Cholovocera attae (Kraatz, 1858)

Figs 4B, 9C, 10B, 12B, 13A, 14B–C, 19

Choluocera Attae Kraatz, 1858: 140.

Coluocera attae – Gemminger & Harold 1868: 905.

Colovocera Attae – Lucas 1874: 239.

Coluocera formicaria – Reitter 1875: 301 (in part).

Colovocera formicaria – Belon 1879: 192 (in part).

Colnocera [sic] attae – Walker 1888: 181.

Cholovocera attae – Rücker 1980: 143, fig. 22 — Audisio et al. 1995: 9.

Differential diagnosis

Cholovocera attae and Ch. occulta sp. nov. are the smallest species in the genus. Size and short wide legs would separate Ch. attae from its sympatric Ch. balcanica. However, the most useful character to distinguish Ch. attae from Ch. balcanica and from all the other species is the morphology of the aedeagus (Fig. 19).

Type material

Cholovocera attae: lectotype male and paralectotype female in the Kraatz Collection, held in SDEI.

Lectotype of Colovocera attae

GREECE • 1 ♂; “Graecia”; Kraatz Collection; SDEI 11869. Designated by Rucker (2011a: 13).

Paralectotype of Colovocera attae

GREECE • 1 ♀; “Graecia”; Kraatz Collection; SDEI 11870.

Notes

As it can be seen in Rücker 2011a (fig. 12), there is a label reading “ Holotypus ” attached to the specimen that Rücker designated as the lectotype. However, this specimen cannot be regarded as the holotype because it was not designated in the original description, which included more than one specimen, i.e., syntypes. Examining the printed Holotypus label, we conclude that it was added at a later date than the description by Kraatz (1858).

The same comment given below under the lectotype, applies to the paralectotype.

Additional material, non-types

GREECE – Peloponnese • 2 ♂♂, 1 ♀; Menalon mts., 5 km E. Vitina; 37º40′46.5″ N, 22º14′52.7″ E; 1380 m a.s.l.; 10 May 2013; Schuh leg.; “Almweide” [Mountain meadow], [one specimen associated with a Messor worker ant]; NHMW – Crete • 1 ♂; West Crete, Vai-Itanos; 13 Apr. 1984; H. Fülscher leg.; NHMW • 4 ♂♂, 1 ♀; West Crete, Levka Ori, r[oa]d. Hania to Omalos, 2.5 km NNE of Omalos; 1150 m a.s.l.; 2 Jun. 2010; Schuh leg.; NHMW • 2 ♂ ♂; Crete; NMPC.

TURKEY – Western Anatolia, Aegean • 1 ♂, 2 ♀♀; Muğla, Göcek; 2 May 1975; Besuchet and Löbl leg.; MHNG.

Reitteria escherichi Wasmann, 1896

TURKEY – Western Anatolia, Aegean • 1 ♂, 1 ♀; Muğla, Göcek; 2 May 1975; Besuchet and Löbl leg.; MHNG.

Type locality

“ Griechenland ”, Greece.

Description

Male as in Fig. 10B. Body length: 1.36 mm average, range 1.20–1.40 mm (N = 10, males and females). Shape of body oval, with the lateral margins of the pronotum slightly widened anteriorly. Elytral apex rounded. Terminal antennomere subtriangular, with round angles (Figs 12B, 13A). Metatibiae relatively short, as in Fig. 14B–C. Prosternal process slightly keeled anteriorly, with a marked median constriction and subquadrangular distally (Fig. 4B). Male last visible ventrite with a slight emargination and bordered by a brush of short setae.

Median lobe of aedeagus curved towards the right in ventral view, tapering to a pointed apex (Fig. 19A). Aedeagus in lateral view as in Fig. 19C. Distal portion of paramere conical, with five or six long apical setae (Fig. 19B–C). The spermatheca was not dissected because of the small number of females available for study.

Geographic distribution

The known distribution of Cholovocera attae is in Greece (Peloponnese, Crete) and western Turkey (Fig. 9C). However, the few available records of this species may not represent its total distributional range.

