Research Article |
Corresponding author: Kojun Kanda ( grabulax@gmail.com ) Academic editor: Warren Steiner
© 2023 Rolf L. Aalbu, Kojun Kanda, Ottó Merkl, Michael A. Ivie, M. Andrew Johnston.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Aalbu RL, Kanda K, Merkl O, Ivie MA, Johnston MA (2023) Reconstitution of some tribes and genera of Lagriinae (Coleoptera, Tenebrionidae). ZooKeys 1172: 155-202. https://doi.org/10.3897/zookeys.1172.103149
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The tribes Goniaderini Lacordaire, 1859 and Lupropini Lesne, 1926 within the tenebrionid subfamily Lagriinae Latreille, 1825 have previously been shown to be non-monophyletic by molecular phylogenetic analyses. The tribes and constituent genera are here reviewed and redefined morphologically. As part of tribal redefinitions, we establish Prateini New Tribe with type genus Prateus LeConte, 1862. We reestablish the subtribe Phobeliina Ardoin, 1961 Revised Status, which is transferred from Goniaderini and placed as a subtribe of Lagriini Latreille, 1825 where it is comprised of Phobelius Blanchard, 1842, and Rhosaces Champion, 1889 (previously in Lagriini: Statirina Blanchard, 1845). The fossil tribe Archaeolupropini Nabozhenko, Perkovsky, & Nazarenko, 2023 is transferred from Lagriinae to Tetratomidae: Tetratominae Billberg, 1820. Keys to extant tribes and subtribes of Lagriinae and genera of Goniaderini, Lupropini, and Prateini are provided. Generic and species-level changes from this work are as follows:
Prateini is comprised of the following 15 genera: Antennoluprops Schawaller, 2007, Ardoiniellus Schawaller, 2013, Bolitrium Gebien, 1914, Enicmosoma Gebien, 1922, Indenicmosoma Ardoin, 1964, Iscanus Fauvel, 1904, Kuschelus Kaszab, 1982, Lorelopsis Champion, 1896, Mesotretis Bates, 1872, Microcalcar Pic, 1925, Micropedinus Lewis, 1894, Paratenetus Spinola, 1845, Prateus, Terametus Motschulsky, 1869, and Tithassa Pascoe, 1860. Lorelus Sharp, 1876 is Returned to Synonymy with Prateus, resulting in the following 49 New Combinations: Prateus angulatus (Doyen & Poinar, 1994), P. angustulus (Champion, 1913), P. armatus (Montrouzier, 1860), P. biroi (Kaszab, 1956), P. blairi (Kaszab, 1955), P. brevicornis (Champion, 1896), P. breviusculus (Champion, 1913), P. caledonicus (Kaszab, 1982), P. carolinensis (Blair, 1940), P. chinensis (Kaszab, 1940), P. clarkei (Kulzer, 1957), P. crassicornis (Broun, 1880), P. crassepunctatus (Kaszab, 1982), P. cribricollis (Kaszab, 1940), P. curvipes (Champion, 1913), P. dybasi (Kulzer, 1957), P. fijianus (Kaszab, 1982), P. fumatus (Lea, 1929), P. glabriventris (Kaszab, 1982), P. greensladei (Kaszab, 1982), P. guadeloupensis (Kaszab, 1940), P. hirtus (Kaszab, 1982), P. ivoirensis (Ardoin, 1969), P. kanak (Kaszab, 1986), P. kaszabi (Watt, 1992), P. laticornis (Watt, 1992), P. latulus (Broun, 1910), P. longicornis (Kaszab, 1982), P. mareensis (Kaszab, 1982), P. marginalis (Broun, 1910), P. niger (Kaszab, 1982), P. norfolkianus (Kaszab, 1982), P. obtusus (Watt, 1992), P. ocularis (Fauvel, 1904), P. opacus (Watt, 1992), P. palauensis (Kulzer, 1957), P. politus (Watt, 1992), P. priscus (Sharp, 1876), P. prosternalis (Kaszab, 1982), P. pubescens (Broun, 1880), P. pubipennis (Lea, 1929), P. punctatus (Watt, 1992), P. quadricollis (Broun, 1886), P. queenslandicus (Kaszab, 1986), P. rugifrons (Champion, 1913), P. solomonis (Kaszab, 1982), P. tarsalis (Broun, 1910), P. unicornis (Kaszab, 1982), and P. watti (Kaszab, 1982). Microlyprops Kaszab, 1939 is placed as a New Synonym of Micropedinus resulting in the following New Combinations: Micropedinus ceylonicus (Kaszab, 1939) and M. maderi (Kaszab, 1940). Lorelopsis Revised Status is revalidated as a genus and eight species formerly in Lorelus are transferred to it resulting in the following six New Combinations: Lorelopsis bicolor (Doyen, 1993), L. glabrata (Doyen, 1993), L. exilis (Champion, 1913), L. foraminosa (Doyen & Poinar, 1994), L. minutulis (Doyen & Poinar, 1994), L. trapezidera (Champion, 1913), and L. wolcotti (Doyen, 1993). Lorelopsis pilosa Champion, 1896 becomes a Restored Combination.
