Cholovocera gallica (Schaufuss, 1876) Delgado & Palma, 2023

Delgado, Juan A. & Palma, Ricardo L., 2023, A revision of the genus Cholovocera Victor, 1838 (Coleoptera: Endomychidae), European Journal of Taxonomy 906, pp. 1-71 : 38-41

Cholovocera gallica (Schaufuss, 1876) new status

Figs 4D , 7D , 9A , 10D , 12D , 13C , 14F–G , 20 , 25A–B

Coluocera gallica Schaufuss, 1876a: 398 .

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

Coluocera fleischeri Reitter, 1902: 5 . Syn. nov.

Cholovocera fleischeri Reitter [sic] – Rücker 1980: 144, fig. 23 — Audisio et al. 1995: 9.

Differential diagnosis

Cholovocera gallica is morphologically and geographically close to Ch. formiceticola , but these species can be separated by the shape of the pronotum ( Fig. 10D against Fig. 11B ) and of the metatibiae ( Fig. 14F–G against Fig. 14J–K ). Furthermore, Cholovocera gallica can be distinguished from all other species by having a bulbous pronotum with a wide anterior margin ( Fig. 10D ), and large, subtriangular terminal antennomeres ( Figs 12D , 13C ).

Also, the shape of the aedeagus and paramere ( Fig. 20 ), as well as of the spermatheca ( Fig. 7D ), are useful characters to distinguish Cholovocera gallica from all other species in the genus.

Type material

Cholovocera gallica : as far as we know, the type material was not examined by any author after the original description of Ch. gallica . In our opinion, that may be the reason for the species having been synonymised and not re-evaluated until now. Dr Bernd Jaeger found two syntypes, one male and one female, in the Schaufuss Collection held at MFNB, which he kindly made available for examination.

Considering the great number of misidentifications of the species of Cholovocera , both in collections and literature, and the fact that the syntypes of Ch. gallica belong to two species, it is advisable to designate a lectotype to give this name taxonomic stability (Article 74.7.3, ICZN 1999). We hereby designate the syntype male from the Schaufuss Collection deposited in MFNB, with labels reading: “ Cholovocera formicaria Motsch., Gall merid” and “ Syntype Coluocera gallica Schaufuss, 1876 , labelled by MFNB 2021” as the lectotype of Cholovocera gallica ( Fig. 25A ). The syntype female becomes a paralectotype ( Fig. 25B ), but it is a misidentified specimen of Ch. punctata .

Coluocera fleischeri : lectotype male and four paralectotypes held in HNHM.

Lectotype of Coluocera gallica

FRANCE – 1 ♂; “ Gallia meridional ”; [Schaufuss Collection], MFNB. Designated below.

Paralectotype of Coluocera gallica

FRANCE – 1 ♀; “ Gallia meridional ”; [Schaufuss Collection], MFNB. Reidentified as Ch. punctata .

Lectotype of Coluocera fleischeri (designated by Rücker (2011a: 13)

CROATIA – Dubrovnik-Neretva • 1 ♂; “Dalmatia, Metkovic”; HMHN.

Paralectotypes of Coluocera fleischeri

CROATIA – Dubrovnik-Neretva • 2 ♂, 2 ♀; “Dalmatia, Metkovic”; HMHN.

Notes

As can be seen in Rücker 2011a (fig. 17), 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 handwriting of the Holotypus label, we conclude that it was added at a later date than the description by Reitter (1902).

Additional material, non-types

SPAIN – Catalonia • 1 ♂, 2 ♀♀, 10 specimens; B[arcelona], Bellaterra; 25 Aug. 1980; X. Espadaler leg.; “nid [nest] Messor barbarus ”; MHNG .

FRANCE – Languedoc-Rousillon • 1 ♂; Collioure; Gambey leg.; ZFMK • 1 ♂; “P.O.” [Pyrénées-Orientales], Collioure; MHNG • 1 ♀; “Pyr. O” [Pyrénées-Orientales], Collioure; Dr Normand leg. [associated with a Messor worker ant labelled: “ Messor sp. , P. Werner det. 2016”]; NMPC • 1 ♂; Agde; MHNG .

