Gambrinus LeConte, 1853: 435

Etzler, Frank E., 2019, Generic Reclassification of Limonius Eschscholtz, 1829 (Elateridae: Dendrometrinae) sensu Candèze 1860 of the World, Zootaxa 4683 (3), pp. 301-335 : 309-313

2. Gambrinus LeConte, 1853: 435 , 58 species

Type species: Elater armus Say, 1839: 171 (= Elater stigma Herbst, 1806)), by monotypy; Nakane and Kishii 1955: 205; Lane 1971 (as subgenus of Limonius ): 21; Kishii 1987: 82; Ôhira 1990: 108; Han et al. 2012: 58 (and references therein).

Gambrinus as synonym of Limonius: Candèze 1860: 377 .

Limoniscus Reitter, 1905: 12 [14] new synonymy.

Type species: Elater violaceus Müller, 1821 (designated by Hyslop, 1921); Schwarz 1907: 195; Leseigneur 1972: 182; Dolin 1978: 43; Ôhira 1990: 108; Sanchez-Ruiz 1996: 107; Cate 2007: 165; Han et al. 2012: 58

Sichuanelater Platia and Gudenzi, 2006: 137 new synonymy.

Type species: Sichuanelater gibbosus Platia and Gudenzi, 2006 , by original designation; Platia 2008: 197.

The following are suggested species groups. I have included generic names in parentheses near the species groups when a type species is included in the group.

Nearctic species (29).

Stigma species-group ( Gambrinus ):

Gambrinus confusus ( LeConte, 1853) : 430 ( Limonius ) new combination.

Gambrinus crotchii (Horn, 1872) : 148 ( Limonius ) new combination.

Gambrinus flavomarginatus (Knull, 1938) : 20 ( Limonius ) new combination.

Gambrinus humidus (Lane, 1941) : 137 ( Limonius ) new combination.

Gambrinus lanchesteri (Lane, 1941) : 135 ( Limonius ) new combination.

Gambrinus meridianus (Knull, 1947) : 177 ( Limonius ) new combination.

Gambrinus mirus ( LeConte, 1853) : 429 ( Limonius ) new combination.

Gambrinus norahae ( Al Dhafer, 2009) : 312 ( Limonius ) new combination.

Gambrinus olentangyi (Knull, 1947) : 179 ( Limonius ) new combination.

Gambrinus rufihumeralis (Lane, 1941) : 133 ( Limonius ) new combination.

Gambrinus stigma (Herbst, 1806): 86 ( Elater ) new combination.

Gambrinus shircki (Lane, 1965) : 183 ( Limonius ) new combination, removed from synonymy.

Gambrinus snakensis (Lane, 1965) : 185 ( Limonius ) new combination, removed from synonymy. Gambrinus pictus ( Van Dyke, 1932) : 343 ( Limonius ) new combination.

Gambrinus ulkei (Horn, 1871) : 314 ( Limonius ) new combination, removed from synonymy.

Gambrinus venablesi (Wickham, 1913) : 27 ( Limonius ) new combination

Griseus species-group:

Gambrinus angulatus (Motschulsky, 1859) : 370 ( Limonius ) new combination

Gambrinus bicolor ( Van Dyke, 1932) : 344 ( Limonius ) new combination.

Gambrinus clypeatus (Motschulsky, 1859) : 369 ( Limonius ) new combination.

Gambrinus cribriceps (Van Dyke, 1943) : 44 ( Limonius ) new combination.

Gambrinus fulvipilis ( Candèze, 1860) : 383 ( Limonius ) new combination.

Gambrinus griseus (Beauvois, 1805) : 1805 ( Elater ) new combination.

Elater cylindriformis Say, 1823: 176 .

Limonius hirticollis Melsheimer, 1846: 215 .

Gambrinus interstitialis (Melsheimer, 1846) : 215 ( Limonius ) new combination, removed from synonymy. Gambrinus plebejus (Say, 1825) : 263 ( Limonius ) new combination.

Gambrinus propexus ( Candèze, 1860) : 386 new combination, removed from synonymy.

Gambrinus rudis (Brown, 1933) : 175 ( Limonius ) new combination.

Gambrinus seminudus ( Van Dyke, 1932) : 348 ( Limonius ) new combination.

Gambrinus sinuifrons (Fall, 1907) : 227 ( Limonius ) new combination.

Gambrinus ursinus ( Van Dyke, 1932) : 349 ( Limonius ) new combination.