In Goniaderini, Aemymone Bates, 1868 Revised Status and Opatresthes Gebien, 1928 Revised Status, which were recently considered as subgenera of Goniadera Perty, 1832, are restored as valid genera based on new character analysis resulting in the following New Combinations: Aemymone hansfranzi (Ferrer & Delatour, 2007), A. simplex (Fairmaire, 1889), A. striatipennis (Pic, 1934) and Restored Combinations: Aemymone cariosa (Bates, 1868), A. crenata Champion, 1893, and A. semirufa Pic, 1917. Gamaxus Bates, 1868 is Returned to Synonymy with Phymatestes Pascoe, 1866, and the type species Gamaxus hauxwelli Bates, 1868 is placed as a New Synonym of Phymatestes brevicornis (Lacordaire, 1859). The following seven genera are placed as New Synonyms of Anaedus Blanchard, 1842: Microanaedus Pic, 1923, Pengaleganus Pic, 1917, Pseudanaedus Gebien, 1921, Pseudolyprops Fairmaire, 1882, Spinolyprops Pic, 1917, Spinadaenus Pic, 1921, and Sphingocorse Gebien, 1921. Fourteen species described by Pic in Aspisoma Duponchel & Chevrolat, 1841 (not Aspisoma Laporte, 1833) are returned to Tenebrionidae as valid species of Anaedus. These synonymies necessitate the following 51 New Combinations: Anaedus albipes (Gebien, 1921), A. amboinensis (Kaszab, 1964), A. amplicollis (Fairmaire, 1896), A. anaedoides (Gebien, 1921), A. angulicollis (Gebien, 1921), A. angustatus (Pic, 1921), A. australiae (Carter, 1930), A. bartolozzii (Ferrer, 2002), A. beloni Fairmaire, 1888), A. biangulatus (Gebien, 1921), A. borneensis (Pic, 1917), A. carinicollis (Gebien, 1921), A. conradti (Gebien, 1921), A. cribricollis (Schawaller, 2012), A. gabonicus (Pic, 1917), A. himalayicus (Kaszab, 1965), A. inaequalis (Pic, 1917), A. jacobsoni (Gebien, 1927), A. lateralis (Pic, 1917), A. latus (Pic, 1917), A. longeplicatus (Gebien, 1921) , A. maculipennis (Schawaller, 2011), A. major (Pic, 1917), A. nepalicus (Kaszab, 1975), A. nigrita (Gebien, 1927), A. notatus (Pic, 1923), A. pakistanicus (Schawaller, 1996), A. pinguis (Gebien, 1927), A. punctatus (Carter, 1914), A. raffrayi (Pic, 1917), A. rufithorax (Pic, 1917), A. rufus (Pic, 1917), A. serrimargo (Gebien, 1914), A. sumatrensis (Pic, 1917), A. terminatus (Gebien, 1921), A. testaceicornis (Pic, 1921), A. testaceipes (Pic, 1917), A. thailandicus (Schawaller, 2012), A. trautneri (Schawaller, 1994); and 13 restored combinations: Anaedus boliviensis (Pic, 1934), A. claveri (Pic, 1917), A. diversicollis (Pic, 1917), A. elongatus (Pic, 1934), A. guyanensis (Pic, 1917), A. holtzi (Pic, 1934), A. inangulatus (Pic, 1934), A. inhumeralis (Pic, 1917), A. mendesensis (Pic, 1917), A. minutus (Pic, 1917), A. rufimembris (Pic, 1932), A. rufipennis (Pic, 1917), A. subelongatus (Pic, 1932). The new synonymies with Anaedus necessitate the following six New Replacement Names Anaedus maculipennis (for Spinolyprops maculatus Kulzer, 1954), A. grimmi (for Aspisoma forticornis Pic, 1917), A. minimus (for Anaedus minutus Pic, 1938), A. merkli (for Anaedus diversicollis Pic, 1938), A. ottomerkli (for Anaedus lateralis Pic, 1923), A. schawalleri (for Anaedus nepalicus Schawaller, 1994).
