Cholovocera punctata Märkel, 1845 (Markel, 1845)

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 : 25-31

Cholovocera punctata Märkel, 1845

Figs 4H , 7G , 9B , 11D , 12H , 13G , 14N–O , 16 , 17 , 23C–E , 24H–I , 25C, 25E

Cholovocera punctata Märkel, 1845: 255 .

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

Coluocera punctata sardoa – Reitter, 1911: 70. Syn. nov.

Colnocera [sic] punctata sardoa – Krausse 1913: 62.

Coluocera punctata ? n. sp.? – Krausse 1915: 120.

Cholovocera sardoa – Rücker 1980: 144, fig. 24.

Differential diagnosis

Cholovocera punctata can be distinguished from all other species in the genus by the shape of the terminal antennomeres, which are shaped like equilateral triangles ( Figs 12H , 13G ). Also, the aedeagus is characteristic ( Fig. 16 ), but similar to that of Ch. attae . However, in lateral view, they clearly differ in shape ( Figs 16C , 19C ), as well as in the shape of the parameres ( Figs 16B , 19B ).

Furthermore, the aedeagus, the paramere and the spermatheca are useful characters to distinguish Cholovocera punctata from all other species in the genus.

Type material

Cholovocera punctata : the type material comprises two syntypes, collected in Sicily by J.W. Helfer, which Märkel (1845) examined and compared with one specimen of Ch. formicaria sent to him by V.I. Motschulsky. In order to locate and recognise the syntypes of Ch. punctata , we had to examine specimens from the Märkel Collection deposited in SMTD, and from the Helfer Collection held in NMPC. A male and female pair was found in SMTD labelled as “ Sicilia – Coll. Märkel” ( Fig. 23E ) and mounted on card triangles ( Fig. 23C , left). Also, five females mounted on the same kind of mounting cards ( Fig. 23C , right) were located in NMPC, all of them with a single label, written by J.W. Helfer, reading “Monte Pellegrino, Palermo” ( Fig. 23D ). From the foregoing, we deduce that J.W. Helfer sent only two specimens to Märkel, which are the syntypes, keeping the others in his collection (see below). Märkel (1845) also examined a third specimen from the Germar Collection, which we were not able to locate.

Considering the great number of misidentifications of the species of Cholovocera , both in collections and in the literature, it is advisable to designate a lectotype from the syntypes of Ch. punctata , to give this name taxonomic stability (Article 74.7.3, ICZN 1999). We hereby designate the male syntype from the Märkel Collection deposited in SMTD, with label reading “ Sicilia – Coll. Märkel”, as the lectotype of Cholovocera punctata ( Fig. 23C , left).

Coluocera punctata sardoa : The type series comprises the lectotype male, five paralectotypes held in HNHM, and twenty-four paralectotypes in SDEI (labelled as “ syntypes ”), all collected in “Assuni”, Sardinia.

Lectotype (designated above)

ITALY – Sicily • 1 ♂, “ Sicilia / Coll. Märkel ”; SMTD.

Paralectotype

ITALY • 1 ♀; Sicily; SMTD.

Lectotype of Coluocera punctata sardoa (designated by Rucker 2011a: 13)

ITALY • 1 ♂; Sardinia, “ Assuni ”; Krausse leg.; HNHM.

Paralectotypes of Coluocera punctata sardoa

ITALY – Sardinia • 2 ♂♂, 3 ♀♀, 19 specimens; “ Assuni ”; Krausse leg.; SDEI: 11985–12004 and 12015–12018 [labelled as “syntypes”] • 5 specimens; “ Assuni ”; Krausse leg.; HNHM .

Notes

As it can be seen in Rücker 2011a (fig. 21), there is a label reading “ Holotypus ” attached to the specimen that Rucker 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 (1911).

The same comment given above under the lectotype, applies to the paralectotypes.

