Anaulacaspis formosa ( ROSENHAUER, 1856 )

Assing, Volker, 2016, Revision of the Anaulacaspis species of the Palaearctic region (Coleoptera: Staphylinidae: Aleocharinae), Beiträge Zur Entomologie = Contributions to Entomology 66 (2), pp. 201-255 : 248-251

Anaulacaspis formosa ( ROSENHAUER, 1856)

( Figs 196–211 , Map 3 )

Falagria formosa ROSENHAUER, 1856: 88 f.

Falagria cirrosa FAUVEL, 1902: 184 f.; syn. n.

Melagria formosa algirica FAGEL, 1969: 30 ; syn. n.

Melagria formosa elkantarensis FAGEL, 1969: 31 f.; syn. n. Melagria formosa macra FAGEL, 1969: 30 f.; syn. n.

Melagria formosa biskrensis FAGEL, 1969: 32 ff.; syn. n. Melagria cirrosa andalusiaca FAGEL, 1969: 36 ; syn. n.

Type material examined: Falagria formosa : Lectotype ♀: “Malaga / Andalusie / Algeciras / Puerto de Gajnrés [?] ( Espagne) / formosa Rosh. type / R.I.Sc.N.B. 17.479, Falagria naevula Er., Coll. et det. A. Fauvel / Ex-Typis / G. Fagel elig., 1968, Falagria formosa Rosh. , Lectotype / Anaulacaspis formosa (Rosenhauer) , det. V. Assing 2016” ( IRSNB).

Falagria cirrosa : Lectotype ♂ [dissected by R. Pace]: “Teboursouk / cirrosa Fvl. / R.I.Sc.N.B. 17.479, Falagria, Coll. et det. A. Fauvel / Ex-Typis / G. Fagel elig., 1968, Falagria cirrosa Fauvel , Lectotype / vidit R. Pace 1985” ( IRSNB) . Paralectotypes: 1 ♀: “ Teboursouk 7 / R.I.Sc. N.B. 17.479, Falagria cirrosa Fauv. Coll. et det. A. Fauvel / Ex-Typis / Falagria cirrosa Fvl. , det. R. Pace 1985” ( IRSNB) ; 1 ♀: “ Palestro , Algerie / R.I.Sc. N.B. 17.479, Falagria cirrosa Fauv. Coll. et det. A. Fauvel / Ex-Typis / Falagria cirrosa Fvl. , det. R. Pace 1985” ( IRSNB) ; 1 ♀: “ Chabounia , Prov. d’Alger / R.I.Sc. N.B. 17.479, Falagria cirrosa Fauv. Coll. et det. A. Fauvel / Ex-Typis / Falagria cirrosa Fvl. , det. R. Pace 1985” ( IRSNB) ; 1 ♀ [dissected by R. Pace ]: “Aïn Draham, 7 Tunisie / R.I.Sc. N.B. 17.479, Falagria cirrosa Fauv. Coll. et det. A. Fauvel / Ex-Typis / Falagria cirrosa Fvl. , det. R. Pace 1985” ( IRSNB) ; 1 ♀ [dissected by R. Pace]: “ Bou Berak près Dellys / R.I. Sc. N.B. 17.479, Falagria cirrosa Fauv. Coll. et det. A. Fauvel / Ex-Typis / Falagria cirrosa Fvl. , det. R. Pace 1985” ( IRSNB) ; 1 ♂: “ El Fedja , 12, Tunisie / Souk el Arba, inond. Madjerda, 1 / R.I. Sc. N.B. 17.479, Falagria cirrosa Fauv. Coll. et det. A. Fauvel / Ex-Typis / Falagria cirrosa Fvl. , det. R. Pace 1985” ( IRSNB) ; 1 ♂ [dissected by R. Pace ], 2 ♀♀: “Les Lacs, Prov. de Const., Algérie / R.I.Sc. N.B. 17.479, Falagria cirrosa Fauv. Coll. et det. A. Fauvel / Ex-Typis / Falagria cirrosa Fvl. , det. R. Pace 1985” ( IRSNB) ; 1 ♀: “ Tunis 9 / R.I.Sc. N.B. 17.479, Falagria cirrosa Fauv. Coll. et det. A. Fauvel / Ex-Typis / Falagria cirrosa Fvl. , det. R. Pace 1985” ( IRSNB) ; 2 ♀♀: “ Fernana 8, Tunisie / R.I.Sc. N.B. 17.479, Falagria cirrosa Fauv. Coll. et det. A. Fauvel / Ex-Typis / Falagria cirrosa Fvl. , det. R. Pace 1985” ( IRSNB) .

