig. 102). 9: tergum VIII posteriorly weakly convex, almost truncate, and with ro w o f d e n s e and rather long thin marginal setae; posterior margin of sternum VIII moderately convex, with row of dense and short, weakly modified marginal setae and with interspersed long thin setae; spermatheca larger and with longer duct than inM . loehli (Fig. 103).

Derivatio nominis
The name refers to the conspicuously large eyes of this species. C om parative notes From all other congeners with a distinct neck, this species is readily distinguished espe cially by the large eyes, the coarse puncturation of the posterior half of the abdominal terga III-V, and by the morphology of the aedeagus.

Distribution and bionomics
The fact that the types were collected in Taiwan and in Nepal (Map 8) suggests thatM. megalops is very widespread in the Eastern Palaearctic region. The specimens were collec ted in April and May at altitudes of 1650 and 2050 m.

The f o r m o s a n a group
The formosana group currently includes two species from Taiwan and Japan, wdiich share the following synapomorphies: head and pronotum with distinct microsculpture, a short and dense pubescence of the forebody, abdominal terga III-VI in posterior half densely punctate. There are probably additional synaporphies in the morphology of the aedeagus, but the male of A. densepunctata is unknown. The formosana group is linked to the loebli group, presumably its adelphotaxon, by several derived characters: antennomere III shorter than II, ligula only apically bifid, mesosternal process broadly truncate, aedeagus with distinctly sclerotized internal structures (see also section 3.3).

Description
Relatively large species, 3.7 -4.1 mm. Body blackish brown to blackish; posterior mar gins of abdominal terga and legs brown to dark brown; elytra dark yellowish brown. Head and pronotum with distinct, elytra with shallower microreticulation; posterior parts abdominal terga III-VI with very shallow and indistinct, terga VII and VIII with more distinct reticulation. Pubescence of forebodv relatively short, but very dense and erect. Head of + subquadrate shape, slightly (approximately 1.10 times) wider than long, not dilated behind eyes; posteriorly without neck, posterior angles weakly marked; eyes large and convex, distinctly projecting from lateral outline of head, their diameter approxima tely as long as postgenae in dorsal view. Puncturation extremely fine, barely noticeable in the microreticulation. Genal carina present, but very fine. Antenna massive and long, antennomere I with pronounced and long dorso-apical furrow; antennomere III shorter than II, antennomere V moderately transverse (less so than in most congeners), connec ting joints between antennomeres relatively narrow (Fig. 110). Third joint of maxillary palpi very long and slender (Fig. 109); ligula as in Fig. 108; anterior margin of labrum in the middle with two very short stout sensilla; mandibles very long and slender, right mandible with indistinct tooth and weakly serrate in dorsal molar region. Pronotum 1.00 -1.05 times as wide as long and approximately 1.2 times as wide as head; maximal width approximately halfway between anterior angles and middle; lateral outli ne near posterior angles weakly concave or straight, posterior angles well-defined; puncturation extremely dense, more distinct than that of head. Elytra at suture 0.80 -0.85 times the length of pronotum and approximately 1.4 times as wide as pronotum; posterior margin near posterior angles indistincdy sinuate; puncturation fine and dense. Legs long and slender; first metatarsomere as long as the combined length of the three following tarsomeres or nearly so. Mesosternum without median carina; mesosternal process apically broadly truncate.
Abdominal terga III-V with moderately deep and broad, tergum VI without anterior impression; sterna III-V anteriorly constricted; puncturation in anterior impressions of terga III-V and near anterior margin of tergum VI very dense and coarse; puncturation of terga III-V on remainder of tergal surface finer than in anterior impression, but rather coarse and dense (much more so than in A. loebli), interstices on average narrower than punctures; puncturation of terga VI and VII finer and sparser, but still very di stinct. Tergum X anteriorly not distinctly elongated, broadly truncate (Fig. I ll ) ; with very dense, stout, long and dark setae on all of its surface. d : posterior margin of tergum VIII truncate, with a dense row of long thin marginal setae; posterior margin of sternum VIII convex with a very dense ro w of long and thin marginal setae and with interspersed long thin submarginal setae; median lobe of aedeagus with long and slender ventral process, at base of ventral process with pair of weakly prominent lateral folds; crista apicalis and crista proximalis present, but rather small; internal sac with very long tube and pair of weakly sclerotized apical structures of di stinctive shape (Figs. 104-105); paramere unmodified, apical lobe of paramere broadly coniform and apically acute (Fig. 106). 9: tergum VIII of similar shape and chaetotaxy as in d ; posterior margin of sternum VIII wreakly convex, row of marginal setae unmodified, thin, and short; spermatheca relatively small, with rather short, wide, and apically distinctly bent duct (Fig. 107).