Capeluprops Schawaller, 2011 is removed from Lupropini and provisionally placed in Laenini Seidlitz, 1895. Plastica Waterhouse, 1903 is transferred from Apocryphini Lacordaire, 1859 to Laenini. Paralorelopsis Marcuzzi, 1994 is removed from Lupropini and provisionally placed in Lagriinae incertae sedis. Pseudesarcus Champion, 1913 is transferred from Lagriinae incertae sedis to Diaperinae incertae sedis. Falsotithassa Pic, 1934 is transferred from Lupropini to Leiochrinini Lewis, 1894 (Diaperinae). Mimocellus Wasmann, 1904 is transferred from Lupropini to Tenebrionidae incertae sedis, and likely belongs in either Diaperinae or Stenochiinae.
Darkling beetles, distribution, higher classification, identification keys, long-jointed beetles, nomenclature.
Dedication
During the preparation of this publication, one of us, our respected colleague and friend Ottó Merkl (1957–2021), passed away suddenly on his way to work. In his honor and in recognition of his help with this and other papers, we have named two species of Anaedus after him.
The family Tenebrionidae Latreille, 1802 presently contains 2,307 valid genera placed in 12 subfamilies (
The composition of lagriine tribes has been problematic for a long time. Many genera have been transferred to and from Lagriinae over the last 40 years. This has gradually led to some progress towards a better understanding of the subfamily and constituent tribes. The current subfamilial concept of Lagriinae is largely based upon the work of
Recent molecular studies have supported the monophyly of Lagriinae (
In the previously described studies, neither Goniaderini nor Lupropini were monophyletic, even after the taxonomic changes made in those papers (Fig.
Phylogeny of Lagriinae from
During independent work on the West Indian tenebrionid fauna, discrepancies between historic determinations of Prateus and Lorelus, by Theodore J. Spilman (1925–1996) and John T. Doyen (
In this study, we redefine Lupropini and Goniaderini, and establish a new tribe containing genera that were previously misclassified in the previous two. Keys to the genera in each of the three tribes and a key to the tribes of Lagriinae are provided. While examining material for this study, the need for new generic synonymies, and reversals of previous synonymies, were revealed. The tribal placements for several other genera are also fixed.
Specimens used in this study are deposited in the California Academy of Sciences, San Francisco, USA (
Specimens were examined with various stereomicroscopes. Photographs were made by use of the following systems: (1) Macropod Pro (Macroscopic Solutions), with a Canon EOS 5dsr camera body and 65mm lens. Images were stacked using Zerene Stacker v. 1.04. (2) A Nikon D5600 camera body mounted on a Stackshot rail system (Cognisys Inc.) equipped with a Laowa 60 mm or 25 mm macro lens. Images were stacked using Zerene Stacker v. 1.04. (3) A Zeiss Discovery.V20 Stereomicroscope with a Zeiss Axiocam 305 Color camera. Images were stacked using Zerene Stacker v. 1.04.
Morphological terminology follows
One character system critical for the definition of the tribes involved herein requires explanation. The closure of the mesocoxal cavity is subject to misinterpretation (Figs
Mesocoxal closure in Lagriinae. 2 Closed state, Prateus fusculus LeConte, 1862 (Prateini) 3 Open state, Phymatestes sp. (Goniaderini) 4 Open state, Capeluprops laenoides Schawaller, 2011 (Laenini) 5 Closed state distorted during preservation, Micropedinus sp. (Prateini). Abbreviation: msv (yellow polygon) = mesoventrite, msa (red polygon) = mesanepisternum, mse (green polygon) = mesepisternum, mtv (blue polygon) = metaventrite.