Melagria formosa algirica : Holotype ♂: “ Philippeville , Algérie, A. Thery / G. Fagel det., formosa algirica nov. / Type / Anaulacaspis formosa (Rosenhauer) , det. V. Assing 2016” ( IRSNB) . Paratypes: 1 ♂: same data as holotype ( IRSNB) ; 1 ♀: “Fil-Fila, Algérie, A. Thery ” ( IRSNB) ; 1 ♀: “ Bône / R.I.Sc. N.B. 17.479, Falagria cirrosa Fauv. Coll. et det. A. Fauvel / Ex-Typis / Anaulacaspis cirrosa ( Fauvel) , det. V. Assing ” ( IRSNB) .

Melagria formosa elkantarensis : Holotype ♂ [slightly teneral]: “ Algérie: Djebel Metlilli , gorges d’El Kantara , V-1954, G. Fagel / formosa elkantarensis nov. / Type / Anaulacaspis formosa (Rosenhauer) , det. V. Assing 2016” ( IRSNB) . Paratypes: 5 ♂♂, 1 ♀ [partly teneral]: same data as holotype ( IRSNB) .

Melagria formosa macra : Holotype ♂: “ Algérie: Algerois , Kaddous, 3-v-1954, G. Fagel / formosa macra nov. / Type / Anaulacaspis formosa (Rosenhauer) , det. V. Assing 2016” ( IRSNB) . Paratypes: 1 ♂, 1 ♀: same data as holotype ( IRSNB); 1 ♂: “ Alger ” ( IRSNB) .

Melagria biskrensis : Holotype ♂: “ Algérie: Ziban , oued Biskra , 17/ 25-v-1954, G. Fagel / biskrensis n. sp. / Type / Anaulacaspis formosa (Rosenhauer) , det. V. Assing 2016” ( IRSNB) . Paratypes: 2 ♂♂ [teneral]: same data as holotype ( IRSNB) ; 1 ♂: “ Algérie: Djebel Metlilli , gorges d’El Kantara, V-1954, G. Fagel ” ( IRSNB) ; 1 ♂: “ Tougourt , 10–20-v-36, Algérien, leg. Vogel ” ( IRSNB) ; 1 ♂: “ Algérie: Ziban , oasis de Filiach, 25-v-1954, G. Fagel ” ( IRSNB) ; 1 ♂: “ Melaha , Trip., 13.4.1936, R. e C. Koch ” ( IRSNB) ; 1 ♂: “ Kairouan 10 / Laghruat [?] / R.I.Sc. N.B. 17.479, Falagria naevula Er., Coll. et det. A. Fauvel ” ( IRSNB) ; 2 ♂♂: “T. Kairouan , Dr. Normand / R.I.Sc. N.B. 17.479, Falagria naevula Er., Coll. et det. A. Fauvel ” ( IRSNB) ; 1 ♂: “ Kairouan 1 / R.I.Sc. N.B. 17.479, Falagria naevula Er., Coll. et det. A. Fauvel ” ( IRSNB) .

Melagria cirrosa andalusiaca : Holotype ♂: “ Andalousie: S. Pedro Alcantara ( Marbella ), V.1963, G. Fagel / G. Fagel det., cirrosa andalusiaca nov. / Type / Anaulacaspis cirrosa (Fauvel) , det. V. Assing 2016” ( IRSNB) . Paratypes: 1 ♂, 1 ♀: same data as holotype; 1 ♀: “ Andalousie : Ronda, rio Grande, 8/ 20-V-1956, G. Fagel / G. Fagel det., cirrosa andalusiaca nov. / Paratype ” ( IRSNB) ; 1 ♂, 1 ♀: “ Andalousie : Lanjaron (prov. Granada), 26.IV.–18.V.1961, G. Fagel, 85 / G. Fagel det., cirrosa andalusiaca nov. / Paratype ” ( IRSNB) ; 1 ♀: “ Andalousie : Ronda, el Tajo, 16-V-1956, G. Fagel / G. Fagel det., cirrosa andalusiaca nov. / Paratype ” ( IRSNB) .

