Oodes (Oodes) helopioides ( Fabricius, 1792 )

Guéorguiev, Borislav & Liang, Hongbin, 2020, Revision of the Palaearctic and Oriental representatives of Lachnocrepis LeConte and Oodes Bonelli (Coleoptera: Carabidae), with special account on Chinese species, Zootaxa 4850 (1), pp. 1-89 : 47-52

14. Oodes (Oodes) helopioides ( Fabricius, 1792)

( Figs 18 A–J, Figs 19 A–H, Table 4)

Carabus helopioides Fabricius, 1792: 155 (type locality: ‘Germania’).

= Oodes helopioides var. varians Letzner, 1851: 135 [unavailable name, cfr. Lorenz 1998: 305]. Placed in synonymy by Csiki (1931: 1009).

= Oodes parallelus Motschulsky, 1858: 172 (type locality: ‘Caucase’), syn. n. The name is junior primary homonym of both Oodes parallelus Say, 1830 and Oodes parallelus LaFerté-Sénectère, 1851 .

= Oodes parallelogrammus Motschulsky, 1858: 172 (type locality: ‘ France méridionale et l’Espagne’), syn. n.

= Oodes thessalonicensis Schatzmayr, 1909: 39 (type locality: ‘Bei Keretschkoi auf der Kalkidischen Halbinsel’ [= ΑσβεσΤΟΧώΡΙ, in Greek; Asvestohori, in English], Greece). Placed in synonymy by Lorenz (1998: 305).

= Oodes helopioides var. Fiorii Porta, 1923: 213 (type locality: ‘Paludi S. Anna’ [Province of Modena, Emilia–Romagna, Italy]). Placed in synonymy by Lorenz (1998: 305).

References.

Oodes helopioides: Dejean 1826: 378 (re-description); LaFerté-Sénectère 1851: 272; Chaudoir 1857: 34; Motschulsky 1858: 172–173 (comparison with O. desertus Motschulsky ); Gemminger & Harold 1868: 232; Bedel 1879: 54 (identification key); Bedel 1880: 160; Chaudoir 1882: 345 (taxonomic notes); Marseul 1882: 28; Ganglbauer 1891a: 383; Bedel 1897: 101; Reitter 1908: 186; Porta 1923: 213; Burmeister 1939: 165 (distribution and ecology); Jedlička 1931: 21 (comparison with Holosoma rambouseki ); Jeannel 1942: 981; Jeannel 1949: 828 (morphology), 833 (distribution); Basilewsky 1953: 161; Habu 1956: 84–85 (comparison with O. helopioides tokyoensis ); Fontolan 1959: 118–121 (re-description and comparison with O. gracilis ); Magistretti 1965: 256; Allen 1973: 32 (Arkansas Insect Collection); Lafer 1973: 848–849 (figures of diagnostic characters and distribution); Ishkov & Kabak 1995: 85; Lafer 1989: 204 (figures of diagnostic characters); Bousquet 1996: 450 (various data, incl. bibliography); 472 (relationships with O. amaroides ); Ortuño 1998: 5–6 (female genitalia); Ortuño et al. 2001: 191 (distribution in Iberian Peninsula); Salari Gougheri et al. 2014: 453; Machard 2019: 197.

Oodes parallelus Motschulsky : Motschulsky 1850: 63; Gemminger & Harold 1868: 232; Marseul 1880: 191; Chaudoir 1882: 346; Marseul 1882: 28.

Oodes holepioides (sic!): Motschulsky 1858: 172.

Oodes parallelogrammus: Gemminger & Harold 1868: 232 ; Marseul 1880: 190.

Oödes helopioides: Bates 1873: 254 (comparison with O. prolixus ).

Oodes (Oodes) helopioides: Ganglbauer 1891b: 51 ; Csiki 1906: 57; Jakobson 1906: 310 (distribution); Csiki 1931: 1008 (‘Europa, Marokko, Kirgisen-steppe, West-Sibi-rien’); Kryzhanovskij et al. 1995: 158 (distribution in ex-USSR).

Oodes (Oodes) gracilis? var. parallelus Motschulsky : Ganglbauer 1891b: 51; Csiki 1906: 57; Jakobson 1906: 310.

