Cholovocera formicaria Victor 1838
Creators: name: Delgado, Juan A.
name: Palma, Ricardo L.
Publisher: name: Zenodo
Year: 2023
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Coleoptera
Endomychidae
Cholovocera
Cholovocera formicaria

description: Cholovocera formicaria Victor, 1838
Figs 4E, 7E, 9C, 11A, 12E, 13D, 14H–I, 15, 23A–B, 24A–B, 24F–G
Cholovocera formicaria Victor, 1838: 179, pl. III.
Colovocera formicaria – Belon 1879: 194.
Cholovocera subterranea – Motchoulsky 1845: 111.
Coluocera formicaria – Heyden et al. 1883: 80.
Coluocera formicaria v. major – Reitter 1887: 10. Syn. nov.
Cholovocera major – Rücker 2011a: 13, figs 18–20.
Differential diagnosis
Cholovocera formicaria is morphologically and geographically close to Ch. afghana and Ch. balcanica, but it can be distinguished from both species by the shape of the metatibiae. Those of Ch. formicaria are club-shaped and sinuous (Fig. 14H–I), but straight and gradually tapering in Ch. afghana (Fig. 14A), and much shorter and wide in Ch. balcanica (Fig. 14D–E).
Furthermore, the aedeagus and the paramere are useful characters to distinguish Cholovocera formicaria from Ch. balcanica and Ch. gallica (Fig. 15 against Figs 20–21).
Type material
Cholovocera formicaria: two female syntypes of Ch. formicaria (Figs23A, 24A–B) held in Motschulsky’s Collection at the University of Moscow (A.A. Gusakov pers. comm. May 2021) were not available for our examination. Instead, we have examined a male of Ch. formicaria from “Derbent” (Fig. 23B), the type locality, held in the Märkel Collection in SMTD (Dresden), which may have been part of Motschulsky’s type series. However, considering its different labelling from the syntypes (Fig. 23A–B), we cannot be absolutely sure of its status. Nevertheless, it is an authenticated specimen of Ch. formicaria, which allows us to make a comparison between this species and the male syntype of Ch. punctata to conclude that they are different species. Victor (1838) also mentioned to have found Ch. formicaria in “Kahétie” (Georgia), but no specimen with such a label have been located.
Coluocera formicaria var. major: lectotype (male) and two paralectotypes (male and female) held in HNHM. The type locality given by Reitter (1887) is “Talish Gebiete” [Talish Region]. However, the label attached to the lectotype reads “Rasano” (see Rücker 2011a: fig. 18, bottom, left), which is located in the region of the Talish Mountains. According to the ICZN Code (1999), the type locality is what is written on the label associated with the type specimen, in this case “Rasano”.
Syntypes, not examined
RUSSIA – Dagestan • 2 ♀♀; Derbent, “Litt. m. csp.” [Caspian Sea region]; ZMUM.
Lectotype of Coluocera formicaria var. major
AZERBAIJAN – Lankaran • 1 ♂; “ Caspi.–M.–Gebiet ” [Caspian Sea region], Rasano [Ancient town in ruins, 15 km southwest of Lerik, 38°40′12.0″ N, 48°18'46.0″ E. See Lazarev (2017)]; Leder leg.; HNHM. Designated by Rücker’s (2011a: 13).
Paralectotypes of Coluocera formicaria var. major
AZERBAIJAN – Lankaran • 2 ♂♂; “ Caspi.–M.–Gebiet ” [Caspian Sea area], Rasano; Leder leg.; [each specimen associated with a Tetramorium worker ant, det. X. Espadaler]; HNHM.
Notes
As it can be seen in Rücker 2011a (fig. 18), 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 (1887).
The same comment given above under the lectotype, applies to the paralectotypes.
Additional material, non-types
RUSSIA – Dagestan • 1 ♀; Derbent; MCNM 198707 • 1 ♂, 1 ♀; Derbent; SMTD • TURKEY – East Anatolia • 2 ♀♀; Kars, Digor; 1650 m a.s.l.; 15 Jun. 1986; MHNG.