Comment: The original description of Falagria formosa is based on an unspecified number of syntypes “Bei P. S. Maria und Malaga ... häufig gefunden” (ROSENHAUER 1856). FAGEL (1969) designated a female from the Fauvel collection, which he regarded as a syntype, as the lectotype. According to HORN et al. (1990), the remaining type specimens should be in the MNHNP.

The original description of Falagria cirrosa was described from an unspecified number of syntypes from “ Algérie et Tunisie ” ( FAUVEL 1902). A lectotype was designated by FAGEL (1969). FAUVEL (1902) already suspected that A. cirrosa represented “une race bien distincte” of A. formosa (which he erroneously referred to as A. naevula ).

The original description of Melagria formosa algirica is based on a male holotype from “Algérie: Philippeville”, a paratype from the same locality, and four paratypes from four additional localities in Tunisia and Algeria ( FAGEL 1969).

Melagria formosa macra was described from a male holotype and two paratypes from “Algérie: Algérois, Kaddous” and five paratypes from three additional localities in Algeria ( FAGEL 1969).

The original description of Melagria formosa elkantarensis is based on a male holotype and “Nombreux exemplaires” from Djebel Metlilli, gorges d’El Kantara”, as well as one paratype from “Aurès, M’Chounech” ( FAGEL 1969).

Melagria biskrensis was described based on a male holotype from “Algérie: Ziban, oued Biskra ” and additional paratypes from various localities in Algeria, Tunisia, and Libya ( FAGEL 1969).

The original description of Melagria cirrosa andalusiaca is based on a male holotype and seven paratypes from “Andalousie: Marbella, San Pedro Alcantara, rives du rio Guadalania” and eight paratypes from four additional localities in Andalucía ( FAGEL 1969).

An examination of the type material of the above names, as well as of the additional material listed below revealed an enormous variation of external characters such as size, coloration, and the length and width of the elytra. However, the male sexual characters are identical. Moreover, different morphs do not correspond to any plausible distribution patterns and were often found in the same region. Part of the type specimens of different species or subspecies were even collected in the same localities. In consequence, the characters emphasized by FAUVEL (1902) and FAGEL (1969) as distinguishing A. cirrosa , A. biskrensis , and the subspecies of A. formosa and A. cirrosa from the nominal subspecies of A. formosa are interpreted as intraspecific variation of a highly variable species. There is no evidence that this species is represented by geographically distinct subspecies, even though the smaller and more slender morph with smaller elytra is the predominant form in southern Spain and rather rare in North Africa. This multiple synonymy may be explained not only with the pronounced intraspecific variation of A. formosa , but also with the observations that some of the type specimens are teneral and that all the specimens originally labelled as A. formosa in the IRSNB are females, whereas the type series of A. biskrensis is exclusively composed of males.

Although part of the type material of Anaulacaspis biskrensis was collected in Libya, neither A. biskrensis nor any of the other names now in the synonymy of A. formosa is listed for Libya in the Palaearctic Catalogue ( SCHÜLKE & SMETANA 2015).

Additional material examined: Spain: Murcia: 2 ♂♂, 1 ♀, Murcia env. [“Sp 202”], leg. Franz ( NHMW, cAss) .

