Cholovocera formiceticola ( Rosenhauer, 1856 ) 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 : 31-34

Cholovocera formiceticola ( Rosenhauer, 1856) new status

Figs 2 , 3C , 4F , 5–6 , 7F , 8A , 9A , 11B , 12F , 13E , 14J–K , 18 , 23F–G , 24E , 25D

Choluocera formiceticola Rosenhauer, 1856: 355 .

Cholovocera formiceticola – Gemminger & Harold 1868: 905.

Coluocera formcieticola – Schaufuss 1876a: 400.

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

Colovocera formicaria – Belon 1879: 191 (in part).

Cholovocera punctata – Shockley et al. 2009b: 65 (in part).

Differential diagnosis

Cholovocera formiceticola is morphologically and geographically close to Ch. gallica , but these species can be separated by the shape of the pronotum and of the metatibiae ( Fig. 14F–G against Fig. 14J–K ). Also, Ch. formiceticola is geographically close to Ch. punctata , but they can be distinguished by the shape of the terminal antennomeres ( Fig. 12F, H against Fig. 13E, G ).

Furthermore, the aedeagus and the paramere are useful characters to distinguish Cholovocera formiceticola from all other species in the genus ( Fig. 18 ).

Type material

Cholovocera formiceticola : the collections of Wilhelm D. Rosenhauer were sold by his family to several European institutions upon his death ( Katter 1881). The beetle collection, which was the base of his 1856 book Die Thiere Andalusiens, was mostly acquired by the Muséum national d’histoire naturelle in Paris, via R. Oberthür ( Cambefort 2006: 282; Bousquet 2016: 450). However, no specimen of Cholovocera from the Rosenhauer Collection could be found in that museum (A. Taghavian pers. comm. Dec. 2019). Alternatively, M. Balke found two male specimens from the Rosenhauer Collection in the SNSB (Berlin) labelled as Choluocera formiceticola and collected in “ Spanien ”. As we could not locate any more specimens from the type series, despite enquiries made to several European museums, we believe that these two beetles are the only extant syntypes of this species.

Considering the great number of misidentifications of the species of Cholovocera , both in collections and literature, it is advisable to designate a lectotype from the syntypes of Ch. formiceticola , to give this name taxonomic stability (Article 74.7.3, ICZN 1999). We hereby designate one syntype male from the Rosenhauer Collection deposited in SNSB, with label reading: “ Choluocera formiceticola Rosenh. Spanien.” as the lectotype of Cholovocera formiceticola ( Fig. 23F–G ). The other syntype male, without a label, becomes a paralectotype.

Lectotype

SPAIN • 1 ♂; “Spanien”; [Rosenhauer Collection], SNSB. Designated above.

Paralectotype

COUNTRY UNKNOWN • 1 ♂; without label; [ Rosenhauer Collection ], SNSB.

Additional material, non-types

FRANCE – Languedoc-Rousillon • 1 ♂, 13 ♀♀; “ Pyr. Or .” [eastern Pyrenees ], Banyuls ; Cl. Besuchet leg.; 25 Apr. 1953; “pierre avec fourmis” [stone with ants], two specimens with a Messor worker ant; MHNG • 2 ♀♀; “ Pyrenäen ” [ Pyrenees ], Collioure ; SFUN .

PORTUGAL – Lisbon • 2 ♂♂, 4 ♀♀, 8 specimens; Apr.–May 1910; A. Schatzmayr leg.; SDEI 10804– 10806 and 10829–10836 • 2 specimens; 1919; Schatzmayr leg.; NHMB • 1 specimen; 1910; SFUN • 1 specimen; “ Schakm.” leg.; SMTD • 2 specimens; 1910; MNHS – Evora • 1 specimen; 1910; SFUN • 3 specimens; 1910; MNHS .