Palearctic species (31).

Stigma species-group ( Limoniscus , Sichuanelater ):

Gambrinus ainu Ôhira, 1968: 9 .

Gambrinus amamiensis Ôhira, 1966: 10 .

Gambrinus atricolor (Lewis, 1879) : 157 ( Limonius ).

Gambrinus elegans (Buysson, 1891) : cxxxvii ( Limonius ) new combination.

Gambrinus gibbosus (Platia and Gudenzi, 2006) : 139 ( Sichuanelater ) new combination.

Gambrinus henanensis ( Schimmel, 2006) : 252 ( Limoniscus ) new combination.

Gambrinus hinakurai (Kishii, 1998) : 2 ( Limoniscus ) new combination.

Gambrinus hiramatsui Ôhira, 1976: 48 .

Gambrinus hosodai Kishii, 1989: 65 .

Gambrinus imitans (Lewis, 1894) : 196 ( Limonius ).

Gambrinus katoi (Kishii, 2002) : 129 ( Limoniscus ) new combination.

Gambrinus kawaharai (Kishii, 2002) : 129 ( Limoniscus ) new combination.

Gambrinus kraatzi kraatzi (Candèze, 1879) : 281 ( Limonius ).

Gambrinus kraatzi nihonicus Kishii, 1966: 46 .

Gambrinus kucerai ( Schimmel, 2006) : 253 ( Limoniscus ) new combination.

Gambrinus limbatipennis Nakane and Kishii, 1955: 43 .

Gambrinus naomii (Kishii, 1997) : 4 ( Limoniscus ) new combination.

Gambrinus nanshanensis (Arimoto and Hiramatsu, 2013) : 85 ( Limoniscus ) new combination.

Gambrinus niponensis (Lewis, 1894) : 194 ( Limonius ).

Gambrinus ogatai Kishii, 1985: 8 .

Gambrinus rufipennis (Lewis, 1894) : 197 ( Limonius ).

Gambrinus rufovittatus Ôhira, 1963: 15 .

Gambrinus shaanxiensis ( Schimmel, 2006) : 253 ( Limoniscus ) new combination.

Gambrinus suturalis (Gebler, 1844) : 99 ( Ampedus ) new combination.

Gambrinus takabai (Kishii, 1997) : 1 ( Limoniscus ) new combination.

Gambrinus violaceus (Müller, 1821) : 184 ( Elater ) new combination.

Gambrinus vittatus (Candèze, 1873) : 22 ( Limonius )

Gambrinus wittmeri (Chassain, 1998) : 120 ( Limoniscus ) new combination.

Gambrinus yamato (Kishii, 1998) : 5 ( Limoniscus ) new combination.

Gambrinus yujii ( Arimoto, 2013) : 79 ( Limoniscus ) new combination.

Gambrinus zhejiangensis ( Schimmel, 2015) : 290 ( Limoniscus ) new combination.

Diagnosis. Species of the genus Gambrinus are readily separated from other genera of the Limonius genus-group by the emarginate posterior margin of the hypomeron (Fig. 4). They are also united by similar female internal genitalia, with colleterial glands incorporated into uterus, bursa copulatrix bearing a narrow expansion, and a basal accessory gland on the bursa copulatrix (Figs. 5 and 6). Generally, species in this genus are larger than the other genera in the Limonius genus group, but exceptions are common. Other characters are quite variable within the genus, and are shared with species in Limonius .

Generic Description. Medium sized, mostly longer than 8mm, sub-cylindrical to cylindrical, lustrous or dull, quite variable in color. Head with simple or umbilicate punctures; frons depressed or rather flattened; frontal margin entire, straight or undulate anterad.Antennae with 2nd and 3rd antennomeres variable, often small, usually sub-equal in size, ratio of length of 2+3 to 4 th variable, often sub-equal, 4th to 10th antennomere serrate, 11 th ovoid, sometimes noticeably longer than others. Pronotum usually longer than wide, sometimes with anterior angles expanded; hind angles broad at base, often with single carina, rarely with carina weak or absent, with a nail-like projection on inner side; posterior margin lacking basal incisures and a basal notch; prosternal sutures double, often distinctly grooved anteriorly; posterior margin of hypomeron emarginate near apex; prosternal process expanded laterally; procoxal cavities narrowly opened. Scutellum subpentagonal, with or without longitudinal elevation. Abdomen with lateral microserrations on each ventrite.Legs slender; tarsomeres simple, often with dense setose pads ventrally; claws simple, lacking setae. Male aedeagus narrow with narrow sinuate parameres with membraneous apical end (Fig. 11, Stigma speciesgroup), or moderate with parameres more sclerotized with apical ends sinuate with distinct lateral expansions (Fig. 10, griseus species-group). Female internal genitalia with uterus bearing moderate sclerotization, colleterial glands incorporated into uterus, bursa copulatrix weakly sclerotized basally with long narrow expansion ending in finger-like glands and long tubular extension to spermatheca (Fig. 5, Stigma species-group), or completely weakly sclerotized, with short narrow expansion (Fig. 6, Griseus species-group); both types with basal accessory gland on bursa copulatrix.