ITALY – Liguria • 2 ♀ ♀; Genova ; Nov. 1892; A. Solari leg.; MFNB – Sardinia • 1 ♂, 1 ♀; North of Bolóntana ; 850 m a.s.l.; 13 Apr. 1992; J. Scheuern leg. [one specimen associated with a Messor worker ant, the second with a Camponotus worker ant] NKME • 1 ♂, 1 ♀; 8 km Northeast of Lula ; 250 m a.s.l.; J. Scheuern leg. [one specimen associated with three Messor worker ants]; NKME • 1 ♂; S.of Teresa ; Jun. 1968; Palm leg.; MZLU 2020-065 • 1 ♀; Lago Baratz ; 23 May 1995; F. Angelini leg.; MCVR • 1 ♀; MCNM 303883 • 1 ♂; Nuoro, Altopiano della Campeda ; 580 m a.s.l.; 18 May 2006; Starke leg.; NHMW • Sicily • 1 ♀; Palermo, Ficuzza ; 700 m a.s.l.; 1–4 May 2000; F. Angelini leg.; “Bosco leccio” [oak forest]; NMPC • 1 ♂; Palermo; NHMB • 1 ♂, 1 specimen; Ficuzza ; 16. Mar. 1942; SMTD • 1 ♂, 1 ♀; Ficuzza ; 16 Mar. 1942; NKME • 1 ♀; Ficuzza ; 16 Mar. 1942; MFNB • 26 specimens; Ficuzza; 1906; O. Leonhard leg.; SDEI 11948–11955 • 3 ♂♂, 4 ♀♀; Randazzo ; 6 May 1933; W. Liebemann leg. [one specimen associated with a Messor worker ant]; SDEI 10860–10866 • 1 ♂; Messina ; 1906; O. Leonhard leg.; SDEI 11940 • 1 ♂; Scanzano, Palermo, Marineo ; 525 m as.l.; 9 Apr. 1993; F. Angelini leg.; MZLU 2020-002 • 8 specimens; Palermo, “ N. Ti ” [North of] Madonie, “dint.” [inside the city of] Isnello ; 700 m a.s.l.; 9 Jun. 1991; F. Angelini leg.; MCVR • 1 ♂; Campofelice ; 28 Apr. 1980; T. Palm leg.; MZLU 2020 /009 • 3 specimens; Erica ; 10 Dec. 1993; Sabella leg.; “Bosco misto” [mixed forest]; NMPC • 6 specimens; Mount Sfaracavallo ; 4 Apr. 1925; Dr Rambousek leg.; NMPC – Sicily, no specific locality • 3 ♂♂; SDEI 11927 and 11934–11935 • 1 ♂; NHMB • 1 specimen; Sicily; SFUN – Tuscany • 3 ♀♀; SMNH • 1 ♂; Bertolini leg.; MFNB – Lazio • 1 ♂, 3 specimens; Maccarese; P. Luigioni leg.; SFUN • 2 ♂♂, 4 ♀♀; Maccarese; P. Luigioni leg.; MFNB • 1 ♀; Roma, Maccarese ; 26 Feb. 1911; P. Luigioni leg.; MFNB – Calabria • 1 ♂; Antonimina ; 1905; SDEI 11909 • 1 ♂; Antonimina ; 1905; Paganetti leg.; SDEI 11908 • 1 ♂; Antonimina; 1905; Paganeti leg.; NMPC • 1 ♂; Gerace ; Paganetti leg.; SDEI 10856 • 1 ♂, 12 specimens; Gerace; Paganetti leg.; NMPC • 1 specimen; Gerace; Paganetti leg.; SMTD • 6 specimens; Aspromonte, San Luca ; 200 m a.s.l.; 28 Apr. 2002; F. Angelini leg.; “Prato” [meadow]; MZLU 2020-003 • 1 ♂, 6 specimens; Aspromonte, Africo ; 50 m a.s.l.; 14 Apr. 1997; F. Angelini leg.; “Prato” [meadow], MCVR • 1 ♂, 5 specimens; Sambiase ; May 1920; C. Minozzi leg.; [each specimen associated with a Messor worker ant]; NHMB – Puglia • 1 ♂, 2 ♀♀; Murgia, San Basilio; Paganetti leg.; SDEI 10818 and 10840–10841 • 1 ♂; Murgia, San Basilio; NHMB • 1 ♂, 3 specimens; Bari ; Nov. 1984; L. De Marzo leg.; MCVR • 1 ♂, 3 specimens; Rutiglieno ; Nov. 1991; L. De Marzo leg.; MCVR – Italy, no specific locality • 1 ♂, 1 ♀; Italia, NMPC .

