Cholovocera balcanica ( Karaman, 1936 )

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 : 41-48

Cholovocera balcanica ( Karaman, 1936) new status

Figs 4C , 7B , 9C , 10C , 12C , 13B , 14D–E , 21 , 23H , 24F–G

Reitteria balcanica Karaman, 1936: 131, figs a–f.

“ Cholovocera major ” – Rücker 2011a: 12. Non Coluocera formicaria v. major Reitter, 1887 .

Differential diagnosis

Cholovocera balcanica is sympatric with Ch. attae , but they can be distinguished by the larger size and longer legs of the former species. Furthermore, Ch. balcanica can be clearly separated from Ch. attae and from all other species in the genus by the morphology of the median lobe of the aedeagus, and the paramere ( Fig. 21 ). The spermatheca of Ch. balcanica is also diagnostic, in particular because of its large nodulus ( Fig. 7B ).

Type material

Cholovocera balcanica : Karaman (1936) examined 18 specimens (syntypes) from two localities for her description of Reitteria balcanica . However, only five of them have been located in the collection of the Faculty of Agricultural Science and Food, Saints Cyril and Methodius University ( CMUS) in Skopje, Republic of North Macedonia ( Fig. 23H ). Furthermore, in the collection of the Senckenberg German Entomological Institute ( SFUN), there are five specimens from Skopje identified as “ Reitteria balcanica ” which may be syntypes but, as their labels do not include a date, their status cannot be confirmed. Alternatively, these five beetles (see Material examined, non types below) may be part of the samples collected after 1936 and reported by Karaman (1964: 33).

Syntypes and voucher specimens of Reitteria balcanica , not examined*

NORTH MACEDONIA – Skopje • 3 syntypes; Vodno ; 7 Jun. 1936; “ Mravinjak ” [ant nest]; CMUS, 245 . • 2 specimens; Vodno ; 7 Jun. 1936; CMUS, C164 • 1 specimen; Rasce ; 5 Feb. 1959; “Mravinjak” [ant nest]; CMUS, 277 • 1 specimen; Stip; 17 Apr. 1960; [associated with an ant labelled as: “ Camponotus ligniperdus, Det. Zora Karaman ”]; CMUS • 1 specimen: Skopska Crna Gora [ Black Mountain of Skopje ]; 10 Jun. 1960; CMUS ( Fig. 23H ) .

* Despite our request to loan the above eight specimens from CMUS, we were not able to examine them. However, we believe it is important to record them here, including their photograph, kindly provided by Vladimir Krpach ( CMUS).

Additional material, non-types

NORTH MACEDONIA – Skopje • 1 ♂, 4 ♀ ♀; “Serbia, S. Makedonia”, Skopje; [one specimen associated with a Camponotus worker ant]; SFUN .

ALBANIA – Vlora • 2 ♂♂, 1 ♀; “M.E.” [ Mount ] Tartarit ; Jun. 1932; Bischoff leg.; MHNB • 1 ♀; “M.E.” [Mount] Tartarit ; Jun. 1932; Bischoff leg.; MFNB – Gjirikaster • 1 ♀; Albania mer. [idional], Tepelene; May 1931; Winkler leg.; [associated with a Messor worker ant]; MHNG.