AZERBAIJAN – Ĝyĝl • 1 ♂; “Caucasus, Helenendorf” [modern Göygöl]; Couřil leg.; NMPC • 1 ♂; “Caucasus“, Helenendorf “; SMTD. – Ganja • 1 ♂, 1 ♀; “Caucasus, Elisabetpol” [modern Ganja]; SDEI 10850 to 10851 • 1 specimen; “Caucasus, Elisabetpol”; NHMW. – Lankaran • 1 ♂, 3 specimens; Leder leg.; NHMB • 1 ♀; Leder leg.; SDEI 11874 • 1 specimen; Leder leg.; MHNG • 2 ♂, 2 ♀; Leder leg.; NMPC • 1 ♂, 1 ♀; “Talyschgeb[er]g [Talish area], Transcaucas[us].”; Leder leg.; SDEI 11875–11876 • 2 ♂♂, 3 specimens; “Talyschgebg., Transcaucas.”; Leder leg.; SFUN • 1 ♂; “Talyschgebg., Transcaucas.”; Leder leg.; SMTD • 2 specimens; “Talyschgebg., Transcaucas.”; Leder leg.; MFNB • 1 specimen; “Caspi.–M.–Gebiet, Rasano”; Leder leg.; SFUN. – Ordubad • 1 ♀, 1 specimen; “Caucasus, Araxesthal” [Arax River Valley]; Leder leg.; NMPC. – Caucasus, no specific locality • 2 ♂♂; “Kaukas”; Leder leg.; [18]86; SDEI 10842– 10843 • 1 ♂, 2 ♀♀; “Kaukas”; Leder leg.; SDEI 10847– 10849 ” • 1 ♀; “Kaukas”; Leder leg.; NMPC • 1 ♂; “Cauc. Sept.”; NMPC • 1 specimen; “Kaukas”; Leder leg.; ZFMK • 1 ♂; “Kauka”; Leder leg.; ZFMK • 1 ♀; “Caucasus”; Leder leg.; MZLU 2020-001 • 1 specimen; “Kaukasus”; ZFMK • 2 ♀♀; “Caucasus”; SMTD • 1 specimen; “Kaukasus”; MFNB.
IRAN – Guilan • 1 ♂, 2 ♀♀; Chalus-Polzoghal; 29 Apr. 1970; Wittmer and Bothmer leg.; MHNB • 1 ♀; Bandar Pahlavi [modern Bandar-e Anzali]; 20 Aug. 1973; S. Vit leg.; “marais” [marshland]; MHNG. – Mazandaran • 1 ♂, 1 ♀; Dadu; NHMB – Unknown localities • 1 ♂; Nov. 1917; SDEI 10793 • 1 ♂, 1 ♀; MFNB.
Type locality
“Derbent, non loin de la mer Caspienne” [Derbent, not far from the Caspian Sea], Dagestan, Russia.
Description
Male as in Fig. 11A. Body length: 1.43 mm average, range 1.30–1.50 mm (N = 41, 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 markedly acute. Terminal antennomeres subtriangular, with round angles (Figs 12E, 13D). Metatibiae as in Figs 14H and 14I, narrower in the proximal half and with sinuous margins, especially in the male. Prosternal process slightly keeled anteriorly, with a marked median constriction and rounded distally (Fig. 4E). Male last visible ventrite with a slight emargination and bordered by a brush of short setae.
Median lobe of aedeagus sinuous and narrow from the first third of its length in ventral view, tapering and acutely pointed distally (Fig. 15A). Aedeagus in lateral view as in Fig. 15C. Distal portion of paramere long, subcylindrical, sinuous, with a round apex bearing a brush of many setae (Figs 15B, 15D). Spermathecal duct and spermathecal reservoir short; ramus long and curved distally, cornu round and nodulus moderately developed, smaller than cornu and ramus together (Fig. 7E).
Geographic distribution
The known distribution of Cholovocera formicaria is on the Caucasus Mountains and the Caspian Sea coast, extending from eastern Anatolia to north-eastern Iran (Fig. 9C). There is a record from Switzerland (L̂bl & Smetana 2007; Shockley et al. 2009b; Rücker 2011b, 2020), which needs to be reviewed, but we regard it as almost certainly erroneous.