Additional material, non-types

FRANCE – Laguedoc -Roussillon • 1 ♂, 3 ♀ ♀; “Süd-Frankreich, Hérault”; NHMB – Provence-Côte D’Azur • 1 ♂, 7 specimens; Marseille; NHMB – Corsica • 5 specimens; “Umgeb.” [environs] Porto-Vecchio; 22– 25 Apr. 1927; SFUN • 4 specimens; Porto Vecchio; MHNG • 2 specimens; “Meerstrand b [near beach] Porto-Vecchio”; 22–25 Apr. 1927; MFNB • 2 specimens; “Umgeb.” [environs] Porto-Vecchio; 22–25 Apr. 1927; MFNB • 3 specimens; Ajaccio; A. Krausse leg.; SFUN • 1 specimen; Ajaccio; 6 Apr. 1911; A. Krausse leg.; NMPC • 1 ♀; Ajaccio; V. Budtz leg.; SMTD • 1 ♂; Ajaccio; SMTD • 2 specimens; Bastia; ZFMK • 9 specimens; Canvia; 24 Apr. 1973; S. Vit leg.; “ss. les pierres” [under stones]; MHNG • 4 specimens; “env.” [environs] Propiano; 30 Apr. 1973; S. Vit leg.; “ss. une pierre” [under a stone], [three specimens associated with a Pheidole worker ant, the fourth with a Messor worker ant]; MHNG • 2 specimens; Piana; 20 Apr. 1973; S. Vit leg.; “ss. une pierre” [under a stone], [one specimen associated with a Camponotus worker ant. The other one with a Lasius worker ant]; MHNG • 2 specimens; “env.” [environs] Sartène; 24 Apr. 1973; S. Vit leg.; “ss. une pierre” [under a stone], [each specimen associated with a Pheidole worker ant]; MHNG • 1 specimen; Ghisonaccia mer[idional]; 26 Apr. 1973; S. Vit leg.; MHNG • 1 specimen; Olmeto; 21 Apr. 1973; S. Vit leg.; “tronc enfoncé” [buried log]; MHNG – Corsica, no specific locality • 1 ♂, 1 specimen; SFUN • 2 ♂♂, 1 ♀, 5 specimens; ZFMK • 1 ♂; MCNM 198708 • 3 ♀♀; NMPC • 4 specimens; SMTD • 3 ♂♂, 7 ♀♀; SDEI 10776– 10777 , 10790–10801 , 10823 and 10844–10846 • 2 ♂♂, 2 ♀♀; Krausse leg.; NKME • 1 ♂; Miller leg.; NHMB • 1 ♂, 6 specimens; NHMB • 16 specimens; SMTD • 1 ♂, 3 ♀♀; MFNB – France, no specific locality • 1 ♂; “ Gallia Meridio ”; NKME • 2 ♀♀; “Frankreich”; SFUN • 1 specimen; “gallia”; MFNB • 1 ♀; “Gall. Mer.”; MFNB .