ALGERIA – Algier • 1 ♂; Lambèze [modern Tazoult]; Jun. 1885; L. Bleuse leg.; ZFMK .

TUNISIA – Mahdia • 1 ♂, 1 specimen; Tunisia, El Djem; 2 Apr. 1925; Dr Rambousek leg.; “fourm.” [ants or ant nest]; NMPC .

CROATIA – Zadar • 1 ♂, 1 ♀; “D”[almatia], Diklo ; Jul. 1913; Novak leg.; NHMB • 1 ♀; “D”[almatia], Diklo ; 7 Jul. 1913; CNHM • 1 specimen; “D”[almatia], Diklo ; Jul. 1913; Novak leg.; MNHS • 1 ♂; “D”[almatia], Diklo ; Jul. 1913; Novak leg.; MFNB • 1 specimen; “D”[almatia], Zara; Novak leg.; MNHS • Split • 1 ♂; “D”[almatia]; Novak leg.; 15 Apr. 1928; [associated with a Tetramorium worker ant]; SFUN • 1 specimen; Salona ; Karaman leg.; MNHS – Dubrovnik-Neretva • 6 specimens; “Dalmatia, Ragusa” [modern Dubrovnik ]; Dr Fleischer leg.; NMPC • 1 ♂, 3 ♀♀; “ Dalmatia ”, Metkovic; SDEI 05781 and 11910 • 1 ♂; “ Dalmatia ”, Metkovic; SFUN • 1 ♀; “Dalmatia” Metkovic ; Formanek leg.; NHMB .

BOSNIA AND HERZEGOVINA – Herzegovina-Neretva • 1 ♂, 1 specimen; Herzegowina, Jablanica; SFUN .

Type locality

“Südfrankreich” [ Gallia meridional ], Southern France.

Description

Male as in Fig. 10D . Body length: 1.33 mm average, range 1.30–1.50 mm (N = 23, males and females). Shape of body oval, pronotum wide and dorsally bulbous, with rounded elytral apex. Terminal antennomeres large, subtriangular. Metatibiae long and narrow, with sinuous margins ( Fig. 14F–G ). Prosternal process markedly keeled anteriorly, with a wide median constriction and subtriangular distally ( Fig. 4D ). Male last visible ventrite with a slight emargination and bordered by a brush of long setae.

Median lobe of aedeagus in ventral view tapering markedly in its distal third, with a round tip ( Fig. 20A ). Aedeagus in lateral view as in Fig. 20C . Distal portion of paramere short, quadrangular, with an irregular tip ( Fig. 20B ), bearing five medium setae ( Fig. 20A–B ). Spermathecal duct short and reservoir straight; ramus short and rounded, cornu short and nodulus long and conical ( Fig. 7D ).

Geographic distribution

The known distribution of Cholovocera gallica is the central Mediterranean, extending from the Balkans in the east to Catalonia in the west, and from northern Italy to Algeria and Tunisia in the south ( Fig. 9A ).

Host ants

There is almost no published information about the ants associated with Cholovocera gallica . Rücker (1980, 1983, 2018) mentioned unidentified species of the genus “ Atta ” as hosts of Ch. fleischeri (now Ch. gallica ). However, species of Atta live exclusively in the Neotropical Region, and therefore cannot be hosts of this beetle species. Lundberg et al. (1987: 123) reported Ch. fleischeri from a large nest of Camponotus in Sicily.