Host ants
Determining the identity of the species of ant hosting Cholovocera formicaria is not an easy task. The main problem is that the name “ Ch. formicaria ” has been incorrectly applied to almost all the other species of the genus for over 170 years (see Taxonomic history below). For example, Wasmann (1894: 133), Bernard (1968: 383) and Kistner (1982: 125), among many others, cited species of Messor as hosts of “ Ch. formicaria ”, which are erroneous due to the misidentification of the beetles. Rücker (1980: 143) goes even further and incorrectly associates “ Ch. formicaria ” with Messor barbarus, Aphaenogaster testaceopilosa (Lucas, 1849) and Pheidole megacephala (Fabricius, 1793).
From literature reports and our own examination of specimens associated with ants, we were able to recognise two ant taxa correctly associated with Ch. formicaria in its true geographic distribution: (1) a species of Tetramorium, based on the identification of two worker ants collected with the paratypes of Coluocera formicaria var. major in the Talish Region, Azerbaijan (see above), and (2) Messor structor (Latreille, 1798) reported by Arakelian & Kalashian (1993: 51) from Armenia.
Junior synonyms
Cholovocera subterranea Motchoulsky, 1845
Motchoulsky (1845: 111) described Ch. subterranea from “ Daghestan ”, distinguishing it from Ch. formicaria by being slightly smaller, darker and brighter. Gemminger & Harold (1868: 905) and Schaufuss (1876b: 413) accepted Ch. subterranea as a good species, without further comment. However, Reitter (1877: 5) found that the differences given by Motchoulsky (1845) did not justify the recognition of Ch. subterranea as a different, valid species. We agree with Reitter (1877) in regarding Ch. subterranea as a junior synonym of Ch. formicaria.
Wasmann (1894: 133), still using the name Ch. subterranea, made a doubtful association of this beetle with a species of the ant genus Aphaenogaster Mayr, 1853, without giving any details or reference. More recently, Shockley et al. (2009b: 65) and Rücker (2009: 14; 2020: 34) accepted Reitter’s (1877) synonymy but, without explanation, they erroneously name it as “ Merophysia subterranea Motschulsky, 1845 ”.
Coluocera formicaria var. major Reitter, 1887
Reitter’s (1887: 10) description of Coluocera formicaria major is very brief, only distinguishing it from the nominate subspecies by size. Leder (1886: 133) included this taxon in his catalogue, qualifying it as identical to Ch. formicaria, but larger. Heyden et al. (1883: 80) and Wasmann (1894: 133) included Co. formicaria major in their catalogues, indicating its location, but without an ant association. Escherich (1897) and Seidlitz (1898: 197) associated this subspecies with the ant Messor structor in his catalogue, but the correct identity of the beetles would have been Ch. balcanica. In his key to species of Cholovocera, Rücker (1980: 144, fig. 26) elevated this taxon to full species, but his interpretation of its type locality was erroneous, placing the Talish Region in “ Angora ”, which is actually in Turkey.
Arakelian & Kalashian (1993: 51) cited Co. f. major from Armenia, around the city of Noubarashen, reporting 10 to 15 individuals inside nests of Messor structor from March to June. We have not examined these beetles, which are deposited in the Entomology Museum of the National Academy of Sciences of Armenia in Yereban, Armenia (G. Arakelian & M. Kalashian pers. comm. 2020). Several checklists and catalogues (i.e., L̂bl & Smetana 2007: 557; Shockley et al. 2009b: 65; Rücker 2009: 14; 2011b; 2020: 34) still regarded Co. f. major as a full species and gave an erroneous geographical distribution, either by citing Turkey or by including several countries where this taxon does not occur, probably confusing it with the distribution of Ch. balcanica (see below).
Rücker (2018: 576, figs 1179–1180) described in detail what he believed to be “ Ch. major ”, but his figure of the aedeagus clearly shows that it was Ch. balcanica, associated with Messor barbarus and M. structor. Stalling (2019: 13) also misidentified material of Ch. balcanica from the Dodecanese Islands (Greece) as “ Ch. major ”, an error repeated by Lapeva-Gjonova & Rücker (2011: 6) and Lapeva-Gjonova (2013: 9) with specimens of Ch. balcanica from Bulgaria, associated with M. structor.