ITALY – Sardinia • 3 specimens; Asuni ; A.H. Krausse leg.; SMTD • 13 specimens; Asuni ; Krausse leg.; SDEI: 12019 to 12031 • 25 specimens; Asuni ; SDEI 11920–11922 , 12033–12044 and 11920– 11922 • 1 specimen; Asuni, Posta Senio ; Krausse leg.; MFNB • 2 specimens; Asuni ; NKME • 3 ♀♀, 8 specimens; Asuni; NMPC • 1 ♀; Asuni ; A.H. Krausse leg; NMPC • 1 ♂, 11 specimens; Asuni; SFUN • 2 specimens; Asuni b. [near] Cagliari ; 1916; J. Krasni leg.; SFUN • 47 specimens; Asuni ; SMTD • 1 ♂, 49 specimens; Asuni; NHMB • 20 specimens; Asuni ; Dr. Krausse leg.; NFNB • 1 specimen; Mount Turitas ; NMPC • 6 specimens; Oristano ; SMTD • 3 specimens; Oristano , NMPC • 1 ♀; Terranova [modern Olbia]; NMPC • 14 specimens; Sardinia; Cagliari; SMTD • 1 ♀; Golfo Aranci ; A. Dodero leg.; NHMB • 1 ♂, 1 ♀; 4 km Notrhwest of Telti ; 350 m a.s.l.; 12 Apr. 1992; J. Scheuern leg.; [one specimen associated with two Messor worker ants]; NKME • 2 specimens; Southwest of Macomer ; 500 m a.s.l.; 13 Apr. 1992; J. Scheuern leg.; [each specimen associated with a Messor worker ant]; NKME • 2 ♂♂, 1 ♀; Sardina, Lago dei Coghinas ; 160 m a.s.l.; 12 Apr. 1992; J. Scheuern leg.; [one specimen associated with a Messo r worker ant, other specimen associated with a Camponotus worker ant]; NKME • 24 specimens; Cagliari ; SMTD • 1 specimen; Cagliari ; A. Fiori leg.; MFNB • 1 specimen; Mount Sette Fratelli ; SMTD • 1 ♂, 8 specimens; Sorgono; NHMB • 1 ♀, 1 specimen; Sorgono ; SDEI 12032 and 11929 • 1 ♂, 5 specimens; Ozieri; NHMB • 2 specimens; Ozieri ; A. Dodero leg.; NHMB • 4 specimens; Sardina, Sassari, Macomer ; 13 Apr. 1952; SFUN • 2 ♂♂, 2 specimens; Maccomer ; 24 May 1920; MFNB • 1 specimen; Golfo Aranci ; A. Dodero leg; SFUN • 1 specimen; Golfo Aranci; SFUN • 1 specimen; Golfo Aranci ; 9 Apr. 1977; S. Vit leg.; “ss. une pierre” [under a stone], [specimen associated with a Camponotus worker ant]; MHNG • 1 ♂, 2 ♀♀, 1 specimen; Alá dei Sardi ; T. Derosas leg.; MCNM 197913–197916 • 1 ♂, 5 specimens; 12 km North of Dorcali ; 250 m a.s.l.; 23 Apr. 1992; J. Scheuern leg.; [two specimens associated with an Aphaenogaster worker ant]; ZFMK • 2 specimens; Noth of Pattada ; 450 m a.s.l.; 12 Apr. 1992; J. Scheuern leg.; [one specimen associated with four Pheidole worker ants]; ZFMK • 5 specimens; North of Pattada ; 300 m a.s.l.; 12 Apr. 1992; J. Scheuern leg.; [each specimen associated with a Messor worker ant]; ZFMK • 4 specimens; South of Pattada ; 300 m a.s.l.; 12 Apr. 1992; J. Scheuern leg.; [each specimen associated with a Messor worker ant]; NKME • 1 ♂, 2 specimens; North of San Vito ; 50 m a.s.l.; 18 Apr. 1992; J. Scheuern leg.; [specimens associated with three Pheidole worker ants]; ZFMK • 15 specimens; Telti, Olbia ; 9 Apr. 1977; S. Vit leg.; “ss. une pierre” [under a stone], [one specimen associated with a Camponotus worker ant]; MHNG • 2 specimens; “M”[ount]. Ferru ; A. Dodero leg.; MFNB • 1 specimen; Seui ; MFNB – Sardinia, no specific locality • 1 ♂, 17 specimens; J. Krausse leg.; NHMB • 4 specimens; SFUN • 2 ♀♀; Bruck leg.; SFUN • 1 specimen; ZFMK • 1 ♀; MCNM 198737 • 10 specimens; SDEI 