SPAIN – Castilla and León • 1 ♂, 4 specimens; Avila, Candeleda ; 30 Mar. 1929; MCNM 198721– 198725 – Madrid • 7 specimens; Madrid; ZFMK • 15 specimens; Madrid; MCNM 198703, 198728, 198730–198731 • 2 specimens; Madrid; 1872; “En el retiro, bajo piedras, con hormigas” [ At the Retiro Park, under stones, with ants]; MCNM 198729 – Castilla and La Mancha • 7 specimens; Toledo, Malpica [Malpica de Tajo]; MCNM 198736 – Extremadura • 1 ♂, 1 ♀; Badajoz ; P. Seiler leg.; NHMB • 2 specimens; Badajoz; NHMB • 1 specimen; Badajoz ; Bleuse leg.; ZFMK • 2 specimens; Badajoz; ZFMK • 13 specimens; Badajoz ; MCNM 198726 – Murcia • 2 specimens; Cartagena ; MCNM 198720 • 1 specimen; Cartagena ; Simon leg.; NMPC • 1 ♂, 7 ♀♀ [four females in alcohol]; Murcia, La Alcayna, Rambla de las Monjas ; 38º05′26.3″ N, 1º10′10.7″ W; 7 Jun. 2018; J. Delgado leg.; CDUM • 1 ♀; Murcia, La Alcayna, Rambla de las Monjas ; 38º05′27.3″ N, 1º10′12.3″ W; 10 Apr. 2018; J.A. Delgado leg.; CDUM • 1 ♀ [in alcohol]; Murcia, La Alcayna, Rambla de las Monjas ; 38º05′27.3″ N, 1º10′12.3″ W; 31 May 2021; J.A. Delgado leg.; [at the entry of an ant nest]; CDUM • 9 ♀♀; Murcia, Molina de Segura, La Alcayna, Rambla de las Monjas ; 38º05′27.3″ N, 1º10′12.3″ W; 7 Jun 2018; J.A. Delgado leg.; [in ant nest]; CDUM – Andalusia • 2 specimens; Cadiz, Villaluenga del Rosario ; 2 May 1960; Cl. Besuchet leg.; MHNG • 2 ♀♀; Tarifa ; 22 Jan. 2005; Lebenbauer and Egger leg.; NHMW • 3 specimens; Cadiz, Los Barrios ; Cl. Besuchet leg.; 22 Apr. 1960; MHNG • 1 ♂, 3 specimens; Cadiz, Cortijo Salomón ; MCNM 198059 and 198063–198064 • 1 ♂, 6 specimens; Cadiz, San Roque, Cortijo Salomón ; 20 Jan. 1990; De Ferrer leg.; MCNM 198058 and 198060–198062 • 5 specimens; Cadiz, Algeciras ; Apr. 1901; Escalera leg.; MCNM 198709–198713 • 1 specimen; Cadiz, Algeciras ; Apr. 1901; Escalera leg.; ZFMK • 7 specimens; Cadiz, Algeciras ; MCNM 198693 and 198714–198717 • 1 specimen; Algeciras , Dieck leg.; MCNM 198732 • 3 specimens; Algeciras ; Arias leg.; MCNM 198733–198735 • 7 specimens; Algeciras; NHMW • 1 specimen; Algeciras; SFUN • 2 ♂♂, 3 ♀♀; Algeciras ; SDEI 10819 , 11924– 11926 and 11944 • 3 specimens; Algeciras; NHMB • 1 specimen; Srra [Range] Carbonera; MCNM 198719 • 18 specimens; Cordoba; SMTD • 2 ♂♂, 12 ♀♀; Cordoba; NMPC • 19 specimens; Malaga, Nerja ; 19–30 Apr. 1987; Scheuern leg.; “Macchia” [Machis], [each specimen associated with a Messo r worker ant]; ZFMK • 4 specimens; Gaucin ; Jan. 1970; “Fourmilière” [Ant nest]; MCNM 198738 • 1 ♀, 1 specimen; Colmenar ; 7 May 2017; Ernst leg.; NMPC • 1 ♂; Sierra Nevada, 10 km Southeast of Orgiva, “Bco.” [Gorge] de las Cuevas de Camacho ; 500 m a.s.l.; Scheuern leg.; ZFMK • 1 specimen; Granada; NHMB • 1 specimen; Mt. [Mount, most likely refers to Sierra Nevada] Granada; SMTD • 4 specimens; Sierra Nevada ; V. Heyden leg.; SFUN – Andalusia, no specific locality • 1 ♂, 1 ♀; NHMB • 3 specimens; ZFMK , • 5 specimens; SFUN – Balearic Islands • 1 ♂; Manacor ; MCNM 198718 • 1 ♀; Miramar ; MFNB – Melilla [Northern Africa] • 2 ♀♀, 1 specimen; Nov. 1908; Arias leg.; MCNM 197992–197994 – Spain, no specific locality • 2 specimens; “Hispania”; SFUN • 2 specimens; “Spanien”; SFUN • 4 specimens; “ Hispania ”; MFNB .