We have examined the holotype male and two paratypes (male and female) of Co. f. major, with type locality in Rasano (Talish Mountains), as well as over 20 males of Ch. formicaria. The body length of the holotype falls within the range measured for males of Ch. formicaria. Also, we have compared the morphology of the aedeagus of the holotype with those of Ch. formicaria. Although the distal tip of the median lobe of the aedeagus and the paramere are missing in the holotype (Fig. 24F–G), the remaining parts are identical to those of the many males of Ch. formicaria, which we have studied, also collected in the Talish Region (Fig. 15A, C). Furthermore, Rasano is located approximately 400 km south of Derbent, the type locality of Ch. formicaria. Therefore, we have no hesitation in placing Coluocera formicaria major as a new junior synonym of Ch. formicaria.
Erroneous synonymies
“ Cerylon lapidarium Dejean ”
This species has not been formally published, therefore it is not taxonomically available, i.e., it is a nomen nudum (A. Slipinski, pers. com. 2021). However, Belon (1879: 194) regarded it as a synonym of Ch. formicaria. Considering the morphology of species of Cerylon Latreille, 1802 (Cerylonidae), it is possible that the material studied by Belon (1879) belonged to a species of the genus Merophysia.
Merophysia ragusae Belon, 1895
Rücker (2020: 34) regards Merophysia ragusae as a junior synonym of Ch. formicaria. This is an error considering that Belon (1895) clearly places M. ragusae from Sicily close to Merophysia sicula Kiesenwetter, 1872, and not to any species of Cholovocera. Furthermore, Rücker (2011a: 18) had already placed Merophysia ragusae as a junior synonym of Merophysia formicaria Lucas, 1852. It is apparent that Rücker (2020: 34) confused Ch. formicaria with M. formicaria.
Taxonomic history and remarks
Although Victor’s (1838) description of Cholovocera formicaria is very brief, his illustrations allow an accurate identification of the taxon. However, as it will be discussed below, most of the records of Ch. formicaria published until recently proved to be incorrect, as well as geographic distributions and names of host ants given for Ch. formicaria in several checklists. Although there are many papers dealing with Ch. formicaria, there is a great deal of incorrect data repeated in them.
Märkel (1845: 255), while describing Ch. punctata, studied a specimen of Ch. formicaria from Russia, sent to him by Motschulsky (see Material examined, above). Lucas (1849: 553) reported “ Ch. formicaria ” from the margins of Lake Tonga (Algeria), on the border with Tunisia, but it was most likely Ch. punctata. Rosenhauer (1856: 355), while describing Ch. formiceticola, compared it with Ch. formicaria and Ch. punctata. Redtenbacher (1858: 380; 1874: 411) discussed the differences between Ch. formicaria and Ch. punctata, in particular their elytral punctuation. Fairmaire (1859: 267) and Dieck (1870a: 399; 1871: 202) recorded “ Ch. formicaria ” from Corsica, but these were misidentifications of Ch. punctata. Saulcy (1862: 291) reported “ Ch. formicaria ” from Banyuls (Southern France, associated with Messor capitatus (Latreille, 1798)) but the correct identity of that material is uncertain because there are four species which occur in Southern France: Ch. punctata, Ch. formiceticola, Ch. gallica and Ch. occulta sp. nov. In their catalogue, Gemminger & Harold (1868: 905) gave the distribution of Ch. formicaria in “Grusia” (Georgia). Piccioli (1871: 304) recorded “ Ch. formicaria ” from Florence and Corsica, and Bargagli (1872: 100) from Sardinia; however, these records are likely to be misidentifications of Ch. gallica or Ch. punctata.
Schaufuss (1876a:396, 400) commented about the morphology of Ch. formicaria and incorrectly regarded it as an endemic of Greece. André (1874: 226) associated Ch. formicaria with Messor barbarus, citing Saulcy (1862), who actually cited Messor capitatus (see above). Reitter (1877: 5) placed Ch. gallica as a junior synonym of Ch. formicaria, a status that we do not agree with. Belon (1879: 191) took a radical view, placing all five species of Cholovocera described until then, as junior synonyms of Ch. formicaria; however, Reitter (1882: 161) rejected such action claiming that Ch. punctata was a different species from Ch. formicaria because of its obvious punctuation, a position which was later accepted by Belon (1884a: 2; 1887: 216). Heyden et al. (1883: 80) incorrectly placed Ch. formicaria in southern Europe in their catalogue. Bolívar (1886: 51) recorded samples from Alicante and Algeciras (Spain), Morocco and Blidah (Algeria) as “ Ch. formicaria ”; we have studied a great part of this material which includes Ch. formiceticola (specimens from Spain and Morocco) and Ch. punctata (from Algeria). Similarly, Oertzen (1886: 201) recorded “ Ch. formicaria ” from Naxos (Greece), which were most likely Ch. balcanica or Ch. attae, Walker (1889: 377) reported it from Gibraltar and Tangier – actually Ch. formiceticola – and Gallois (1893: 18) cited it incorrectly from Nantes (France).