Derivatio nom inis
The name (Lat., adj.) is derived from Formosa, the former name of Taiwan.
Comparative notes and comments A. formosana is distinguished from all its congeners, except for A. densepunctata (see the following species section), by a number of characters, especially the densely punctured pronotum, the distinct microsculpture of the whole forebody, the short, dense, and erect pubescence, the densely and coarsely punctured abdominal terga III-V, and by the primary sexual characters in both sexes.

Description
In size (3.7 mm), coloration, and general appearance somewhat resembling A. inermis. Head and antenna dark brown with the basal antennomere slightly lighter, pronotum, legs, and abdomen (except for the lighter tergal margins and abdominal apex) brown; elytra slightly lighter brown; maxillary palpi testaceous. Microreticulation on head weak, on pronotum very weak (visible only at high magnifications), on elytra absent, and on abdomen very shallow. Pubescence of forebody relatively short and dense, weakly erect, almost decumbent. Head of subcircular shape, approximately as wide as long, not dilated behind eyes; po steriorly without neck, posterior angles weakly marked; eyes moderately large and wea kly convex, weakly projecting from lateral outline of head, their diameter slightly shorter than postgenae in dorsal view Puncturation very fine and rather sparse. Genal carina pro nounced. Antenna massive and long, antennomere I with pronounced and long dorsoapical furrow; antennomere III slightly shorter than II, antennomeres IV-X of similar proportions as in A. formosana, but IV and V more transverse, and X less transverse and longer than IX, connecting joints between antennomeres relatively narrow. Maxillary pal pi, labium, and labrum of similar morphology as in A. formosana (Fig. 113); mandibles apically less long and acute than in A. formosana', right mandible without tooth. Pronotum 1.08 times as wide as long and approximately 1,4 times as wide as head; maximal width before the middle; lateral outline near posterior angles ± straight, pos terior angles rather well-defined; puncturation very fine, more ill-defined and much den ser than that of head. Elytra at suture 0.82 times the length of pronotum and approximately 1.2 times as wide as pronotum; posterior margin near posterior angles weakly, but noticeably sinuate; punctu ration slightly granulose, very dense, and much more distinct than that of pronotum. Legs long and slender; first metatarsomere very long, longer than the combined length of the three following tarsomeres. Mesosternum without median carina; mesosternal process apically broadly truncate. Anterior impressions of abdominal terga III-V of similar depth and breadth as in A. formosana, tergum VI without anterior impression; sterna III-V anteriorly constricted; puncturation in anterior impressions of terga III-V and near anterior margin of tergum VI very dense and coarse; puncturation on remainder of dorsal surface finer than in anterior impressions of terga III-V, but conspicuously coarse, somewhat granulose, and dense in comparison to other Amarochara species, interstices on terga III-VI distinctly narrower than punctures. Tergum X anteriorly distinctly elongated, narrowly truncate; with very dense, stout, long and dark yellowish setae; posterior median area without setae (Fig. 114). cf: unknown. $: posterior margin of tergum VIII almost truncate and with rather dense row of mar ginal setae (broken off in the holotype); posterior margin of sternum VIII with dense row of short unmodified marginal setae; spermatheca of characteristic derived mor phology, relatively small, with apically constricted capsule, the latter connected to duct by a short and broad tube; duct rather short, subapically thin and distinctly bent, and apically noticeably dilated (Fig. 112).

Derivatio nominis
The name (Lat., adj.) refers to the conspicuously dense puncturation of the abdomen.

Comparative notes and comments
A. densepunctata is distinguished from all its congeners especially by the dense puncturation of the abdomen and by the morphology of the spermatheca. It additionally differs from other Eastern Palaearctic species in the lighter coloration and the large pronotum, which is much wider than the head and only slightly narrower than the elytra.

Distribution and bionomics
The holotype was collected near Tsumagoi (Gunma, Honshu), some 140 km NW Tokyo at an altitude of approximately 1000m.