MOROCCO – Tangier-Tétouan • 1 ♂, 85 specimens; Tangier ; M. Escalera leg.; MCNM 197922– 197949 and 197950–197952 • 14 specimens; Tangier ; M. Escalera leg.; MCNM 197995–198006 and 198008 • 1 specimen; Tangier ; Sep. 1957; ZFMK • 4 specimens; Tangier ; E. Vaucher leg.; Sep. 1957; ZFMK • 1 specimen; Tangier , Herrmann leg.; NHMW • 3 specimens; Tangier; MHNG • 1 ♀; Tangier , 1899; NMPC • 1 ♀; Tangier; NMPC • 1 ♂, 2 ♀♀; Bani-Msuar [Beni Mesauar, near Tetouan]; M. Escalera leg.; MCNM 197921 • 1 ♂, 2 ♀♀; Cas. Blnc. [Casablanca]; NKME • 1 ♂, 2 ♀♀, 1 specimen; F. Taourirt; 20 Mar. 1923; E. Handschin leg.; B.? “ Ameisen ” [With ants]; NHMB – Fez-Meknes • 1 specimen; “ Maroc, prés de [near] Meknes ”; 24 Apr. 1961; Munard leg.; MHNG • 1 ♂, 3 ♀♀; Moyen Atlas, Azrou, Ito env.; 33º54.759′ E, 005º32.884′ W; 1440 m a.s.l.; 9 May 2009; Hlaváč, leg.; Under rocks on open meadow; NMPC • 2 ♀♀; Moyen Atlas, Azrou, Forêt de Cedres ; 33º43.490′ E, 5º18.418′ W; 1600 m a.s.l.; 9 May 2009; Hlaváč leg.; under stone in forest; NMPC – Souss-Massa • 1 ♂; Agadir ; 16–29 Jan. 1976; T. Palm leg.; “hos myror” [among ants]; MZLU 2020-006 – Rabat • 1 ♂; Rabat ; NHMB • 2 ♀ ♀; Ouimes [Oulmés], El Harcha “n̂rdl.” [north of]; H. Franz leg.; NHMW • 1 specimen; O.[ued] Beth entre Aïn-el Orma et Khemisset ; 24 May 1961; Munard leg.; MHNG – Marrakech-Safí • 1 ♀; Mogador ; Nov. 1905; Escalera leg.; MCNM 197991 – Laâyoune-Sakia El Hamra • 3 ♀♀; “ Algérie ” [error for Morocco], Tarfaïa; NHMB – Morocco, no specific locality • 1 ♂, 1 ♀; “ Marocco ”; SFUN • 1 ♂; “ Marocco ”; SDEI 10828 • 2 ♂♂, 1 ♀; “ Marokko ”; SDEI 10825–10827 • 2 specimens; “ Maroc ”; Gambey leg.; ZFMK • 6 specimens; “Maroc”; MHNG.

ALGERIA – Oran • 1 ♂, 2 ♀ ♀; Oran; SDEI 11911 and 10779–10780 • 2 ♀♀; Oran; SFUN – Algier • 1 ♂; Laverdure; C. De Barros leg.; SMTD .

Type locality

“Algeciras, Malaga, Estepona und Ronda” [Andalusia], Spain.

Description

Male as in Fig. 11B . Body length: 1.34 mm average, range 1.30–1.50 mm (N = 54, 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 rounded. Terminal antennomeres subtriangular, with round angles ( Figs 12F , 13E ). Metatibiae as in Fig. 14J–K . Prosternal process slightly keeled anteriorly, with a slight median constriction and triangular distally ( Fig. 4F ). Male last visible ventrite with a marked emargination and bordered by a brush of long setae.

Median lobe of aedeagus in ventral view, wide, subconical, short, with a pointed apex and slightly serrated on its right side ( Fig. 18A ). Aedeagus in lateral view as in Fig. 18C . Distal portion of paramere very short, wide and conical, with only two short subapical setae ( Fig. 18B, D ). Spermathecal duct short and reservoir c-shaped; ramus long and curved distally, cornu round and nodulus short and wide ( Fig. 7F ).

Geographic distribution

The known distribution of Cholovocera formiceticola is western Mediterranean, extending from southern France in the northeast to the Balearic Islands, and from continental Spain to Algeria, and Morocco in the south, reaching the Atlantic coast ( Fig. 9A ).

Host ants

Rosenhauer (1856: 355) associated Cholovocera formiceticola with “ Oecophthora pusilla Heer, 1852 ” (now Pheidole megacephala ) in several locations of Andalusia ( Spain). The presence of Ph. megacephala in Andalusia is doubtful; although myrmecologists cannot rule out the possibility that it was established in the Iberian Peninsula, it has not been found in the last 100 years despite intense search ( Espadaler & Collingwood 2001: 260). Most likely, the ant that Rosenhauer (1856: 355) referred to was Pheidole pallidula (Nylander, 1849) , an abundant species in southern Spain ( Martínez-Ibáñez & Espadaler-Gelabert 1986: 1026; Bernard 1968: 153).