Andalucía: 3 ♂♂, 3 ♀♀, Cádiz, Sierra de Grazalema, El Bosque , rio Malaceite , 26.VI.2003, leg. Santamaria (cAss); 1 ♀, Cádiz, 20 km N Tarifa, 36°09'N, 5°38'W , 100 m, loamy shore of reservoir, flood debris, sifted, 29.XII.2009, leg. Assing (cAss) ; 1 ♂, 1 ♀, Malaga, Jubrique, 500 m, stream bank, debris, 26.III.1994, leg. Wunderle (cWun) ; 9 ♂♂, 11 ♀♀, Ronda, 500 m, flood debris, 6.III.1990, leg. Matern (cWun, cAss); 1 ♂, 2 ♀♀, Sierra de Cazorla [“ Sp 687”], leg. Franz ( NMW) ; 2 ♀♀, Cádiz, 9 km SE Alcala de los Gazules, bank of Rio Alberite , 7.V.1991, leg. Hieke ( MNB) ; 1 ♀, Estepona env. [“Sp 645”], leg. Franz ( NHMW) ; 1 ♂, 3 ♀♀, Estepona, trail to Ronda , 1000 m, X.1986, leg. Benick ( MNB) ; 4 ♀♀, Estepona , 20.X.1968, leg. Benick ( MHNG) ; 5 ♀♀, Estepona , X.1968, leg. Benick ( MHNG) ; 1 ♂, 1 ♀, Estepona , X.1968, leg. Benick ( MNB) ; 1 ♀, Malaga , leg. Weise ( NHMW) ; 4 ♂♂, 4 ♀♀, Malaga, 20.XI.1980, leg. Simon ( NHMW, cAss) ; 1 ♀, Algeciras ( MHNG) ; 1 ♀, bank of Galadjaro river , 28.–31.V.1968, leg. Benick ( MHNG) ; 1 ♂, locality not specified (MNB).

Morocco: 4 ♂♂, 4 ♀♀, Haut Atlas, NE Tizi-nTest , 30°54'N, 8°19'W, 1710 m, stream bank, sifted, 29.XII.2002, leg. Assing (cAss) ; 2 ♂♂, 4 ♀♀, same data, but leg. Wunderle (cWun) ; 1 ♂, 1 ♀, Haut Atlas , S Tizin-Test, Tachguette, 30°51'N, 8°23'W, 1600 m, 30.IV.2009, leg. Behne (cAss) ; 2 ♂♂, 2 ♀♀, Haut Atlas , Tinmal, XI.1985, leg. Rougemont (cAss) ; 1 ♀, Ar Rif , road Chefchaouen – Ksal el Kebr, 35°05'N, 5°25'W, 680 m, 3.–7. VI.2007, leg. Hlaváč (cAss) ; 1 ♂, 1 ♀, Ar Rif , Oued Laou env., 35°20'N, 5°07'W, 200 m, 7.–9.VI.2007, leg. Hlaváč (cAss) ; 5 ♂♂, Ar Rif , Chefchaouen env., 35°12'N, 5°19'W, 270 m, 3.VI.2007, leg. Hlaváč (cAss) ; 1 ♂, 2 ♀♀, Forêt Mamora , stream on road to Tiflet, 34°04'N, 6°06'W, 160 m, 26.III.2008, leg. Andújar et al. (cAss); 1 ♀, 30 km NW Berkane, M. de Kebdana, 350 m, 35°03'N, 2°36'W, 31.XII.2001, leg. Bayer ( MNB); 1 ♀, Larache env., Jemis del Sahel , 3.IV.1959, leg. Besuchet ( IRSNB); 1 ♀, Esmir , leg. Walker ( IRSNB); 1 ♂, 1 ♀, Bni Tadjite, Oued Ait Aissa, 32°18'N, 3°28'W, 17.I.2015, leg. Daoudi (cTro) ; 1 ♀, Mansouria , IV.1936, leg. Grosclaude (cTro) ; 2 ♂♂, Ouirgane , UV light, 19.V.2015, leg. Miessen (cTro) ; 5 exs., Marrakesh, leg. Quedenfeldft ( MNB); 1 ♀, Morbeya river , leg. Quedenfeldt ( MNB); 3 ♀♀, Larache env. , Jenis del Sahel , 3.IV.1959, leg. Besuchet ( MHNG); 2 ♂♂, 2 ♀♀, “ Fl. Issil ”, leg. Quedenfeldt ( NHMW, cAss); 1 ♂, locality not specified, leg. Simon ( NHMW) .