Under normal closed circumstances, the arms of the ventrites do clearly touch laterad the mesocoxa (Fig.
This “popped”-open condition is exemplified in Fig.
Prateus LeConte, 1862.
Body length: 1.5–6.0 mm, stout to elongate, glabrous or setose. Most species are unicolored, fuscous to piceous, but a few species are patterned.
Head : Eyes round to ovoid, at most feebly notched anteriorly by epistomal canthus. Antennae usually reaching the middle of the pronotum, sometimes extending just past the base of the pronotum; antennomeres obconical to moniliform with last three to five forming a weak to strong club.
Thorax
: Pronotum shape variable, usually quadrate to rectangular. Lateral margin complete, smooth to dentate. Procoxae clearly separated by prosternal process. Mesocoxal cavity laterally closed by meeting of lateral arms of meso- and metaventrite (Fig.
Abdomen : Intersegmental membranes visible between sternites V–VII, abdominal hinging tenebrionoid. Defensive glands absent. Ovipositor slender, with three to four clearly separated coxite lobes, terminal coxite digitate, gonostyli apical or subapical.
Dorsal habitus of representatives of Prateini genera. 6 Bolitrium chinensis (Kaszab, 1940), holotype 7 Enicmosoma sp. 8 Indenicmosoma punctator Kaszab, 1979 9 Iscanus trukensis (Kulzer, 1957), paratype 10 Mesotretis ferruginea Bates, 1872, syntype 11 Microcalcar instriatum (Pic, 1925) 12 Micropedinus sp. 13 Paratenetus punctatus Spinola, 1844 14 Tithassa corynomelas Pascoe, 1860. Scale bars: 0.5 mm (7, 8); 1 mm (9, 12, 13); images lacking scale bars were produced by Otto Merkl and sizes of specimens were not recorded before he passed.
Prateini is distinguished from Goniaderini and Lupropini by having the mesocoxal cavity closed (i.e., laterally closed by meeting of meso- and metaventrite) and absence of abdominal defensive glands.
In Lagriinae, this character combination is only shared with Cossyphini Latreille, 1802. These two tribes can easily be distinguished from each other by the general habitus; all species of Cossyphini have prominent pronotal and elytral flanges, and the pronotal flange covers the head. In Prateini, the pronotum never covers the head. Cossyphini also has medial hinging between abdominal sternites V–VII (i.e., tentyrioid hinging) and intersegmental membranes are not visible, while Prateini has lateral hinging between abdominal sternite V–VII (i.e., tenebrionoid hinging) and intersegmental membranes are visible.
Antennoluprops Schawaller, 2007a, Ardoiniellus* Schawaller, 2013, Bolitrium Gebien, 1914, Enicmosoma Gebien, 1922, Indenicmosoma Ardoin, 1964, Iscanus Fauvel, 1904, Kuschelus* Kaszab, 1982a, Lorelopsis Champion, 1896, Mesotretis Bates, 1872, Microcalcar Pic, 1925, Micropedinus Lewis, 1894, Paratenetus Spinola, 1845, Prateus LeConte, 1862, Terametus* Motschulsky, 1869 and Tithassa Pascoe, 1860.
Dorsal habitus of species of Prateus LeConte, 1862 and Lorelopsis Champion, 1896. 15 Prateus fusculus LeConte, 1862, type species of Prateus 16 P. priscus (Sharp, 1876), type species of Lorelus Sharp, 1876 17 Lorelopsis exilis (Champion, 1913) 18 L. trapeziderus (Champion, 1913). Scale bars: 1 mm.
Prateus LeConte, 1862: 238. Type species: Prateus fusculus Leconte, 1862.
=Lorelus Sharp, 1876. Type species: Lorelus priscus Sharp, 1876. syn. rest. (original synonymy by
The rarely collected Prateus fusculus of North America has not been critically studied since its description. Its placement in Goniaderini has led to the expectation that it has open meoscoxal cavities, but they are clearly closed (Fig.
Prateus angulatus† (Doyen & Poinar, 1994), comb. nov.
Prateus angustulus (Champion, 1913), comb. nov.