Oodes gracilis: Fiori 1903: 200 (‘paludi di S. Anna: nell’ Emilia’; see Porta 1923: 213, Note 1).

Oodes (Oodes) helopioides var. varians: Jakobson 1906: 310 .

Oodes thessalonicensis: Müller 1921: 139 (taxonomic notes and correction to Schatzmayr 1909).

Oodes helopioides var. thessalonicensis: Müller 1926: 137 ; Burmeister 1939: 165.

Oodes (Oodes) helopioides var. fiorii: Csiki 1931: 1009 .

Oodes (Oodes) helopioides var. thessalonicensis: Csiki 1931: 1009 .

Oodes helopioides var. fiorii: Burmeister 1939: 165 ; Magistretti 1965: 256.

Oodes helopioides helopioides: Lorenz 1998: 305 ; Bousquet 2003: 445; Lorenz 2005: 325; Bousquet 2017: 636.

Type material. Carabus helopioides Fabricius : Not examined.

Type material. Oodes parallelus Motschulsky : lectotype ♀, ‘Caucasus [r, h] // Oodes parallelus mihi Caucas [w, h] // Zool. Mus. Mosc. Univ. (Moscow, RUSSIA) ex coll. V.I. Motschulsky [w, p] // [red label without data]’ (ZMMU) ( Figs 18A, B ).

Having not studied the type of V.I. Motschulsky, Chaudoir (1882: 346) suggested that O. parallelus is identical to O. gracilis . Ganglbauer (1891b), Csiki (1906) and Jakobson (1906) placed the former as variation and questionable synonym of the latter. The synonymy of both names was validated by Csiki (1931). Study of the type specimen at hand showed that the ratio EL/EW is 1.36, and metepisternum is distinctly punctured. Comparisons with the available samples of O. helopioides and O. gracilis ascertained it is conspecific with the former, and not with the latter: Oodes parallelus Motschulsky, 1858 , syn. n. of Oodes helopioides ( Fabricius, 1792) .

There is no indication as to the number of specimens in the hands of Motschulsky for the original description of O. parallelus . Kelejnikova (1976: 210) mentions that Motschulsky’s collection contains a single syntype. This is why we find it appropriate to follow Recommendation 73F of the Code for avoidance of assumption of holotype ( ICZN 1999) and designated the specimen as lectotype.

Type material. Oodes parallelogrammus Motschulsky : lectotype ♀,‘Marseille[w, h] // Oodes paralellogrammus [sic] Motsch. Gall. mer. [w, h] // Zool. Mus. Mosc. Univ. (Moscow, RUSSIA) ex coll. V.I. Motschulsky [w, p] // [red label without data]’ (ZMMU) ( Figs 18C, D ).

Prior to this study, O. parallelogrammus was believed to be synonym of O. gracilis . Having not studied the type of V.I. Motschulsky, Chaudoir (1882: 346) supposed that both forms are identical. In the same year, their synonymy was formalized by Marseul (1882: 28), and this treatment was later accepted in the various catalogues of the Palaearctic Carabidae ( Ganglbauer 1891a, 1891b; Csiki 1906; Jakobson 1906; etc.). Study of the type specimen showed that it has: (1) ratio EL/EW: 1.46; (2) stria 7 at anterior third of elytron as developed as other medial striae; (3) granulation in elytral marginal furrow continuous; (4) metepisternum densely punctured, with lateral border slightly concave; and (5) metacoxal basal sulcus extended on lateral third. These characters demonstrate that it is conspecific with O. helopioides , and not with O. gracilis : Oodes parallelogrammus Motschulsky, 1858 , syn. n. of Oodes helopioides ( Fabricius, 1792) . It is worth noting that its EL/EW: 1.46 is at a border value for O. helopioides which overlaps with that of O. gracilis (see Table 4).

No data were found for number of studied specimens in the description of O. parallelogrammus . Chaudoir (1882: 346) stated that Motschulsky (1858) based his description on a single individual. On the contrary, Kelejnikova (1976: 210) noted that the specimen preserved in ZMMU is a syntype. Having no certainty on what source to trust, we followed Recommendation 73F of the Code ( ICZN 1999) and designated the specimen available as lectotype.