Algeria: 1 ♂, Biskra, Hammam Salahine, 24.–25.V.1971, leg. Hoffer & Horák (cAss); 1 ♂, Ain Zatout, Djebel Aures , 26.V.1971 (cAss); 1 ♀, “Lalla Marnia” ( IRSNB) ; 1 ♀, Philippeville, Bou Saada ( IRSNB ); 1 ♂, 2 ♀♀, Bejaia [“ Bougie ”; 36°45N, 05°05E], leg. Théry ( NHMW) ; 1 ♀, Hammam Righa [36°22'N, 2°23'E] ( MNB) ; 1 ♂, locality not specified, leg. Fauvel ( NHMW) ; 1 ♀, Mustapha , leg. Peyerimhoff ( NHMW) ; 1 ♀, Annaba ( MHNG) ; 3 ♂♂, 1 ♀, Blidah ( MHNG).

Tunisia: 1 ♀, Jendouba, Ben M’Tir, 8 km S Aïn Draham, 36°43'N, 8°42'E, 610 m, 2.III.2012, leg. Hetzel (cFel) ; 5 ♂♂, 3 ♀♀, Aïn Draham, IX.1945, leg. Demoflys ( MHNG, cTro); 1 ♀, Sousse, 18 km SW Hammamet, Bouficha , 1.XII.1994, leg. Wrase (cAss); 1 ♀, Teboursouk, VII. ( IRSNB) ; 1 ♂, Jerid , 4 km NE Tozeur, 30–40 m, light trap, 21.X.2009, leg. Lehmann (cAss); 1 ♀, Nabeul, coast, 0–30 m, 4.–16.IV.1999, leg. Winkelmann (cAss); 2 ♂♂, 3 ♀♀, Le Kef ( MHNG, NHMW); 1 ♀, Le Kef, leg. Normand (cTro) ; 1 ♂, 4 ♀♀, Le Kef, X.1941, leg. Demoflys (cTro); 3 ♂♂, 1 ♀, Le Kef, IX.1935, leg. Normand (cTro); 6 ♂♂, 4 ♀♀, Rades , XII.1936, leg. Grosclaude (cTro, cAss); 3 ♂♂, 2 ♀♀, same data, but X.1943 (cTro); 1 ♂, 2 ♀♀, Rades, X.1941, leg. Demoflys (cTro); 1 ♀, Gabès, I.1946, leg. Demoflys (cTro); 1 ♀, Kairouan, III.1925, leg. Rambousek ( MHNG) .

Presumably mislabelled: 1 ♀, “Pyr.-Or.” ( NHMW); 1 ♀, “ Herault, Palavas ” ( MHNG).

Redescription: Habitus as in Fig. 211 . Body length 2.4–3.3 mm; length of forebody 1.2–1.5 mm. Coloration: head reddish-brown to blackish-brown; pronotum reddish to reddish-brown; elytra reddish-yellow with a more or less distinct and more or less extensive mediolateral infuscate spot; abdomen bicoloured: segments III and IV reddish, segment V usually at least partly darkbrown to blackish; segments VI–VIII blackish-brown to black; legs yellowish; antennae brown to dark-brown with the basal 3–4 antennomeres yellowish to reddish.

Head relatively weakly transverse, approximately 1.1 times as broad as long, with weakly pronounced sexual dimorphism. Eyes slightly longer than postocular region in dorsal view. Antennae slender, 0.9–1.1 mm long; antennomeres IV–V usually at least weakly oblong, VI approximately as long as broad, VII–X increasingly transverse, X approximately 1.5 times as broad as long.

Pronotum approximately as broad as long or weakly transverse, and approximately as broad as head; sexual dimorphism usually weakly pronounced; punctation rather dense.

Elytra of hightly variable width and length, 0.8–1.0 times as long as pronotum; punctation very fine and moderately dense. Hind wings present, possibly poly- or dimorphic.

Abdomen as broad as, or slightly narrower than, elytra; punctation fine, but distinct, rather dense, and more or less regularly distributed; posterior margin of tergite VIII ( Figs 196–197 ) weakly convex, with a row of distinctly modified short and stout setae.