Prateus armatus (Montrouzier, 1860) [Trogosita], comb. nov.
Prateus biroi (Kaszab, 1956), comb. nov.
Prateus blairi (Kaszab, 1955), comb. nov.
Prateus brevicornis (Champion, 1896), comb. nov.
Prateus breviusculus (Champion, 1913), comb. nov.
Prateus caledonicus (Kaszab, 1982b), comb. nov.
Prateus carolinensis (Blair, 1940), comb. nov.
Prateus chinensis (Kaszab, 1940), comb. nov.
Prateus clarkei (Kulzer, 1957), comb. nov.
Prateus crassicornis (Broun, 1880), comb. nov.
= Lorelus sternalis Broun, 1910. Synonymy by
Prateus crassepunctatus (Kaszab, 1982b), comb. nov.
Prateus cribricollis (Kaszab, 1940), comb. nov.
Prateus curvipes (Champion, 1913), comb. nov.
Prateus dybasi (Kulzer, 1957), comb. nov.
Prateus fijianus (Kaszab, 1982b), comb. nov.
Prateus fumatus (Lea, 1929) [Mesotretis], comb. nov.
Prateus glabriventris (Kaszab, 1982b), comb. nov.
Prateus greensladei (Kaszab, 1982b), comb. nov.
Prateus guadeloupensis (Kaszab, 1940), comb. nov.
Prateus hirtus (Kaszab, 1982b), comb. nov.
Prateus ivoirensis (Ardoin, 1969), comb. nov.
Prateus kanak (Kaszab, 1986), comb. nov.
Prateus kaszabi (Watt, 1992), comb. nov.
Prateus laticornis (Watt, 1992), comb. nov.
Prateus latulus (Broun, 1910), comb. nov.
Prateus longicornis (Kaszab, 1982b), comb. nov.
Prateus mareensis (Kaszab, 1982b), comb. nov.
Prateus marginalis (Broun, 1910), comb. nov.
Prateus niger (Kaszab, 1982b), comb. nov.
Prateus norfolkianus (Kaszab, 1982b), comb. nov.
Prateus obtusus (Watt, 1992), comb. nov.
Prateus ocularis (Fauvel, 1904), comb. nov.
Prateus opacus (Watt, 1992), comb. nov.
Prateus palauensis (Kulzer, 1957), comb. nov.
Prateus politus (Watt, 1992), comb. nov.
Prateus priscus (Sharp, 1876), comb. nov.
Prateus prosternalis (Kaszab, 1982b), comb. nov.
Prateus pubescens (Broun, 1880), comb. nov.
Prateus pubipennis (Lea, 1929) [Mesotretis], comb. nov.
Prateus punctatus (Watt, 1992), comb. nov.
Prateus quadricollis (Broun, 1886), comb. nov.
Prateus queenslandicus (Kaszab, 1986), comb. nov.
Prateus rugifrons (Champion, 1913), comb. nov.
Prateus solomonis (Kaszab, 1982b), comb. nov.
Prateus tarsalis (Broun, 1910), comb. nov.
= Lorelus nigrescens Broun, 1910. Synonymy by
Prateus unicornis (Kaszab, 1982b), comb. nov.
Prateus watti (Kaszab, 1982b), comb. nov.
Micropedinus Lewis, 1894: 370. Type species: Micropedinus algae Lewis, 1894.
=Notoprataeus Carter, 1924:37. Type species: Notoprataeus litoralis Carter, 1924. Synonymy by
=Microlyprops Kaszab, 1939: 108. Type species: Microlyprops ceylonicus Kaszab, 1939. syn. nov.
This genus is known from littoral habitats in the Australasian, Indomalayan, and eastern Palearctic regions. The synonymy of Microlyprops was first suggested by Kaszab in his unpublished annotations within his physical copy of Gebien’s Catalog (
Lorelopsis Champion, 1896: 15. Type species: Lorelopsis pilosa Champion, 1896.
We reestablish Lorelopsis as a valid genus in Prateini based upon several characters mentioned in the key and discussion below. Further, we move several species described in Lorelus by
Lorelopsis bicolor (Doyen, 1993), comb. nov.
Lorelopsis glabrata (Doyen, 1993), comb. nov.
Lorelopsis exilis (Champion, 1913), comb. nov.