Host ants

Smith (1874) cited studies by Moggridge (1873), who collected Ch. attae in nests of Messor ibericus Santschi, 1925 (as Atta structor) in Menton (southern France). Messor ibericus is the only species of the Messor structor complex present in that region (Steiner et al. 2018). However, according to our present knowledge, the beetles could not have been Ch. attae, but Ch. punctata, Ch. gallica or Ch. occulta sp. nov. Subsequent authors perpetuated the incorrect identifications of both beetles and ants (e.g., Lucas 1874; Rücker 1980; Angelini & Rücker 1999). However, Rücker (2018: 576) correctly associated Ch. attae with Messor.

At present, it is not possible to know the species of ant associated with Cholovocera attae in Greece. We have examined one beetle preserved together with an ant worker of the genus Messor but, considering the large number of species of this genus living in Greece and the lack of good descriptions, we cannot identify it to species (X. Espadaler pers. comm. 13 Mar. 2021).

Taxonomic history and remarks

Kraatz’s (1858: 140) description of Cholovocera attae is brief, but he accurately distinguished it from both Ch. punctata and Ch. formiceticola by its body shape and size. In their catalogue, Gemminger & Harold (1868: 905) gave the distribution of Ch. attae as Greece, and Schaufuss (1876a: 396) made comments about its morphology. Reitter (1875: 301) examined the type of Ch. attae and found it similar to Ch. formiceticola, but he listed both Ch. attae and Ch. formiceticola as junior synonyms of Ch. formicaria, an action accepted by Heyden et al. (1883: 80) in their catalogue. Walker (1888: 181, 1892: 248) reported “ Ch. attae ” from Gibraltar, but it was most likely Ch. formiceticola, and Fauvel (1890: 338) resurrected Ch. attae as a valid taxon. A report by Wasmann (1890: 298) in Tunisia was most likely of Ch. punctata or Ch. gallica. Bodemeyer (1900: 24) recorded “ Ch. attae ” in “Karakeuy” (Istanbul), but we have examined some of Bodemeyer’s material and have identified it as Ch. balcanica. Reitter (1902: 5) compared the morphology of Ch. attae with that of C. gallica.

Rücker (1980: 144) included Ch. attae in his key for the identification of Cholovocera species, illustrating the median lobe of the aedeagus (Rücker 1980: 145, fig. 22); also, he gave the geographic distribution of this species as: southern Europe, Greece and northern Africa, probably taken from previous publications, and associated it with Messor barbarus and M. structor. Later, Rücker (1983: 4) reported Ch. attae from southern Hungary, including a habitus figure with metatibiae equal to those of Ch. balcanica, and an illustration of the distal part of the aedeagus (Rücker 1983: 5, fig. f), which also matches that of Ch. balcanica. Actually, Rücker’s (1983) figure f, labelled as Ch. attae, represents the same species as his figure e, labelled as Ch. balcanica, but both shown in different orientations.

D̂bler (1987: 14) recorded two “ syntypes ” of Ch. attae in the SDEI collection, although one of them had a label reading “ Holotypus ” (see Rücker 2011a: fig. 12). Audisio et al. (1995: 9) and Angelini & Rücker (1999: 218) cited Italy as the location of Ch. attae, again an error by confusing this species with Ch. punctata. We have examined over 30 specimens collected by F. Angelini in Sicily, which we have identified as Ch. punctata (see Material examined above).

Sár et al. (2004: 331) expressed doubt about the record of Ch. attae in Rücker (1983) from Hungary, commenting that some records and specimens found in museum collections labelled as from “ Hungary ”, may have originated from territories which once were part of the Austro-Hungarian Empire, such as Serbia, Romania and nearby countries. Therefore, we believe that, unless specimens are re-examined, reports from those countries should be considered doubtful, probably referring to Ch. balcanica.

Several subsequent catalogues and checklists gave large geographic distributions for Ch. attae, including France, Greece, Italy, Hungary, Serbia, Montenegro, Algeria, Morocco and Tunisia, which we regard as erroneous, except for Greece (L̂bl & Smetana 2007: 557; Rücker 2009: 14, 2011b; Shockley et al. 2009b: 65). However, Rücker (2018: 576, 2020: 34) restricted that distribution to only Crete and Rhodes, which is partially correct. Also, Rücker (2018: 576, fig. 1187) gave a detailed description of Ch. attae, including a figure of the aedeagus in lateral view. Finally, Stalling (2019: 13) reported the island of Kos (Dodecanese Islands) as a new locality for Ch. attae in Greece.