Type material. Oodes thessalonicensis Schatzmayr : lectotype ♂, ‘Keretschkoi (Makedonien) [w, p] // O. helopioides thessalonicensis Schatzm. [w, p] // Tipo [r, h]’ (MSNM). The sole specimen examined is conspecific with O. helopioides ( Figs 18E, F , 19A, B ).

The description of Schatzmayr (1909) does not contain designation for either holotype or type nor any hint of the number of specimens examined. Hence, as with the cases above, we followed Recommendation 73F of the Code ( ICZN 1999) and the specimen available is designated as lectotype.

Type material. Oodes helopioides var. fiorii Porta : consists of 1♂, 3♀♀ syntypes preserved in MSNM, all of them conspecific with each other and with O. helopioides . The four specimens are glued on cards and pinned two by two. The only male and one female specimen bear a label ‘Paludi S. Anna Modena. IV.1894 ex coll. A. Porta [w, h/p]’; the male specimen is chosen for lectotype ( Fig. 18G ), the female for paralectotype. The remaining two females, also designated paralectotypes, bear two labels, ‘Paludi di Sala Bologna— 12.3.1893 ex coll. A. Porta [w, h/p] // Oodes helopioides Fabr. Antonio Porta [w, h/p]’. Considering the two localities of var. fiorii, M. Pavesi (pers. comm.) stated that the two wetlands no longer exist.

Other material examined. FRANCE: I m p re c i s e localities: 1♀, ‘Donville 1–5–1900.’ ( IZAS). P ro v e n c e– A l p e s–C ô t e d ’ A z u r: 1♂, ‘les Collettes 30.6.86’ ( NMNHS).

BULGARIA: Silistra Province: 1♀, ‘BG—Srebarna 12/08/1989 N. Kodjabachev’ (IBER); 1♂, ‘BG— Srebarna 13/08/1991 N. Kodjabachev’ (IBER); 1♂, 1♀, ‘Srebarna Lake, wharf near the lake, 44.10331, 27.06392, 28–29.V.2017, G. Georgiev, W. Rossi & D. Stoianova’ ( NMNHS). Va r n a Province: 1♀, ‘ BULGARIA Kamchiya Outfall 3.VIII.2000 Picea sp. forest leg. Plamen Mitov’ ( NMNHS). Kyustendil Province: 1♀, ‘BG, Zemen gorge, 5.1986 G. Blagoev’ ( NMNHS). Burgas Province: 1♂, ‘BG, Kiten 16– 22.12.1984 P. Beron’ ( NMNHS); 3♂♂, ‘BG: Slānčev brjag b. Burgas Sűsswassertümpel (2) 07.V.2000 Handfang leg. Wolfg. Beier ( Germany)’ (cWB); 1♂, ‘BG, Strandza Mt., Veleka River outfall, 42.06100, 27.96533, 16.04.2009, longoz forest, leg. R. Kostova’ ( NMNHS) and 5♂♂, 4♀♀, same data but collected 15.04– 02.07.2009 ( NMNHS); 1♀, ‘BG, Strandza Mt., Malko Tarnovo district, Indipaskha, 42.00469, 27.65256, 18.04.2009, leg. R. Kostova’ ( NMNHS).

KAZAKHSTAN: West Kazakhstan Region: 1♀, ‘Kazakhstan NW Uralsk env. 4. 7. 91’ (cSF).

TME: 31 specimens. TGE: 2♂♂, 3♀♀.

Diagnosis. This species differs from the other Oodes species studied, except O. echigonus , by the lack of longer and denser, yellowish hairiness on the ventral surface of meso- and metatarsomeres 2–4 and rather wide elytra (EL/ EW ≤1.46, Table 4). It differs from O. echigonus in its small size, less than 10 mm.