Lorelopsis foraminosa† (Doyen & Poinar, 1994), comb. nov.
Lorelopsis minutulis† (Doyen & Poinar, 1994), comb. nov.
Lorelopsis pilosa Champion, 1896, comb. rest.
Lorelopsis trapezidera (Champion, 1913), comb. nov.
Lorelopsis wolcotti (Doyen, 1993), comb. nov.
Besides having the characters of Prateini, Lorelopsis species are small, elongate, parallel-sided, and covered in fine, silky, erect to suberect setae. The pronotum is slightly to distinctly narrower than the base of the elytra and microspiculate on the lateral margin, each spicule with an associated projecting seta forming a fringing row of projecting setae. A distinct and newly observed character is a long, stout projecting seta on the dorsum of the head close to the hind edge of the eye. This seta is clearly visible in species with relatively sparse and short setae on the head (Fig.
1 | Antenna with 10 antennomeres | 2 |
– | Antenna with 11 antennomeres | 4 |
2 | Eye oval in lateral view, never prominent, narrowed by gena; pronotum with lateral margins of disc dorso-ventrally flattened; antennae highly and differently modified between sexes [Madagascar] (see |
Antennoluprops |
– | Eye round, prominent, sometimes conical; pronotal disc convex to lateral margins, not flattened at sides; antennae unmodified in both sexes | 3 |
3 | Body surface pubescent [Tropical Africa, Madagascar] (Fig. |
Enicmosoma |
– | Body surface glabrous [Southeast Asia, India] (Fig. |
Indenicmosoma |
4 | Body oval, posterior margin of pronotum extended medially, closely appressed to elytra; elytral epipleuron very wide, nearly as wide as width of metacoxa [South Africa] (see |
Terametus |
– | Body not oval, posterior margin of thorax not extended medially; elytral epipleuron much narrower than width of metacoxa | 5 |
5 | Front edge of clypeus produced and very shallowly emarginate at middle; tarsi short and broad, all preapical tarsomeres transverse; antenna short, not reaching middle of pronotum. Length: 3–4 mm. [Australia on beaches] (Fig. |
Mesotretis |
– | Front edge of clypeus produced or not, but never emarginate; tarsi narrower, at least some preapical tarsomeres longer than wide; antenna reaching middle of pronotum | 6 |
6 | Labial palp with terminal palpomere larger than subterminal; antennae from 7th antennomere to end gradually widened; body length: 1.2–5.4 mm [SE Asia, Australia, Pacific on beaches] (Fig. |
Micropedinus |
– | Labial palp with subterminal palpomere enlarged, terminal palpomere small and parallel-sided; antenna more abruptly broadened from antennomere 8 or 9, last 3–4 ca. equal in width | 7 |
7 | Scutellum not visible; species very small (1.8 mm) [New Caledonia] (see |
Kuschelus |
– | Scutellum usually visible (except one species of Prateus); species larger (> 2 mm) | 8 |
8 | Body surface fully pubescent; pronotum laterally setose | 9 |
– | Body glabrous or partially pubescent; pronotum laterally asetose | 11 |
9 | Eye very large, distance between eye and anterior edge of pronotum dorsally less than length of eye [North and South America] (Fig. |
Paratenetus |
– | Eye much smaller, distance between eye and anterior edge of pronotum dorsally equal or greater than length of eye | 10 |
10 | Body very elongate, parallel sided; head with a long stout seta dorsally near posterior edge of eye (obscured by other long setae when head is more densely setose); distance between eye and anterior edge of pronotum dorsally ca. equal to length of eye; pronotum laterally microspiculate; elytra not globose. [Tropical America] (Figs |
Lorelopsis |
– | Body rounded, wide; head without stout seta above eye; distance between eye and anterior edge of pronotum dorsally usually much greater than length of eye; pronotum distinctly constricted posteriorly; elytra globose [tropical America] (Fig. |
Tithassa |
11 | Pronotal base slightly narrower than elytral base, humeri rounded; body setose or not | 12 |
– | Pronotal base not or barely narrower and elytral base, humeri variable, often sub-angulate; body always glabrous | 13 |
12 | Antennal club with 3 antennomeres [widespread, mainly tropical Asia, Pacific, and Americas] (Figs |
Prateus |
– | Antennal club with 4 antennomeres [Southeast Asia] (Fig. |
Bolitrium |
13 | Humeri angulate, not rounded; eyes small; basal membrane of labrum covered by broadened edge of clypeus. [Pacific Islands] (Fig. |
Iscanus |
– | Humeri clearly rounded, not angulate; eyes larger; basal membrane of labrum visible or not | 14 |
14 | Head with transverse impression between eyes; small species (1.8–2.5 mm) [South Africa] (see |
Ardoiniellus |
– | Head without transverse impression between eyes; larger species (4–5.5 mm) [Madagascar] (Fig. |
Microcalcar |
Goniadera Perty, 1832.