BULGARIA – Dobrich • 1 specimen; Albena; 31 Jan. 1990; Batelka leg.; NMPC • 6 specimens; Albena; 14 Sep. 1990; Batelka leg.; NMPC – Lovech • 1 ♂; Tirnowa [modern Veliko-Tarnovo]; May 1994; Flach leg.; SMTD • 4 ♀♀; Tirnowa [modern Veliko-Tarnovo]; May 1994; Flach leg.; SFUN – Burgas • 1 ♂, 1 ♀; Karabajir, Rumel; 3 Apr. 1909; Rambousek leg.; SDEI 10773–10769 • 1 ♂, 10 specimens; Karabajir, Rumel; 3Apr. 1909; Rambousek leg.; [three specimens, each associated with a Messor worker ant labelled as: “ Messor structor (Latreille) P. Werner det. 2016”]; NMPC • 3 ♀ ♀; Burgas; Mar. 1909; Rambousek leg.; [two specimens, each associated with a Messor worker ant]; SDEI 10770 and 10774–10775 • 3 ♂ ♂, 2 ♀♀; Flach leg.; SDEI 10854–10855 and 10782–10784 • 1 ♂, 18 specimens; Burgas; May 1895; Flach leg.; SMTD • 1 specimen; Burgas; May 1895; Flach leg.; SFUN • 1 specimen; Burgas; Flach leg.; SFUN • 1 ♂, 19 specimens; Burgas; May 1894; Flach leg.; SFUN • 1 ♂, 1 ♀; Burgas, May 1894; Flach leg.; NMPC • 1 specimen; Burgas; NMPC • 2 specimens; Burgas; Rambousek leg.; [associated with a Messor worker ant]; NMPC • 2 specimens; Burgas “maritime”; Apr. 1909; F. Rambousek leg.; NMPC • 1 specimen; Rumel, Bačkovo; 11 May 1909; Rambousek leg.; [associated with a Messor worker ant, labelled as: “ Messor structor (Latreille) P. Werner det. 2016”]; NMPC • 4 specimens; Bulgaria Oriental, Zeitinburun [Mount Zeytin Burnu]; Apr. 1933; “Mař Táb.” [Mařan and Táborský leg.]; NMPC • 1 ♂, 4 specimens; Bulg. Mac.; Jun. 1933; “Mař Táb.” [Mařan and Táborský leg.]; NMPC • 1 specimen; Burgas; Rambousek leg.; MNHS • 5 specimens; Bulgaria or.[iental], Bozurec; Ernest leg.; NMPC.

GREECE – Corfu • 2 ♂♂; S. Corfu, Paxos , 2 km NW of Galos; 17 Apr. 1981; Scheuern leg.; “Oliven hain” [olive grove]; [one specimen assocaited with a Messor worker ant]; ZFMK • 1 ♀; Corfu monats., Paleaskastritza ; 10 Apr. 1972; V. Mahnert leg.; MHNG – Tessaly • 1 ♂, 1 ♀, 16 specimens; Pelion; MHNB – Ionian Islands • 3 ♀♀; Cephalonia, Argostoli; NHMB • 1 ♂; Cephalonia ; Paganetti leg.; SMTD • 1 ♂; Cephalonia ; Paganetti leg.; NMPC – Central Greece • 1 ♂, 2 ♀♀; Parnassos; SDEI 10790–10791 and 10856 • 2 ♂♂, 1 ♀, 3 specimens; “Balkan Parnass.”; 1903; Paganetti leg.; SMTD • 1 ♂; “ Parnass ”; Paganetti leg.; NMPC – Attica • 1 ♀; Greece, Atica; SMTD – Peloponnese • 5 ♂♂, 9 ♀♀, 25 specimens; Ahaïa, Erimanthos–Geb., Kaletzi ; 37°57′04″ N, 21°46′17″ W; 1100 m a.s.l.; 23 Apr. 1998; Zerche leg.; “Tannenwald oberh.[alb]” [above fir forest],; [each specimen associated with a Messor worker ant]; SDEI 11792–11816 and 11877–11907 • 1 ♂, 2 specimens; Ahaïa, surroundings of Panahaiko, near Ano Kastrisi ; 38º15′54″ N, 21º50′42″ W; 900 m a.s.l.; 28 Mar. 1997; L. Zerche leg.; “unter Steinen” [under stones]; [each specimen associated with a Messor worker ant]; SDEI 11807 and 11811–10815 • 1 ♂, 11 specimens; Kalavyrta, Morea; MHNB • 2 ♀; Nauplia; SFUN • 1 ♂, 1 specimen; Nauplia ; Mar. 1890; Hänel leg.; SMTD • 1 ♀; Ilia, Erymanthos mts , 1.5 km NNW of Orini; 1290– 1370 m a.s.l.; 9 May 2013; Schuh leg.; NHMW – Thrace • 2 ♂♂, 13 specimens; “ Alexandrupolis ”; 2 May 1937; Bartoň leg.; NMPC – Crete • 2 ♂♂, 17 specimens; Omalos, Lefka Ora; 27 May–1 Jun. 1980; Brodský and Bílý leg; NMPC • 2 ♂♂; Omalos, Lefka Ori Mts ; 3 Jun. 1984; Bílý leg.; NMPC • 1 ♀; NW Crete, Dikti Oros, Limnakaro ; 35º08′08″ N, 25º29′00″ E; 1170 m a.s.l.; 7 Apr. 2012; V. Assing leg.; Messor nest; NMPC • 2 ♀♀; Crete, Lasithi Geb. [iet] [surroundings of]; MFNB • 3 ♂♂, 5 ♀♀, 10 specimens; East Crete, Lasithi Plateau, road Kaminaki– Embaros , Mesovouni Pass ; 1150 m a.s.l.; 5 Jun. 2010; Schuh leg.; NHMW • 1 ♂; Crete; NMPC – Sporades Islands • 4 ♂♂, 2 ♀♀; Samos, “val. des Rossignols” [Nightingale valley], “sous” [near] Manolates ; 100 m a.s.l.; 17 May 1985; Besuchet leg.; MHNG • 2 ♀ ♀; South Sporades, Karpathos; V. Oertzen leg.; MFNB – Dodecanese • 2 ♂♂, 1 ♀; Rhodas, Ebonas ; 15 Apr. 1977; Cl. Besuchet leg.; MHNG – Greece, no specific locality • 1 ♀; Greece; 1901; Sabel leg.; SFUN • 1 ♂, 2 specimens; Greece; SMTD • 1 ♂, 1 ♀; Greece; SMTD .

