Goniaderini Lacordaire, 1859, 1925

Aalbu, Rolf L., Kanda, Kojun, Merkl, Otto, Ivie, Michael A. & Johnston, M. Andrew, 2023, Reconstitution of some tribes and genera of Lagriinae (Coleoptera, Tenebrionidae), ZooKeys 1172, pp. 155-202 : 155

Tribe Goniaderini Lacordaire, 1859

Figs 21-28 , 29 , 30-41

Type genus.

Goniadera Perty, 1832.

Description.

Body length: 3-19 mm; stout to elongate, dorsoventrally flattened to having elytra strongly inflated, glabrous or setose. Most species are unicolored, some are bicolored (e.g., pronotum and elytra with different coloration) or have patterned elytra.

Head: Eyes reniform, anteriorly notched by canthus, rarely completely divided. Antennae moderately long, usually reaching past base of pronotum; antennomeres obconical to filiform.

Thorax: Pronotum shape variable, usually cordate, constricted at base, sometimes quadrate to rectangular. Lateral margins complete. Procoxae clearly separated by prosternal process. Mesocoxal cavity laterally closed, at least partially, by mesepimeron. Elytra striate or not. Metathoracic wings well developed (in all species examined by us). Legs slender, not fossorial, penultimate tarsomeres lobed or cupuliform.

Abdomen: Intersegmental membranes visible between sternites V-VII, abdominal hinging tenebrionoid. Defensive glands absent. Ovipositor either stout with four distinct gonocoxites and terminal gonocoxite digitate or greatly reduced with gonocoxites fused (e.g., Anaedus punctatus (Carter, 1914) see Matthews and Bouchard 2008).

Diagnosis.

Goniaderini can be distinguished from Lupropini and Prateini by having the mesocoxal cavities laterally open (i.e., laterally, at least partially closed by mesepimeron) and abdominal defensive glands absent.

In Lagriinae , this combination of characters is shared with Belopini Reitter, 1917, Chaerodini Doyen, Matthews & Lawrence, 1990, Eschatoporini , and Laenini (Fig. 1 ). Goniaderini can be distinguished from these tribes as follows.

In Belopini , abdominal hinging between sternites V-VII is medial (tentyrioid hinging), and no intersegmental membrane is visible between the sternites; the aedeagus is oriented so the tegmen is ventral, as in the majority of Pimeliinae ; penultimate tarsomeres are not lobed or cupuliform. Goniaderini has lateral abdominal hinging between sternites V-VII (tenebrionoid hinging), and the intersegmental membranes between these segments are visible; aedeagus is oriented so the tegmen is dorsal; penultimate tarsomere is either lobed or cupuliform.

Chaerodini contains just two genera found on sandy shores in Australia and New Zealand. They exhibit features typical of psammophiles, including having a globose body, fossorial protibiae, and shortened antennae. Chaerodini also has an antennal club composed of five antennomeres and very reduced ovipositors that lack apical gonostyli. Goniaderini is not globose, at most only the elytra are inflated; protibiae are not fossorial; and antennae extend past the anterior margin of the pronotum and are not clubbed. The ovipositor is shortened and reduced in some groups (e.g., Anaedus Blanchard, 1842), but gonostyli are always present.

Eschatoporini contains just one genus with two species restricted to Northern California. These species inhabit caves with natural water and are sometimes found at entrances to underground springs. The eyes are completely absent. Goniaderini possesses well-developed reniform eyes. Although Eschatoporini and Goniaderini both lack sternal defensive glands, the former possesses a pair of cuticular sac-like reservoirs between tergites VII and VIII. This character seems to be unique within Tenebrionidae , and their function is unknown ( Aalbu et al. 2017).

Most Laenini has small, rounded eyes that are not anteriorly notched by the epistomal canthus; body shape elongate, semi cylindrical but with strong constriction between thorax and abdomen making thorax rounded and abdomen elongate rounded; all species are apterous. Goniaderini has reniform eyes that are anteriorly notched by the epistomal canthus and although the body shape is highly variable, all examined species are winged.

Genera included.

Acropachia * Mäklin, 1875, Aemymone Bates, 1868, Anaedus Blanchard, 1842, Ancylopoma Pascoe, 1871, Goniadera Perty, 1832, Lyprochelyda Fairmaire, 1899, Microgoniadera * Pic, 1917a, Myrmecopeltoides Kaszab, 1973, Opatresthes Gebien, 1928, Phymatestes Pascoe, 1866, Spinolagriella Pic, 1955, and Xanthicles Champion, 1886.

Taxonomic changes among Goniaderini genera

Ferrer and Delatour (2007) revised the genera Goniadera and Microgoniadera , and placed both Aemymone and Opatresthes as subgenera of Goniadera mainly based on external surface characters. The characters listed in the former work to diagnose the tribe Goniaderini included mostly generalized lagriine or other variable characters. Their tribal concept also included Eschatoporis ( Eschatoporini , see Aalbu et al. 2017). No other genus was mentioned other than Microgoniadera , which was separated in their key as a distinct species based only on size. They did not consider Anaedus to belong to Goniaderini but rather to Lupropini .