12005–12014 • 1 specimen; Baudi leg.; SDEI 11938 • 1 specimen; SMTD • 1 ♀, 6 specimens; MFNB – Sicily • 3 specimens; Erica ; 10 Dec. 1993; Sabella leg.; “Bosco misto” [mixed forest]; NMPC • 6 specimens; Calabria, Sambiase ; May 1925; C. Minozzi leg.; [each specimen associated with a Messor worker ant]; SMTD • 1 specimen; Agricento, Valle dei Templi ; 18 Mar. 2013; P. Hlavác leg.; Under rock, ant nest [specimen associated with a Cataglyphis worker ant]; NMPC • 12 specimens; Ficuzza ; SDEI 12033–12044 • 2 ♂♂; Ficuzza ; NHMB • 1 ♂, 7 specimens; Ficuzza ; May 1895, Flach leg.; SFUN • 13 specimens; Ficuzza ; 14 May 1912; Fiori leg.; MHNG • 3 ♂♂, 1 ♀; Ficuzza ; 16 Mar 1942; NKME • 2 ♀♀; Ficuzza ; 16 Mar. 1942; MFNB • 1 ♂, 5 specimens; Palermo, Ficuzza ; 700–900 m a.s.l; 21–24 May 1996; F. Angelini leg.; MZLU • 1 ♂, 6 specimens; Ficuzza; NHMB • 21 specimens; Ficuzza ; 5 Apr. 1925; Dr. Rambousek leg.; NMPC • 3 ♀♀; Palermo, Ficuzza ; 700 m a.s.l.; 1–4 May 2000; “Bosco leccio” [oak forest], F. Angelini leg.; NMPC • 1 ♂, 6 specimens; Palermo, Marineo, L. [ake] Scanzano ; 525 m a.s.l.; 9 Apr. 1993; F. Angelini leg.; MCVR • 1 ♂, 5 specimens; Palermo, Madonie, Road Castelbuono–Gerace , Pitorno ; 30 Apr. 1996; Angelini leg.; MCVR • 1 ♂, 1 ♀, 1 specimen; Palermo, Lago di Piana degli Albanesi ; 610 m a.s.l.; 21 May 1996; F. Angelini leg.; MCVR • 5 ♀ ♀; “G.” [environs] Palermo, Monti Pellegrino ; 1852; Helger leg.; NMPC [Helfer Collection] • 1 ♂; Palermo ; NHMB • 2 ♂♂, 2 ♀♀, 1 specimen; Palermo; SDEI 11914–11919 • 1 ♂, 2 specimens; Palermo ; SDEI 11912–11915 • 1 ♂; Palermo ; E. Ragusa leg.; SFUN • 2 specimens; Palermo ; 1906; O. Leonhard leg.; SMTD • 1 ♂, 2 specimens; Randazza ; 6 May 1933; W. Liebmann leg.; SDEI 11972–11974 • 3 ♂♂, 2 ♀♀, 4 specimens; Catania; SDEI 11976–11984 • 5 specimens; Catania ; SDEI 05764–05768 • 11 specimens; Catania ; NMPC • 3 specimens; Catania ; Prof. Lera leg.; MFNB • 1 specimen; Catania ; MFNB • 1 ♂, 4 specimens; Catania ; SDEI 11976–11980 • 1 ♂, 18 specimens; Gibilmanna ; 800 m a.s.l.; 6 May 1982; T. Palm leg.; “Sten” [stone]; MZLU • 1 ♂, 2 specimens; Siracusa, Melilli ; 100 m a.s.l.; 5 Apr. 1997; F. Angelini leg.; MCVR • 1 ♀; Trapani, Mount S. Giuliano ; 24 Jan 1913; A. Dodero leg.; NHMB • 1 ♂, 2 specimens; Bertolia; SFUN • 4 specimens; Licata ; ZFMK • 2 specimens; “M.”[ount] Busambra ; 16 May 1912; A. Fiori leg.; MHNG • 2 ♀; Calabria, Cimina ; Paganeti leg.; NMPC – Sicily, no specific locality • 1 ♂; SDEI 11932 • 2 ♂♂; NHMB • 1 ♀; SFUN • 1 ♂, 1 ♀; Flach leg.; SFUN • 1 ♂, 1 ♀, 1 specimen; ZFMK • 1 ♂; “C. Rissen ” leg.; SDEI 11941 • 4 specimens; Sicily; SMTD • 1 specimen; 21 Nov. ; Füge leg.; SMTD • 1 specimen; NMPC – Tuscany • 1 ♂, 4 ♀♀; Mt. Argentario ; May 1907; Dr. Stolz leg.; NHMW – Lazio • 1 ♂, 3 ♀♀, 3 specimens; Mte. Circeo ; H. Franz leg.; NHMW • 1 specimen; Maccarese ; 20 Feb. 1910; Luigioni leg.; MFNB • 1 ♀; Roma ; 23 Apr. 1904; A. Fiori leg.; MFNB • 1 ♂, 1 ♀; Roma, Acilia; Castel. ; Nov. 1931; MFNB – Campania • 1 ♀; Salerno, Pioppi ; Nov. 1964; W. Liebmann leg.; MFNB – Calabria • 2 ♀ ♀; Alli; 24 Apr. [19?]84; MFNB .