Wasmann (1894: 133) summarised published associations of what he regarded as “ Ch. formicaria ” with several species of ants: with Messor barbarus in eastern Pyrenees (from Saulcy 1862), in Gibraltar and Tangier (from Walker 1888) and in Tunisia (from Wasmann 1890); with a species of Pheidole in Andalusia, Spain (from Rosenhauer 1856), and with Aphaenogaster testaceopilosa in Menton, France (from Walker 1888). Even if the ant identities were correct, the beetles were certainly not Ch. formicaria.
Other reports of beetle samples misidentified as “ Ch. formicaria ” are:
Escherich & Emery (1897) cited it from “Brussa” (Bursa), associated with Messor structor, and from Angora (both localities in Anatolia, Turkey), but the correct identity of the beetles would have been Ch. balcanica; Escherich (1897) reported it from Anatolia again, but as “ Coluocera formicaria var. major ” and associated with Messor barbarus; Sahlberg (1903: 31) recorded a great number of specimens near Constantine (Algeria) associated with Messor barbarus, but we have examined material from this locality, which is Ch. punctata; Luigioni & Adelchi (1910: 69) recorded about 100 specimens under a rock in Lazio (Italia) in February, which we believe were Ch. punctata (see Material examined, below); Donisthorpe (1927: 9) collected specimens in Sicily, associated with “ Camponotus atlantis nylanderi Emery, 1921 ”, although this ant does occur in Sicily, the beetles were either Ch. gallica or Ch. punctata; Luigioni (1929: 528) reported material from Liguria, Toscana, Umbria, Lazio, Puglia, Calabria, Sardinia and Sicily (Italy), but the correct identity would have been Ch. gallica or Ch. punctata; Martínez de la Escalera (1914: 123) collected beetles from several localities in northern Morocco, which we have examined and identified as Ch. formiceticola; Angelini & Rücker (1999: 218) reported “ Ch. formicaria ” from Puglia, Calabria and Sicily, but it was most likely Ch. punctata or Ch. gallica, equally for Sabella & Sparacio’s (2004: 498) report from Sicily, and for the listed record in Fadda et al. (2007: 70) from Provence (France), which may also refer to Ch. occulta sp. nov. and/or Ch. formiceticola. Ponel (2011: 254) reported “ Ch. formicaria ” from Villeneuve à Fréjus, near Nice (France), but it may have been Ch. punctata, Ch. gallica or Ch. occulta sp. nov. Prieto-Manzanares (2018: 464) recorded “ Ch. formicaria ” in Barcelona (Spain), but it was either Ch. formiceticola or Ch. gallica. Parmentier et al. (2020: 589) reported “ Ch. formicaria ” from Córdoba (Spain) but, again, it was a misidentification of Ch. formiceticola.
Rücker (1980: 143) published a key for the identification of the six species which he recognised as belonging to Cholovocera, illustrating the median lobe of the aedeagus of all the species (Rücker 1980: 145, figs 21–26), including that of Ch. formicaria; however, the geographic distribution of this species is erroneous, and was repeated by Rücker (1983: 3).
Several recent checklists included Ch. formicaria, but repeated incorrect geographic distributions published previously (Rücker 2009: 14, 2008: 576, 2020: 34; Shockley et al. 2009b: 65). Finally, Rücker (2018: 576, figs 1183–1184) gave a detailed description of Ch. formicaria, including figures of the aedeagus; however, we believe these figures actually represent the aedeagus of Ch. punctata (Fig. 16).
description: Published as part of Delgado, Juan A. & Palma, Ricardo L., 2023, A revision of the genus Cholovocera Victor, 1838 (Coleoptera: Endomychidae), pp. 1-71 in European Journal of Taxonomy 906 on pages 20-25, DOI: 10.5852/ejt.2023.906.2329, http://zenodo.org/record/10424508