Our examination of specimens of Ch. formiceticola showed two records, each associated with an unidentified species of Messor , in southern France and in Andalusia respectively.

Taxonomic history and remarks

Rosenhauer’s (1856: 355) original description of Cholovocera formiceticola is long and detailed, but he only dealt with external morphology and mouth-parts, which are not sufficient to reliably separate species within Cholovocera . In their catalogue, Gemminger & Harold (1868: 905) correctly cited Spain as the geographic distribution of Ch. formiceticola , but Piccioli (1871: 304) cited Sardinia in error, probably referring to Ch. punctata or Ch. gallica . Schaufuss (1876a: 400) gave the distribution of Ch. formiceticola as Spain, Balearic Islands, Corsica, Sardinia and Algeria; according to our data, Corsica and Sardinia are incorrect, most likely referring to Ch. punctata or Ch. gallica . Dieck (1870b: 173) reported specimens from Algeciras (Andalusia, Spain), and Verdiani-Bandi (1874: 149) from Tuscany ( Italy), but the latter is, again, a likely an error for Ch. gallica or Ch. punctata .

Reitter (1875: 301) discussed the external similarity of Ch. formiceticola with Ch. attae , but he synonymised both species under Ch. formicaria , a synonymy accepted by Heyden et al. (1883: 80) in their catalogue. Dieck (1888: 41) reported Ch. formiceticola from Tangier, a new locality, and Fauvel (1890: 338) resurrected it from the synonymy in his rectification of Heyden et al. (1883) catalogue. However, over a century later, Rücker (2009: 14) synonymised Ch. formiceticola under Ch. punctata in his checklist, a status which was accepted by Shockley et al. (2009b: 65) and Rücker (2020: 34).

As shown above, our examination of many specimens of Ch. formicaria , Ch. punctata and Ch. formiceticola , including type material, showed that Ch. formiceticola is a distinct species, which we herewith resurrect as a valid taxon.

Figures

Fig. 2. Mouth parts, antenna and distal metatarsomere of Cholovocera formiceticola (Rosenhauer, 1856). A. Labrum, dorsal view. B. Epypharynx, ventral view. C. Left antenna, ventral view. D. Left maxilla, ventral view. E. Left mandible, dorsal view. F. Right mandible, dorsal view. G. Labium, ventral view. H. Hypopharynx, ventral view. I. Distal metatarsomere, dorsolateral view. Scale bar = 0.1 mm.

Fig. 3. Head. A. Tentorium, ventral view. B. Tentorium, lateral view. C. Hind wing of Cholovocera formiceticola (Rosenhauer, 1856). D. Stylised aedeagus, ventral view. Abbreviations: see Material and methods. Scale bars = 0.1 mm.

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. 5. Terminal abdominal segments of Cholovocera formiceticola (Rosenhauer, 1856), ♀. A. Last visible ventrite, ventral view. B. Gonocoxites, paraprocts and proctiger, ventral view. C. Tergite 8, dorsal view. D. Sternite 8, ventral view. E. Proctiger, dorsal view. Abbreviations: see Material and methods. Scale bar = 0.1 mm.

Fig. 6. Terminal abdominal segments of Cholovocera formiceticola (Rosenhauer, 1856), ♂. A. Last visible ventrite, ventral view. B. Left hemitergite, dorsal view. C. Tergite 8, dorsal view. D. Spiculum gastrale, ventral view.E. Sternite 8, ventral view. F. Hemitergites and proctiger, dorsal view. Scale bar = 0.1 mm.

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. 8. Cholovocera Victor, 1838 feeding, parasites and commensals. A. Nest chamber of Messor barbarus (Linnaeus, 1767) with seeds and one specimen of Ch. formiceticola (Rosenhauer, 1856).B. Gut content: particulate material. C–D. Fungal spores. E. Hyphae. F. Arthropod remains. G. Parvomyces cf. merophysiae Santamaria, 1995 attached to a leg of Ch. formiceticola (Rosenhauer, 1856). H. Specimen of Parvomyces cf. merophysiae Santamaria, 1995. I. Malpighian tubules of Ch. formiceticola (Rosenhauer, 1856) with cyst of parasitic amoebae. J. Cysts enlarged.K. Rhizoglyphine mite, phoretic on Ch. punctata (Märkel, 1845), ventral view. L. Same mite, lateral view. Scale bars: A–F = 20 µm; G–L = 30 µm.

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. 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.

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. 18. Aedeagus and paramere of Cholovocera formiceticola (Rosenhauer, 1856). Lectotype (Spain). 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.

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.