TURKEY – Marmara • 1 ♂; “Asia minor”, Karakeuy ; V. Vodemeyer leg. [for Bodemeyer]; [associated with two Messor worker ants]; SFUN • 1 ♂; Bursa; 1 Apr. 1917; [associated with a Messor worker ant]; SMTD – Western Anatolia, Aegean • 1 ♀; Smyrna, SMNH • 1 ♀; Smyrna; NMPC • 1 ♀; Smyrna; MFNB • 2 ♀ ♀; Ephesus; J. Sahlb[erg]. leg.; HMHN • 1 ♀; Ephesus; J. Salhb [erg]. Leg.; SFUN – Mediterranean • 1 ♀; Mersin, 10[sic] km NW of Silifke ; 27 Apr. 1978; Besuchet and Löbl leg.; MHNG .

UKRAINE • 1 ♀; Odesa; Dr Lgocki leg.; SMTD • 1 ♀; Kherson; 17 Apr. 1941; Lasovko leg.; NHMW .

Description

Male as in Fig. 10C . Body length: 1.40 mm average, range 1.30–1.50 mm (N = 30, males and females). Shape of body elliptical, with the lateral margins of the pronotum continuous with those of the elytra, i.e., without an indentation. Elytral apex pointed. Terminal antennomeres subtriangular, sexually dimorphic ( Figs 12C , 13B ). Metatibiae with straight margins, diverging distally ( Fig. 14D–E ). Prosternal process slightly keeled anteriorly, with a slight median constriction and distally rhomboidal ( Fig. 4C ). Male last visible ventrite with a slight emargination and bordered by a brush of short setae.

Median lobe of aedeagus long and sinuous in ventral view, tapering and acutely pointed distally ( Fig. 21A ). Aedeagus in lateral view as in Fig. 21C . Distal portion of paramere long, conical, with a round apex bearing a brush of many long setae in ventral view ( Fig. 21B ), but polygonal in lateral view ( Fig. 21D ). Spermathecal duct short, spermathecal reservoir long and T-shaped; ramus long and curved distally, cornu round and nodulus greatly developed, almost as long as cornu and ramus together ( Fig. 7B ).

Type localities

“Schlucht von Topolka bei Veles, Vodno-Berg bei Skoplje” [Canyon of Topolka near Veles, and Vodno near Skopje], Republic of North Macedonia.

Geographic distribution

The known distribution of Cholovocera balcanica extends from the Balkan Peninsula in the west to eastern Turkey, and from the northern Black Sea coast and the Crimean Peninsula in the north to Crete and other Greek islands in the south ( Fig. 9C ).

Host ants

Karaman (1964: 33) reported that the specimens from Mount Vodno, the type locality of Ch. balcanica , were collected in nests of Tetramorium caespitum (Linnaeus, 1758) . Also, Karaman (1964: 33) recorded Ch. balcanica in nests of T. caespitum and Camponotus ligniperda (Latreille, 1802) from Skopska Crna gora (Black Mountain, near Skopje) and Rasce (near Skopje), but associated with Messor in Stip (E. of Macedonia) and around Mostar ( Bosnia and Herzegovina).