♂: punctation of head and pronotum on average more distinct than in female; pronotum sometimes more or less depressed or shallowly impressed and/or with short fine median sulcus in postero-median portion; sternite VIII ( Fig. 198 ) with convex posterior margin; median lobe of aedeagus ( Figs 199–208 ) usually 0.36–0.40 mm long (exceptionally larger; see notes on intraspecific variation below), with weakly pronounced crista apicalis; ventral process apically weakly incised in ventral view.

♀: posterior margin of sternite VIII ( Fig. 209 ) concave in the middle; spermatheca ( Fig. 210 ) minute and commashaped, not distinctive.

Intraspecific variation: Anaulacaspis formosa is subject to enormous intraspecific variation both within and between populations, not only of size, coloration, the length and width of the elytra, as mentioned above, but also of the male modifications of the head and pronotum, and even the size of the aedeagus. In two males from Tunisia and Morocco, the median lobe is distinctly larger (length 0.47 mm; see Figs 199–202 ) and the pronotum of the male from Tunisia has a postero-median impression. However , the male from Morocco was collected together with other males with aedeagi of normal size and no additional evidence was found suggesting that these specimens represent a distinct species. In any case, the extent of intraspecific variations indicates a patchy distribution and low gene flow between populations.

Comparative notes: As can be inferred from the similarly derived chaetotaxy of the abdominal tergite VIII (posterior margin with strongly modified marginal setae) and from the similar morphology of the aedeagus, A. formosa is evidently the adelphotaxon of the allopatric A. elegans . For characters distinguishing both species see the comparative notes in the following section. The only geographically close and possibly partly sympatric congener is A. desertorum , from which A. formosa differs by a less transverse head, a more slender and more convex (cross-section) pronotum with a much less pronounced sexual dimorphism, the coloration particularly of the elytra (medio-lateral infuscate spot less defined and less extensive), the modifications of tergite VIII, and by the completely different morphology of the aedeagus.

Distribution and natural history: Based on the examined material, A. formosa is widespread and rather common in South Spain and North Africa ( Morocco, Algeria, Tunisia, Libya) ( Map 3 ). Since this species has been misinterpreted or confounded with other species in the past, literature records from other regions are doubtful. The same applies to the two specimens categorized above as presumably mislabelled.

The specimens for which ecological data are available were hand-collected or sifted from debris on the banks of rivers and streams, or they were caught with light traps. Teneral specimens were found in May. The altitudes range from 100 to 1710 m.

Figures

Map 3: Distributions of Anaulacaspis formosa (black circles), A. elegans (white circles), A. nigrina (black diamonds), A. eminens (white diamonds), and A. nigerrima (stars) in the West Palaearctic and Middle Asia, based on revised records.

Figs 187–210: Anaulacaspsis gilva (187–190), A. pectinata (191–195), and A. formosa (196–210; 199–200: Tunisia; 201–203: Morocco; 204–205: holotype of A. macra; 206: Murcia; 207–208: Andalucía): median lobe of aedeagus in lateral and in ventral view (187–188, 194–195, 199–208); female tergite VIII (189); female sternite VIII (190, 209); male tergite VIII (191, 196); posterior margin of male tergite VIII (192, 197); male sternite VIII (193, 198); spermatheca (210). Scale bars: 187–191, 193–196, 198–210: 0.1 mm; 192, 197: 0.05 mm.

Figs 211–232: Anaulacaspsis formosa (211), A. elegans (212–220), A. beesoni (221–228; 224–225: lectotype), and A. inexpectata (229–232; 231–232: holotype): habitus (211; photo by Marc Tronquet); male tergite VIII (212, 221); male sternite VIII (213, 222); median lobe of aedeagus in lateral and in ventral view (214–217, 224–227, 229–232); female tergite VIII (218); posterior margin of female tergite VIII (219); female sternite VIII (220, 228); postero-lateral portion of sternite VIII (223). Scale bars: 211: 1.0 mm; 212–218, 220–222, 224–232: 0.1 mm; 219, 223: 0.05 mm.