MALTA – Northern Region • 2 ♀♀; Bidnija, NW Mosta; 1 Apr. 2002; Schuh and Mifsud leg.; NHMW .

ALGERIA – Oran • 1 ♂; Oran; NMPC – Algiers • 1 ♂; Quendef.; SDEI 10768 • 1 ♀; SDEI 11928 • 1 ♂, 10 specimens; Kabilie, Bou-Berak; L. Puel leg.¸ NHMB • 1 ♂, 1 ♀; St Charles; NHMB • 1 ♂, 1 ♀; Biskra; May 1898; L. Vareilles leg.; NHMB • 2 ♂♂, 1 ♀; Philippeville [now Skikda]; May 1898; L. Vareilles leg.; [each specimen associated with a Cataglyphis worker ant]; NHMB • 1 ♂, 1 ♀; Constantine; Le Thierry leg.; MCNM 198706 • 5 ♀♀; Constantine; Sep 1957; ZFMK • 4 ♀ ♀; Blida; 19 Apr. 1896; L. Bleuse leg.; “dans detritus où ils se trouvent des fourmis” [in an ant refuse pile]; ZFMK • 1 ♂; “ Bliad ” [Blida?]; MCNM 198704 • 1 ♂; Lambèze [now Tazoult-Lambése]; Jun. 1885; L. Bleuse leg.; ZFMK • 1 ♂, 1 ♀, 9 specimens; Bone [now Annaba]; ZFMK • 1 specimen; Gorges de la Chiffa, Ruisseau des Singes ; 280– 380 m a.s.l.; 4 May 1988; Besuchet, L ̂bl and Burckh. leg.; MHNG • 2 specimens; Djurdjura , 4 km Southwest of Tikjda; 1200 m a.s.l.; 7 May 1988; Besuchet, L ̂bl, Burckh. leg.; MHNG • 1 ♂, 2 ♀ ♀; G.[orge] de Kabylie, Djebel Bou-Berak ; 350 m a.s.l.; 19 May 1988; Besuchet, Löbl and Burckh. leg.; MHNG • 3 ♂, 7 specimens; G.[orge] de Kabylie, Adekar ; 900 m a.s.l.; 15 May 1988; Besuchet, L̂bl and Burckh. leg.; MHNG • 1 specimen; G. de Kabylie, L’Arbatache sur El Kseur ; 300–400 m a.s.l.; 18. May 1988; Besuchet, L̂bl and Burckh. leg.; MHNG • 4 specimens; G.[orge] de Kabylie, 9 km East of Yakouren ; 930 m a.s.l.; 13 May 1988; Besuchet, L̂bl and Burckh. leg.; MHNG • 3 ♀♀; MFNB • 1 ♂, 2 ♀♀, 33 specimens; Laverdure ; 30 Apr–14 May 1927; Mařan leg.; [six specimens associated with a Tetramorium worker ant, and an additional label: “ Tetramorium sp. ; P. Werner det. 2016”]; NMPC • 7 specimens; Batna; NMPC – Algeria , no specific locality • 1 ♂; Algier; NHMB • 1 ♂; Doufou leg.; MFNB .