Discussion. This definition encompases the species traditionally used with the generic name Limoniscus . While Limoniscus has been used consistently since it was described in 1905, its synonymy with Gambrinus requires a change in names. A reversal of precedence under Art. 23.9.1 of the International Code of Zoological Nomeclature (ICZN 1999) is precluded by the use of Gambrinus as a valid genus by Ôhira (1990), Han et al. (2012), and others within the last 50 years.

The relationship between these two names has been recognized since the 1950s (Ôhira 1954), with a direct comparison of type species done by Ôhira (1990). Ôhira moved the Japanese species to Limoniscus based on the lack of a carina on the posterior pronotal angles and the less serrated antennomeres present in G. stigma, stating that they were a better match to Limoniscus violaceus . However, close examination of many specimens of many species from both regions show that the strength of the hind angle carina is variable within the group, and the strength of serration of the antennomeres is species specific. I also disagree with Ôhira (1990) when he states that the antennae of G. stigma are weakly serrate (see Fig. 7) and there is a carina, although weaker than in G. violaceus , present on the hind angle (Fig. 8). Due to the variability of both characters, I can only recognize one genus.

The monotypic Sichuanelater is a highly derived member of this genus. While it has many unique characters, which led to the type species being described in its own genus, the male genitalia are typical for Gambrinus (Platia 1998, see his Fig. 11). Gambrinus gibbosus , from mainland China and type species of Sichuanelater , also appears similar to the Taiwanese G. nanshanensis further strengthening this synonymy.

This genus is defined by the emargination of the posterior edge of the hypomeron. In addition, these species share characters of both the male and female internal genitalia. All species can be linked by the female internal genitalia, a character that Becker (1956) noted was a strong character to discover natural groups of species.

Within Gambrinus , there appear to be two species groups, here designated the Stigma species-group and the Griseus species-group. Despite the differences in bursa copulatrix shape (Figs. 5 and 6), both species groups have colleterial glands as subtle expansions of the uterus, and bear accessory glands near the junction of the uterus and bursa copulatrix. Additionally, the male genitalia were used here to form species groups, matching those of the female internal genitalia. The Stigma species-group has aedeagi that possess narrow parameres (Fig. 11) and includes all Palearctic species and part of the Nearctic species. The Griseus species-group has relatively wide parameres (Fig. 10), and is limited to the remaining Nearctic species. Other general trends in the species groups are that the Stigma species-group contains species with completely piceous pronota, and elytra bearing orange humeral markings, while the Griseus species-group contains species with bicolored pronota and unicolorous brown elytra. Exceptions to these color patterns are not uncommon, so these can only be used as general trends and not definitions of the groups.

Larvae are only confidently associated for a few Palearctic species. Important works are Dolin (1978) and Whitehead (2003) for Western Eurasian species, and Ôhira (1962) for Japanese species. Known larvae are very close in appearance to Limonius , but the outer urogomphal prong is usually quite long relative to Limonius species (Fig. 9 in Whitehead 2003, cf. Fig. 72 here). While never specifically mentioned in keys, known species lack stemmata (Paul F. Whitehead, in litt.). In the Nearctic, only G. confusus has tentatively associated larvae ( Glen 1950). These larvae possess stemmata, differing from known Palearctic species, but were found in decayed wood and in well decayed leaf-litter ( Glen 1950), similar to the habitat of G. violaceus ( Whitehead 2003) . However, the examined larvae in Glen (1950) lack associated adults, so these observations should be used with caution.

This reorganization exposes a need for considerable work to be done at the species level, particularly in the Nearctic. Notable examples include the previously excluded G. angulatus ( Al Dhafer 2009) , the G. crotchi complex, and the G. griseus complex of species.