Anaedus clearly belongs morphologically within Goniaderini , which is consistent with molecular analyses ( Aalbu et al. 2017). In fact, Aemymone (Fig. 19 ) is likely more closely related to Anaedus due to both possessing very elongate basal hind tarsomeres (not mentioned by Ferrer and Delatour 2007) as well as a lack of tubercles. Size is not reliable as certain species of Anaedus , like An. robusticollis (Pic, 1921), are larger than most Aemymone . Aemymone differs from Anaedus by (1) having clearly defined, punctate elytral striae, (2) lacking posterior pointing denticles on the lateral margin of elytra near the base, and (3) by having a slight metallic sheen in some species.

Ferrer and Delatour (2007) separated Goniadera and Opatresthes , as subgenera in their work, based upon the presence of setae (we find that both genera have setae), color of the integument (we find this character unreliable), and the sides of pronotum (we find this character reliable, although not adequately described in their key). Both Goniadera and Opatresthes , unlike Aemymone , have the basal tarsomere of the hind tarsi equal or subequal to the terminal tarsomere. These two genera can be further separated from each other by (1), the strongly explanate anterior two-thirds of the pronotum in Opatresthes (only at most slightly explanate sides of the pronotum in Goniadera ), (2) the lateral aspect of both the pronotum and elytra being strongly dentate/tuberculate in Opatresthes (lateral aspect at most with a few dentitions on the pronotum in Goniadera ), (3) the metaventrite is equal to or shorter than the first visible abdominal ventrite in Opatresthes (metaventrite longer than length of first abdominal ventrite in Goniadera ), and (4) general shape, Goniadera being narrower and more elongate than Opatresthes .

The reinstatement of Aemymone and Opatresthes is summarized in the following checklists. Note that many authorship and year attributions of Ferrer and Delatour (2007) were incorrect.

Figures

Figure 1. Phylogeny of Lagriinae from Aalbu et al. (2017) with revised generic names, tribal classification, and diagnostic morphological characters. Prev [ious]. Tribal Placement: COS = Cossyphini, BEL = Belopini, ADE = Adeliini, LAG = Lagriini, CHA = Chaerodini, LUP = Lupropini, GON = Goniaderini, PYC = Pycnocerini, LAE = Laenini. Defensive glands: location and number of abdominal defensive gland reservoirs: 8 / 9, paired = a pair of gland reservoirs with openings between abdominal sternites 8 and 9; 7 / 8, paired = a pair of gland reservoirs with openings between abdominal sternites 7 and 8; 7 / 8, single = single gland reservoir with opening between abdominal sternites VII and VIII. Mesocox [al]. Closure: Closed = lateral arms of meso- and metaventrites touching laterad of mesocoxa; Open = lateral arms not touching.

Figure 29. Dorsal habitus of holotype of Gamaxus hauxwellii Bates, 1868 [= Phymatestes brevicornis (Lacordaire, 1859)]. Scale bar: 2.5 mm.

Figures 21 - 28. Dorsal habitus of representatives of Goniaderini genera. 21) Aemymone cariosa Fairmaire 1873 22 Goniadera repanda (Fabricius, 1801) 23 Phymatestes spathifer Gebien, 1928 24 Spinolagriella sp. 25 Opatresthes quadrinodosus Gebien, 1928 26 Xanthicles caraboides Champion, 1886 27 Lyprochelyda sp. 28 Ancylopoma punctigera Pascoe, 1871, holotype. Scale bars: 1 mm (22, 26); 2 mm (19, 23 - 25); 5 mm (20, 21).

Figures 30 - 41. Dorsal habitus of species of Anaedus Blanchard, 1842 including species belonging to genera synonymized with Anaedus in this paper. 30 Anaedus punctatissimus Blanchard, 1842, type species of Anaedus 31 A. brunneus (Ziegler, 1844) 32 A. expansicollis Gebien, 1913, paratype 33 A. explanatus Pic, 1917 34 A. leleupi Ardoin, 1876, paratype 35 A. robusticollis Pic, 1921 36 A. notatus (Pic, 1923), syntype, type species of Microanaedus Pic, 1923 37 A. conradti (Gebien, 1921), originally described in Pseudanaedus Gebien, 1921 38 A. dilaticollis (Fairmaire, 1882), holotype, type species of Pseudolyprops Fairmaire, 1882 39 A. himalayicus (Kaszab, 1965), originally described in Spinolyprops Pic, 1917 40 A. serrimargo (Gebien, 1914), senior subjective synonym of Spinadaenus singularis Pic, 1921, the type species of Spinadaenus Pic, 1921. 41 A. nepalicus (Kaszab, 1975), originally described in Sphingocorse Gebien, 1921. Scale bars: 1 mm. Images lacking scale bars were produced by Otto Merkl and sizes of specimens were not recorded before he passed. Figs 36, 38 taken by Christophe Rivier (MNHN).

Figures 19 - 20. Head of two Lorelopsis species. 19 Lorelopsis sp. with clearly discernible supraorbital setae (SupS) 20 L. trapeziderus (Champion, 1913), a densely setose species in which the supraorbital setae are not discernible.