Our examination of Ch. balcanica material preserved with ants showed that it is frequently found with species of Messor in Albania and Greece, and with Messor structor (det. P. Werner) in Bulgaria. The association with Camponotus ligniperda given by Karaman (1964) needs confirmation of the species, although the genus is correct. We have examined one specimen from North Macedonia identified by Zatia Karaman (see above) and we agree that it belongs to Camponotus .

Taxonomic history and remarks

The original description of Reitteria balcanica by Karaman (1936: 131, figs a–f) is well illustrated, including figures of an antenna, two metatibiae, the aedeagus and habitus in dorsal and ventral views; these illustrations agree with the same characters of the material we have examined and identified as Cholovocera balcanica . Karaman (1964: 33) published new locality records from the Balkans, based on additional samples collected after her 1936 paper and from her examination of museum collections; these records were Skopska Crna Gora (Black Mountain, near Skopje), Rasce (near Skopje), Stip (Eastern Macedonia) and around Mostar ( Bosnia and Herzegovina). Also, Karaman (1964: 33) mentioned to have studied many specimens from Spalato (Split, Croatia) deposited in the collection of the Institute of Crop Protection of Belgrade (now Institute for Plant Protection and Environment). This large collection was later divided between the Natural History Museum in Zagreb (I. Toševski, pers. comm. 17 Jun. 2021) and the Museum of Natural History in Split (V. Mičetić, pers. comm. 3 Sep. 2021). However, despite our enquiries, Karaman’s specimens have not been located.

Without examining the types of Reitteria balcanica, Rücker (2011a: 12) placed this species as a junior synonym of Cholovocera major .Apparently, Rücker (2011a: 13) missed the similarity between Ch. major and Ch. formicaria (see synonymy above) and, instead, he associated Ch. major with Ch. balcanica based on the morphology of the aedeagus, and on an earlier misinterpretation of type localities, as follows: Rücker (1980: 144) gave the type locality of Ch. major as “ Angora (Talysch-Gebiet)”; however, “ Angora ” is situated in Anatolia ( Turkey), and not in the Talish Region, which is located in the south of the Caucasus in Azerbaijan. Thus, Rücker (2011a: 12) believed that Ch. major and Ch. balcanica were sympatric and synonymised them. This synonymy was maintained by Rücker (2020: 34) in his latest checklist. Although we do not agree with the placement of Reitteria balcanica as a junior synonym of Ch. major as proposed by Rücker (2011a: 12), we agree that it belongs to the genus Cholovocera .

From our examination of five specimens from the type locality and many other samples from six countries (see Material examined) we conclude that Cholovocera balcanica 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. 21. Aedeagus and paramere of Cholovocera balcanica (Karaman, 1936). Male from type locality (Skopje). A–B. Ventral views. C–D. Lateral views. Scale bar = 0.1 mm.

Fig. 23. Types and relevant specimens of species of Cholovocera Victor, 1838. A. Syntypes of Ch. formicaria Victor, 1838 on original Motchoulsky’s mounting card (courtesy of A.A. Gusakov). B. Male specimen of Ch. formicaria Victor, 1838 from the type locality. C. Lectotype of Ch. punctata (Märkel, 1845) on left, and a specimen from the J.W. Helfer’s Collection (NMPC) on right, showing identical pins and mounting cards. D. Single label attached to five females of Ch. punctata (Märkel, 1845) from the Helfer Collection. E. Label of Ch. punctata (Märkel, 1845) lectotype. F. Lectotype (above) and paralectotype (below) of Ch. formiceticola (Rosenhauer, 1856). G. Label of lectotype of Ch. formiceticola (Rosenhauer, 1856). H. Five syntypes of Ch. balcanica (Karaman, 1936), from left: second, third and fourth pins (courtesy of V. Krpach).

Fig. 24. A. A syntype of Cholovocera formicaria Victor, 1838, dorsal view (courtesy of K.V. Makarov). B. Same specimen, ventral view (courtesy of K.V. Makarov). C. Holotype of Ch. brevicornis Johnson, 1977 (= Displotera beloni (Wasmann, 1899)), dorsal view. D. Same specimen, ocular area, lateral view. E. Ocular area of Ch. formiceticola (Rosenhauer, 1856) (Lisbon), lateral view. F. Aedeagus of Ch. formicaria major, lectotype, ventral view. G. Same specimen, lateral view. H. Aedeagus of Co punctata sardoa, lectotype, ventral view. I. Same specimen, lateral view. Scale bars: A–C = 0.5 mm; D–I = 0.1 mm.