TUNISIA – Bizerte • 2 ♀♀; Bordj Djedid; SFUN • 1 ♂, 1 ♀; Tabarka, Cap Negro ; 17 Mar. 1984; H. Meybohm leg.; NHMW • 1 specimen; Belif, “prés” [near] Cap Negro ; 5 Apr 1962; Cl. Besuchet leg.; MHNG • 1 ♂; Teskraia bei Bizerte ; 16 Mar. 1984; H. Meybohm leg.; NHMW • 2 ♂♂, 2 ♀♀; Galita I[sland]; H. Franz leg.; NHMW • 1 ♀; Monts de la Mejerda, Fôret Quedzen , ca. 20 km Southeast of Ain Draham; 26 Jan. 2004; Lebenbauer leg.; NHMW – Jendouba • 2 ♂♂; Umg. Ain Draham ; 18 Mar. 1984; H. Meybohm leg.; NHMW • 1 ♂, 13 specimens; Ain Draham ; 10 Mar. 1925; Dr. Rambousek leg.; NMPC • 3 specimens; Ain Draham ; 16–18. Apr. 1927; Mařan leg.; NMPC • 1 ♀; Ain Drahan ; B. v. Bodemeyer leg.; SDEI 11975 – Tunis • 1 specimen; Souk El Arba; NMPC • 1 ♀; Dijedid ; NMPC • 3 specimens; Dr Normand leg.; NMPC • 1 specimen; Belvedere ; 3 Mar. 1962; Cl. Besuchet leg.; MHNG • 1 specimen; Oued Mitiane près de Tunis; 26 Mar. 1962; Cl. Besuchet leg.; MHNG • 3 ♂♂; Hanman-El-Def [for Hamman-Lif]; 10. Jul. 1914; CNHM – Kef • 1 specimen; Le kef; Dr Normand leg.; NMPC – Kairouan • 1 ♂, 7 specimens; Ain Jioula; 26 Mar. 1984; leg. H. Meybohm; NHMW – Tunisia, no specific locality • 1 ♂, 1 ♀; “Tunis”; 8 Nov. [18?]85; ZFMK • 1 specimen; “Tunis”; NMPC .

Type locality

“in Sicilien ” [Sicily], Italy.

Description

Male as in Fig. 11D . Body length: 1.40 mm average, range 1.30–1.50 mm (N = 49, males and females). Shape of body subelliptical, pronotum almost rectangular, with angular lateral margins. Elytral apex rounded. Terminal antennomeres triangular, equilateral, in both sexes. Metatibiae as in Fig. 14N, O , narrower in the proximal half and with almost straight margins. Prosternal process slightly keeled anteriorly, with a marked median constriction and distally rhomboidal ( Fig. 4H ). Male last visible ventrite with a marked emargination and bordered by a brush of long setae.

Median lobe of aedeagus short, subrectangular in the first third of its length and distally oblique, tapering with a round apex in ventral view ( Fig. 16A ). Aedeagus in lateral view as in Fig. 16C . Distal portion of paramere long, conical, with a pointed apex bearing a brush of setae of variable number ( Fig. 16B–C ). The parameral apex and the number of apical setae show a clinal variation in shape and number respectively ( Fig. 17A–F ). Specimens from North Africa and central Italy have a parallelsided, rounded apex with five or six setae ( Fig. 17A–B ). The apex gradually tapers, becoming more triangular and pointed, with a decreasing number of setae in more northern European populations ( Fig. 17C–F ). Both the spermathecal duct and the spermathecal reservoir are short; ramus short, round and globose distally, cornu round, and nodulus weakly developed, much smaller than cornu and ramus together ( Fig. 7G ).

Geographic distribution

The known distribution of Cholovocera punctata is typically Mediterranean, extending from southern France in the northwest, to Corsica, Sardinia, the west coast of continental Italy and Sicily in the east, and Algeria and Tunisia in the south ( Fig. 9B ).

Host ants

Sahlberg (1903: 31) reported specimens of Ch. punctata associated with Messor barbarus in Lake El Bahira ( Algeria) and in Hamman-Lif ( Tunisia); considering that we have examined three males of Ch. punctata from Hamman-Lif (see above), we consider this association to be most likely correct. Confirmation of that association can be found in Krausse (1911), who found Ch. punctata (as Co. punctata sardoa ) associated with Messor barbarus in Sardinia, and both species do occur in this island. Rücker (2018) associated Ch. punctata with the ant genera Messor and Atta ; however, the genus Atta is exclusively Neotropical in distribution, making it impossible to be associated with a beetle of Mediterranean distribution!