Description. Habitus. Specimens of middle size (BL: 8.5–9.8 mm, BW: 3.5–4.0 mm), with ovate and moderately convex body ( Figs 18A, C, E, G, H ). Ratios and measurements. See Table 4. Color and luster. Body including appendages black with extremities of last palpomeres usually paler. Integument moderately shiny, without iridescence. Punctuation. Dorsal surface without punctuation; sides of prosternum and proepisternum as well as abdominal ventrites at sides rugose; mesosternum and abdominal ventrite 6 at apex punctate and rugose; sides of metasternum and metepisternum with coarse punctuation. Head. About half as wide as pronotum (see Table 4). Mentum tooth with distinct paramedial border ( Fig. 18I ). Thorax. Pronotum with sides slightly rounded toward posterior angles (PW/PB: 1.03–1.08); maximum width in posterior third; laterobasal impressions indistinct; base moderately sinuate; anterior angles rounded, not much projected anteriorly. Prosternum with median longitudinal sulcus very shallow; prosternal process bordered throughout, barely pointed at apex ( Fig. 18J ). Metepisternum slightly longer than wide (MA/MM: 0.93–0.97), with lateral margin concave posteriorly and coadunation with epipleuron very short, located anteriorly ( Fig. 18D ). Elytra. Apical sinuation weak, poorly defined. Basal margin distinct laterally, forming a small denticle at shoulder, disappearing medially at level of stria 3. Granulation in marginal furrow continuous. Parascutellar striola punctiform, other striae punctate anteriorly. Stria 7 as distinct as stria 6. Intervals 1–7 subconvex, interval 8 more convex than others. Legs. Metacoxal basal sulcus extending to lateral fifth ( Figs 18B, D ). Male mesotibia ( Fig. 18F ) moderately modified, with slight swelling in apical half. Protarsomeres 1– 3 of male moderately dilated, with second tarsomere slightly wider than long (W/Lp2: 1.06–1.10). Male genitalia. Median lobe ( Figs 19A, B, C, D, E, F ) with basal bulb long and wide; angle between basal bulb and shaft acute; shaft long and not swollen compared with basal bulb; apex short, tapered, and curved ventrally; apical lamella short, left-orientated, widely rounded; ostium long, reaching basal bulb. Female genitalia (see also Ortuño 1998). Basal gonocoxite with five lateroapical setae arranged in line. Apical gonocoxite without dorsolateral ensiform setae, with nematiform setae ( Fig. 18G ). Spermatheca coiled apically ( Fig. 18H ).

Distribution. Morocco, greater part of Europe (including the Caucasus Major), Turkey, Iran, Central Asia ( Kazakhstan, Turkmenistan, Tajikistan), western Siberia, in the east up to Abakan City, south-central Siberia ( Lafer 1973; Lindroth 1992; Ishkov & Kabak 1995; Kryzhanovskij et al. 1995; Bousquet 2017). The records for Turkmenistan and Tajikistan come from old sources and need confirmation, as well as information for Morocco (see Machard 2019). The record from North Iran (Salari Gougheri et al. 2014) is questionable too because no other reliable records for O. helopioides are known south of the line Greater Caucasus—North Kazakhstan.

Figures

FIGURE 18. Oodes helopioides (Fabricius, 1792) (A–B: Oodes parallelus Motschulsky, 1858, lectotype and its labels; C–D: Oodes parallelogrammus Motschulsky, 1858, lectotype and its labels; E–F: Oodes thessalonicensis Schatzmayr, 1909, lectotype and its labels; G: Oodes helopioides var. fiorii Porta, 1923, lectotype and its labels; H: male specimen, Bulgaria, Slānčev brjag; I: female specimen, Strandza Mt., Indipaskha; J: male specimen, France, les Collettes).A, C, E, G–H: habitus; B: left metacoxa; D: right metepisternum and metacoxa; F: left protarsomeres; I: mentum; J: prosternal process. Scale lines = 2 mm (Figs A, C, E, G–H); = 1 mm (Figs B, D); = 0.5 mm (Figs F, I–J).

FIGURE 19. Oodes helopioides (Fabricius, 1792) (A–B: Oodes thessalonicensis Schatzmayr, 1909, lectotype; C–D: male specimen, Bulgaria, Veleka River outfall; E–F: male specimen, Bulgaria, Srebarna Lake; G: female specimen, same locality; H: female specimen, same locality). A, E: median lobe of aedeagus, left lateral view; B–C: same, anterior dorsal view; D: same, posterior dorsal view; F: same, ventral view; G: left gonocoxite, ventral view; H: spermathecal complex and gonocoxites, ventral view. Scale lines = 0.5 mm (Figs A–F, H); = 0.2 mm (Fig. G).