Our examination of Ch. gallica material preserved with ant specimens showed the following associations: (1) with an unidentified species of Messor in southern France, in Andalusia, in Sardinia and in Sicily; (2) with Messor barbarus (det. X. Espadaler) in Spain; (3) with an unidentified species of Camponotus in Sardinia; (4) with an unidentified species of Tetramorium in Croatia.

Junior synonym

Coluocera fleischeri Reitter, 1902

Reitter (1902: 5) described Co. fleischeri from specimens collected near Metkovic, Dalmatia ( Croatia). Rücker (1980: 144) included Ch. fleischeri in his key for the identification of Cholovocera species, illustrating the median lobe of the aedeagus ( Rücker 1980: 145, fig. 23); also, he gave the geographic distribution of this species as Dalmatia, Yugoslavia. Further, Rücker (1983: 4–5) added Herzegovina to the distribution and included a figure of a partial aedeagus. Lundberg et al. (1987: 123) reported Ch. fleischeri from Sicily. Audisio et al. (1995: 9) mentioned Ch. fleischeri in Italy, and Angelini & Rücker (1999: 218) in Puglia y Basilicata ( Italy), but both records were based on the same material collected by F. Angelini in association with “ ants ”, without an identification. An additional locality was reported by Lo Cascio et al. (2006: 325) who recorded Ch. fleischeri in Lipari Island (Aeolian Islands, north of Sicily, Italy). Subsequent catalogues and checklists increased the geographic distributions of Ch. fleischeri even more, adding Macedonia (L̂bl & Smetana (2007: 557), Malta, Montenegro and Serbia ( Shockley et al. (2009b: 65), Hungary ( Rücker 2011b), and Corsica and Tunisia ( Rücker 2020: 34). Although we have not seen material of Ch. fleischeri (as Ch. gallica ) from Corsica, Malta, Macedonia, Montenegro and Serbia, we cannot rule out the possibility that this beetle occurs in those localities. However, we believe that the record from Hungary needs confirmation as it is unlikely to be correct. Finally, Rücker (2018: 578, figs 1188–1189) gave a detailed description of Ch. fleischeri , including a figure of the aedeagus in lateral and ventral views.

Notwithstanding the many reports of this species as Ch. fleischeri , we have examined its holotype male and compared it with many males of Ch. gallica , including the lectotype, without finding any significant morphological difference that would justify the separation of these species. Therefore, we have no hesitation in placing Coluocera fleischeri as a new junior synonym of Ch. gallica .

Taxonomic history and remarks

Schaufuss (1876a: 398) described Cholovocera gallica from southern France in great detail, comparing it with material from Corsica ( Fig. 25C ), the Balearic Islands ( Fig. 25D ), Algeria ( Fig. 25E ) and Sardinia. Although his identifications of those specimens were not all correct, our study showed that he had Ch. punctata and Ch. formiceticola for comparison. However, one year later, Reitter (1877: 5) placed Ch. gallica as a junior synonym of Ch. formicaria , a status which was accepted by Belon (1879: 192), and a number of subsequent catalogues, such as those by Heyden et al. (1883: 80), Rücker (2009: 14), Shockley et al. (2009b: 65) and Rücker (2020: 34). Other authors, with the exception of L̂bl & Smetana (2007: 557), did not mention Ch. gallica at all, but recorded and listed Ch. fleischeri instead.

However, from our examination of the lectotype of Ch. gallica and many other samples from a wide geographical area ( Fig. 9A ), we believe that Ch. gallica is a distinct species, which we herewith resurrect as a valid taxon.

Figures

Fig. 4. Prosternal processes. A. Cholovocera afghana Johnson, 1977. B. Ch. attae (Kraatz, 1858). C. Ch. balcánica (Karaman, 1936). D. Ch. gallica (Schaufuss, 1876). E. Ch. formicaria Victor, 1838. F. Ch. formiceticola (Rosenhauer, 1856). G. Ch. occulta sp. nov. H. Ch. punctata (Märkel, 1845).