Besides its association with Messor barbarus , we were able to recognise another six ant genera associated with Ch. punctata in its geographic distribution (see Material examined above), as follows: (1) Pheidole in Corsica and Sardinia, (2) Camponotus in southern France and Sardinia, (3) Cataglyphis in Sardinia and Algeria, (4) Lasius in Corsica, (5) Aphaenogaster in Sardinia, and (6) Tetramorium in Algeria. Among all the species of Cholovocera , Ch. punctata has the greatest range of associations with ant taxa, especially in Sardinia, where it has been recorded with species of five different ant genera.

Junior synonym

Coluocera punctata sardoa Reitter, 1911

Reitter (1911: 70) described Co. punctata sardoa from specimens collected in “Assuni”, Sardinia, provided by A.H. Krausse, distinguishing it from nominate Ch. punctata by external features, such as being slightly longer, more punctured and lacking two dark spots on the base of the pronotum. Krausse (1911: 148, 1913: 62, 1915: 120, 1917: 51) recorded Co. p. sardoa in Sardinia, adding Sorgono as a new locality for it. Seidlitz (1912: 67) included Co. p. sardoa in his list of Coleoptera described until 1911.

Rücker (1980: 144) raised this taxon to full species, including it in an identification key, with a figure of the median lobe of the aedeagus; however, Rücker’s (1980: 145, figs 24–25) illustrations of the median lobes of Ch. sardoa and Ch. punctata appear different because they are presented in different orientations. D̂bler (1987: 16) reported the presence of 24 “ syntypes ” (in fact, paratypes) de Co. p. sardoa in the collection of the SDEI. Subsequent catalogues and checklists include Ch. sardoa as a valid taxon and endemic to Sardinia (L̂bl & Smetana 2007: 557; Rücker 2009: 14, 2011b, 2018: 579, 2020: 34; Shockley et al. 2009b: 65).

We have examined the holotype male of Co. punctata sardoa and compared it with many males of Ch. punctata from the type locality, without finding any significant morphological difference that would justify the separation of this species into two subspecies. The terminal antennomere of the holotype is identical to that of Ch. punctata , and its aedeagus, although partially broken in the basal piece ( Fig. 24H– I ), is otherwise like those of several males of Ch. punctata , which we have studied ( Fig. 16 ). Therefore, we have no hesitation in placing Coluocera punctata sardoa as a new junior synonym of Ch. punctata .

Taxonomic history and remarks

Märkel’s (1845: 255) description of Cholovocera punctata is in Latin and brief, qualifying it as “elongate, oval, convex, reddish-yellow, punctured, with elytral apex attenuated, and habitat in Sicily”, but he emphasized the main difference from Ch. formicaria being the abundance of punctures on the entire surface of the body; also, Märkel (1845) implied a possible association with ants but could not confirm it. Erichson (1845: 125) and Ragusa (1873: 176) cited Ch. punctata in Sicily again; Redtenbacher (1858: 380, 1874: 411) mentioned its punctation as the main difference with Ch. formicaria , and Fairmaire & Coquerel (1860: 170) published an illustration of the mouth parts of Ch. punctata . In their catalogue, Gemminger & Harold (1868: 905) mentioned Sicily as the distribution of Ch. punctata . Piccioli (1871: 304) and Bargagli (1872: 100) reported Ch. punctata from Sardinia for the first time. Schaufuss (1876a: 396) commented on the morphology of Ch. punctata ; Reitter (1875: 302) gave a brief description of Ch. punctata , and later sank Ch. formiceticola as a junior synonym of it ( Reitter (1877: 5). Belon (1879: 191) proposed to sink Ch. punctata under Ch. formicaria , but Reitter (1882: 161) disagreed with Belon (1879) alluding to the denser punctuation of Ch. punctata . Then, Belon (1884a: 2, 1887: 216) gave other characters to separate Ch. punctata from Ch. formicaria . The catalogue of European beetles by Heyden et al. (1883: 80) gave the distribution of Ch. punctata as Sicily and Sardinia only, but Riggio (1885: 31) added a new locality for this species in the Island of Ustica, 100 km north of Sicily, and Heyden (1886: 38) recorded it in Algeria. Ciofalo (1886: 188) recorded it in Termini Imerese, northern Sicily. However, the catalogue published by Wasmann (1894: 133) only mentioned Sicily as the distribution of Ch. punctata .