Redescription
Relatively large species, 4.6 mm. Colour of body distinctive: head, pronotum, and abdo men blackish, except for the slightly lighter abdominal apex; elytra bicoloured, with the anterior and lateral margins blackish and with the posterior and interior two thirds of each elytron brightly rufous, thus forming a clear and large reddish spot; antennae and legs light brown to yellowish brown. Whole body without appreciable microsculpture. Pubescence of forebody relatively long, decumbent to suberect. Head of subcircular shape, approximately as wide as long; eyes very large, moderately convex, weakly projecting from lateral outline of head, almost as long as postgenae in dorsal view; postgenae smoothly curved towards posterior margin. Puncturation moderately sparse and moderately fine, but distinct, with interspersed micropunctures. Genal carina present and distinct. Antennae very long and massive, similar to those in M . siculifera A SSIN G , antennomere I with dorso-apical furrow reduced, very short and inconspicuous; antennomere III even slightly longer than II; IV and V moderately transverse; VI -X weakly transverse. Third joint of maxillary palpus very long and slender, similar to that in AE siculifera. Pronotum large in relation to head, 1.15 times as wide as long and 1.37 times as wide as head; maximal with a short distance anterior to middle; posterior angles weakly defined; puncturation similar to that of head, but much denser, interstices mostly narrower than punctures. Elytra relatively long, at suture 0.85 times the length of pronotum and approximately 1.20 times as wide as pronotum; posterior margin near posterior angles weakly sinuate; puncturation, especially in anterior half, coarse and very dense, interstices reduced to narrow ridges. Hind wings fully developed. Mesosternum without carina, mesosternal process long and apically acute. Legs very long and slender, metatarsus almost as long as metatibia and first metatarsomere as long as the combined length of the three following tarsomeres.
Abdominal terga III-V with deep anterior impressions, tergum VI without anterior im pression; sterna III-V anteriorly weakly constricted; puncturation in impressions of ter ga III-V and near anterior margin of tergum VI very coarse and dense, on remainder of dorsal abdominal surface moderately dense and relatively coarse, denser and much coar ser than in M. siculifera and AE forticornis. Tergum and sternum VIII long and slender. Tergum X with long anterior extension.
d : posterior margin of tergum VIII moderately convex, that of sternum VIII strongly convex, with a dense row of relatively long marginal and a sparse row of very long submarginal thin setae ; median lobe of aedeagus long and slender, at base of ventral process with distinct median process best seen in lateral shew; crista apicalis completely, crista proximalis partly reduced; internal tube with relatively long basal part; internal sac without distinctly sclerotized structures ; apical lobe of paramere slightly modified, the two apical setae very short (Fig. 124). $: unknown.

Comparative notes and comments
A. splendens is distinguished from other species of the genus by the distinctive coloration alone. Based on the morphology of the median lobe of the aedeagus (especially the presence of a median process at the base of the ventral process), the morphology of the antennae and mouthparts, the extremely long and slender legs, and other morphological characters, A. splendens is closely related to A. siculifera sp.n. and A. inermis sp.n, from the eastern Mediterranean. It is additionally separated from these species by much larger eyes, the more distinct puncturation of head, pronotum, elytra, and of the posterior areas of the abdominal terga III -VII, by the more slender median lobe of the aedeagus, and by the different shape and chaetotaxy of the apical lobe of the paramere.

Distribution and bionomics
The species is known only from the type locality in the Haut Atlas, Marocco, where it was collected at an altitude of 1400 m in April.

A m a r o c h a r a t i n g i t a n a J a r r i g e , 1952
A m arochara {Mniobates) tingitana JARRIGE, 1952: 138.

Comments
The holotype of this species was apparently lost in the process of posting. The labels read as follows: 94 Tanger / Maroc ex Musae E. Vaucher 1908 / Amarochara tingitana Jarr. / TYPE / Muséum Paris Ex Collection J. Jarrige 1976 (MNHNP). Unfortunately, the original description is based on a single specimen, so that an interpretation of this species and a neotype designation will have to wait until additional material becomes available.

5.
Key to the A m arochara species o f the Holarctic region The following key does not account for A. tingitana JA RRIG E (see chapter 4.7).