Fig. 7. Spermathecae, lateral view. A. Cholovocera afghana Johnson, 1977. B. Ch. balcánica (Karaman, 1936). C. Ch. occulta sp. nov. D. Ch. gallica (Schaufuss, 1876). E. Ch. formicaria Victor, 1838. F. Ch. formiceticola (Rosenhauer, 1856). G. Ch. punctata (Märkel, 1845). Abbreviations: See Material and methods. Scale bar = 0.1 mm.

Fig. 9. Known geographical distribution of the species ofCholovocera Victor, 1838.A. Ch. formiceticola (Rosenhauer, 1856) (red dots) and Ch. gallica (Schaufuss, 1876) (yellow dots). B. Ch. punctata (Märkel, 1845) (red dots) and Ch. occulta sp. nov. (yellow dots). C. Ch. balcanica (Karaman, 1936) (red dots), Ch. attae (Kraatz, 1858) (yellow dots), Ch. formicaria (dark blue dots) and Ch. afghana Johnson, 1977 (orange dot).

Fig. 10. Habitus, dorsal view. A. Cholovocera afghana Johnson, 1977, ♀. B. Ch. attae (Kraatz, 1858), ♂. C. Ch. balcanica (Karaman, 1936), ♂. D. Ch. gallica (Schaufuss, 1876), ♂. Scale bar = 0.5 mm.

Fig. 12. Terminal female antennomeres. A. Cholovocera afghana Johnson, 1977. B. Ch. attae (Kraatz, 1858). C. Ch. balcánica (Karaman, 1936). D. Ch. gallica (Schaufuss, 1876). E. Ch. formicaria Victor, 1838. F. Ch. formiceticola (Rosenhauer, 1856). G. Ch. occulta sp. nov. H. Ch. punctata (Märkel, 1845). Scale bar = 0.1 mm.

Fig. 13. Terminal male antennomeres.A. Cholovocera attae (Kraatz, 1858). B. Ch. balcánica(Karaman, 1936). C. Ch. gallica (Schaufuss, 1876). D. Ch. formicaria Victor, 1838. E. Ch. formiceticola (Rosenhauer, 1856). F. Ch. occulta sp. nov. G. Ch. punctata (Märkel, 1845). Scale bar = 0.1 mm.

Fig. 14. Metatibiae. A. Cholovocera afghana Johnson, 1977, ♀. B. Ch. attae (Kraatz, 1858), ♀. C. Ch. attae (Kraatz, 1858), ♂. D. Ch. balcanica (Karaman, 1936), ♀. E. Ch. balcanica (Karaman, 1936), ♂. F. Ch. gallica (Schaufuss, 1876), ♀. G. Ch. gallica (Schaufuss, 1876), ♂. H. Ch. formicaria Victor, 1838, ♀. I. Ch. formicaria Victor, 1838, ♂. J. Ch. formiceticola (Rosenhauer, 1856), ♀. K. Ch. formiceticola (Rosenhauer, 1856), ♂. L. Ch. occulta sp. nov., ♀. M. Ch. occulta sp. nov., ♂. N. Ch. punctata (Märkel, 1845), ♀. O. Ch. punctata (Märkel, 1845), ♂. Scale bar = 0.1 mm.

Fig. 20. Aedeagus and paramere of Cholovocera gallica (Schaufuss, 1876). Lectotype (Southern France). A–B. Ventral views. C. Lateral view. Scale bar = 0.1 mm.

Fig. 25. Specimens of Cholovocera Victor, 1838 from the Schaufuss Collection. A. Syntype (now lectotype) of Ch. gallica (Schaufuss, 1876) from Southern France. B. Misidentified syntype (now a paralectotype) of Ch. gallica from Southern France (actually a female of Ch. punctata (Märkel, 1845)). C. Specimen of Ch. punctata from Corsica. D. Specimen of Ch. formiceticola (Rosenhauer, 1856) from the Balearic Islands. E. Specimen of Ch. punctata from Algeria.

Fig. 11. Male habitus, dorsal view. A. Cholovocera formicaria Victor, 1838. B. Ch. formiceticola (Rosenhauer, 1856). C. Ch. occulta sp. nov., paratype. D. Ch. punctata (Märkel, 1845). Scale bar = 0.5 mm.