Ganglbauer (1899: 821) and Belon (1902: 5) published redescriptions of Ch. punctata and its distribution, while Vitale (1904: 75) commented that this beetle is very rare in Messina (East Sicily). Luigioni & Adelchi (1910: 70, 1913: 152) recorded a few specimens in moist soil under Eucalyptus trees near Rome and others in Sicily, respectively. In his catalogue of Italian Coleoptera, Luigioni (1929: 528) mentioned Liguria, Toscana, Capri Island, Lazio, Campania, Calabria and Sicily, as localities for Ch. punctata ; we have examined specimens from Lazio, Calabria and Sicily which confirm Luigioni’s (1929) records.

Martínez de la Escalera (1914: 123) reported “ Ch. punctata ” from several localities in northern Morocco, but the correct identification of the beetles would have been Ch. formiceticola . In his catalogue, Sainte-Claire Deville (1914: 255) suggested that Ch. punctata occurs in western Mediterranean islands, while Ch. formicaria lives in southern France, but the latter statement is incorrect. Rücker (1980: 144) included Ch. punctata in his key for the identification of Cholovocera species, illustrating the median lobe of the aedeagus ( Rücker 1980: 145, fig. 25); also, he gave the geographic distribution of this species as: France, Spain, Corsica, Sardinia, Sicily, Algeria and Tunisia, which was repeated by Rücker (1983: 4). We agree with that distribution except for Spain, where we have not found any record of Ch. punctata ( Fig. 9B ). Angelini & Rücker (1999: 218) reported Ch. punctata from Basilicata (southern Italy) and Sicily.

Catalogues and checklists which include Ch. punctata have been published by L̂bl & Smetana (2007: 557), Rücker (2009: 14), Shockley et al. (2009b: 65) and Rücker (2011b, 2018: 576, 2020: 34); all these papers gave the geographic distribution as: southern France, Corsica, Italy, Sardinia, Sicily, Hungary, Spain, Switzerland, Algeria and Tunisia. Our examination of many specimens shows that these localities are mostly correct, except for Spain, Hungary and Switzerland. Records from Spain are most likely of Ch. formiceticola (see below), but those from Hungary and Switzerland are not supported by specimens; therefore, they need to be confirmed. Rücker (2018: 576, figs 1181–1182) gave a detailed description of Ch. punctata , including a figure of the aedeagus; however, we believe these figures actually represent the aedeagus of Ch. formicaria ( Fig. 15 ).

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. 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. 16. Aedeagus and paramere of Cholovocera punctata (Märkel, 1845). Lectotype (Sicily). A– B. Ventral views. C. Lateral view. Scale bar = 0.1 mm.

Fig. 17. Distal portions of parameres of Cholovocera punctata (Märkel, 1845) from various localities. A. Philippeville (Algeria). B. Monte Argentario (Central Italy). C. Palermo (Sicily). D. Corsica. E. Herault (Southern France). F. Marseille (Southern France).

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.

Fig. 19. Aedeagus and paramere of Cholovocera attae (Kraatz, 1858) (Greece). A–B. Ventral views. C. Lateral view.. Scale bar = 0.1 mm.

Fig. 15. Aedeagus and paramere of Cholovocera formicaria Victor, 1838. Male from type locality (Derbent). A–B. Ventral views. C–D. Lateral views. Scale bar = 0.1mm.