8.
Coloration of body uniformly blackish. Head and pronotum with distinct micro sculpture. Puncturation in impressions of terga III-V and near anterior margin of tergum VI coarse and dense. Spermatheca as in Fig. 68 At least head and abdomen distinctly darker. Antennae longer than 0.75 mm. Eyes larger and more bulging, distincdy projecting from lateral outline of head; postgenae less than 1.5 times as long as eyes in dorsal view. 18. Larger species, 3.0 -3.8 mm. Coloration on the whole usually darker, pronotum at most only slightly lighter than the dark brown to blackish head; antennae not distinctly bicoloured. Antennae longer and with less transverse antennomeres; antennomeres VII-IX approximately twice as wide as long (Fig. 37). Pronotum relatively large, 1.25-1.30 times as wide as head. Elytra relatively short, at suture approximately 0.75 times as long as pronotum. Legs distinctly longer, length of metatarsus even in small specimens at least 0.37 mm; first metatarsomere very elongate, as long as the combined length of the three following tarsomeres or nearly so (Fig. 35). Abdomen with more clearly defined and less dense punctu ration, and with pronounced shine, d : aedeagus of completely different shape, with much longer and more slender ventral process (Figs. 29-30). 9: spermatheca smaller (Fig. 32) Pronotum light brown to brown, distinctly lighter than the blackish head; antennae distincdy bicoloured, dark brown to blackish, with yellowish to ferrugineous basal antennomeres. Antennae shorter and with more transverse antennomeres; antennomeres VII-IX almost 3 times as wide as long. Pronotum relatively smaller. Elytra longer, at suture >0.80 times as long as pronotum. Legs much shorter, length of metatarsus approximately 0.30 mm; first metatarsomere of similar length as in A. umbrosa, shorter than the three following tarsomeres. Abdomen with less defined and much denser puncturation, and the refore with reduced shine, d : median lobe of aedeagus much shorter and broader ventral process . ¥: spermatheca larger (Fig. 42) 20. Aedeagus and paramera strongly modified. Ventral process of median lobe short and small in relation to basal part, in lateral view with short process or obtuse projection near base of ventral process; internal sac with several strongly sclerotized structures. Apical lobe of paramere distinctly flattened (Figs. 84,85,95). .. 21 Aedeagus and paramere not strongly modified. Ventral process of median lobe longer in relation to basal part, in lateral view without short process or obtuse projection near base of ventral process; internal sac with weakly sclerotized struc tures. Apical lobe of paramere not distinctly flattened 22 21.
Ventral process of median lobe longer, in lateral view near base of ventral lobe with short and acute process; sclerotized structures of aedeagus of distinctive shape 86); apical lobe of paramere not transverse, apically rounded ; spermatheca as in Fig. 87

Comments
The holotype of A. japónica is not a member of the oxypodine genus Amarochara, but belongs to the Athetini and is here attributed to the genus Alevonota TH O M SO N based on its general appearance (slender body, slender pronotum), the rather stout antennae, the mouthparts (deeply bifid ligula), and the morphology of the primary sexual characters, especially of the spermatheca. It differs from Western Palaearctic congeners particularly in the shape of antennomere I, which is rather large, somewhat flattened, and which has a pronounced wide dorsal furrow.

Comments
Prior to the present study, the mouthparts and the genitalia of the holotype had been dissected and glued to the mounting label. Various solvents were tried, but a glue had been used that proved to be resistant to all these agents, so that it was impossible to assess many important characters properly and to examine them under a compound microscope (mouthparts, primary and secondary sexual characters, pro-and mesotarsi, ventral aspect of body). The holotype is definitely no Amarochara; its antennae, abdo men, metatarsi, maxillary palpi, and aedeagus are of completely different morphology. Judging from the aedeagus and other characters, it apparently belongs to an athetine genus, characterized by relatively massive antennae (with weakly transverse antennomeres IV-X, short and stout antennomeres II and III, a short antennomere XI, and a distinct dorsal furrow on antennomere I), maxillary palpi with the third joints distinctly dilated, relatively short and stout mandibles, a transverse pronotum, a short first metatarsomere (not longer than second), and anteriorly weakly impressed abdominal terga III-V. The generic affiliations of this species may remain uncertain until new material becomes available for examination.

Comments
Nasirema humilis, the type species of Nasirema C ASEY, 1893, was previously attributed to Amarochara (e. g. BERNHAUER & SCHEERPELTZ 1926). However, an examination of the holotype revealed that the species belongs to Calodera M AN N ERH EIM , 1830 and not to Amarochara. Consequently, Nasirema, which has previously been regarded as a syno-nym of A marochara, is here placed in the synonymy of the senior name Calodera. C. humilis CASEY is a secondary junior homonym of C. humilis E RICH SO N , 1837 (which again is a junior synonym of Caloderaprotensa M AN N ERH EIM , 1830), so that the replace ment name Calodera caseyi nom. n. is proposed for C. humilis (CA SE Y ). C. caseyi belongs to the group of species without posteriorly distinctly constricted head. From C. lapponica SAHLBERG and C. protensa M AN N ERH EIM , the species is distinguished by its distinctly smaller size (in this respect similar to C. aethiops (G-RAVENHORST)), the lighter coloration (light brown, with the elytra and the appendages testaceous), the shorter antennae with more transverse antennomeres V-X (antennomere more than twice as wide as long), and by the denser and coarser puncturation of the pronotum.  In order to fix a single name-bearing type and to preserve the present interpretation of the species, the syntype from the Bernhauer collection is designated as lectotype. Illu strations of the genitalia of A. subtumida are provided by GUSAROV (1991), who studied the types. Two years after the description of 0. reitteri, BERNHAUER (1902) attributed the species to the new subgenus Sorecocephala, with 0. reitteri as the sole representative and type species by monotypy. No further species of Sorecocephala have become known. Since O. reitteri is a synonym of a species of Aleochara, the following synonymy is here esta blished: Aleochara GrAVENHORST, 1802 = Sorecocephala BERNHAUER, 1902, syn. n.