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
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.
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.
Dorsal habitus of representatives of Goniaderini genera. 21) Aemymone cariosa
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 (
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.
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.
Anaedus clearly belongs morphologically within Goniaderini, which is consistent with molecular analyses (
The reinstatement of Aemymone and Opatresthes is summarized in the following checklists. Note that many authorship and year attributions of
Aemymone Bates, 1868: 314. Type species: Goniadera cariosa Bates, 1868.
Aemymone cariosa (Bates, 1868) [Goniadera], comb. rest. Note:
Aemymone crenata Champion, 1893, comb. rest.
= Goniadera championi Ferrer & Delatour, 2007. Replacement name due to secondary homonym. Note: When
Aemymone hansfranzi (Ferrer & Delatour, 2007) [Goniadera], comb. nov.
Aemymone semirufa Pic, 1917a, comb. rest.
Aemymone simplex (Fairmaire, 1889) [Goniadera], comb. nov.
= Aemymone bordoni Marcuzzi, 1994. Synonymy by
Aemymone striatipennis (Pic, 1934) [Anaedus], comb. nov. Synonymy with A. cariosa Fairmaire, 1873 by
= Goniadera cariosa Fairmaire, 1873. Junior primary homonym (in Goniadera) and secondary homonym (in Aemymone) of Goniadera cariosa Bates, 1868.
= Aemymone silvanae Marcuzzi, 1994. Synonymy by
As noted above, Goniadera cariosa Fairmaire, 1873 is a primary homonym of Goniaderia cariosa Bates, 1868, and now that both species are included in Aemymone, it is also a secondary homonym. Although both species may have been described to accommodate an unavailable Dejean species by the same name (Bates 1968;
Opatresthes Gebien, 1928: 192. Type species: Opatresthes binodosa Gebien, 1928.
Opatresthes binodosa Gebien, 1928, comb. rest.
Opatresthes quadrinodosa Gebien, 1928, comb. rest.
Opatresthes maesi (Ferrer & Delatour, 2007) [Goniadera], comb. nov.
Opatresthes tuberculifera (Fairmaire, 1889) [Goniadera], comb. nov.
Phymatestes Pascoe, 1866: 142. Type species: Lagria tuberculata Fabricius, 1787.
=Gamaxus Bates, 1868: 315. Type species: Gamaxus hauxwellii Bates, 1868. syn. rest. (original synonymy by
During this study, the holotype of Gamaxus hauxwelli Bates, 1868 (Fig.
Anaedus
Blanchard, 1842: pl. 14. Type species: Anaedus punctatissimus Blanchard, 1842 (Fig.
=Aspisoma Duponchel & Chevrolat, 1841: 240. Type species: Aspisoma fulvipenne Duponchel & Chevrolat, 1841. Synonymy by
=Anaedes Agassiz, 1846: 20. Type species: Anaedus punctatissimus Blanchard, 1842. Unjustified emendation, not in prevailing usage (
=Aspidosoma Agassiz, 1846: 36. Type species: Aspisoma fulcipenne Duponchel & Chevrolat, 1841. Unjustified emendation, not in prevailing usage (
=Microanaedus Pic, 1923: 16. Type species: Microanaedus notatus Pic, 1923. syn. nov. (Fig.
=Pengalenganus Pic, 1917a: 10. Type species: Pengalenganus inaequalis Pic, 1917a. syn. nov.
=Pseudanaedus Gebien, 1921: 107. Type species: Pseudanaedus biangulatus Gebien, 1921. syn. nov.(Fig.
=Pseudolyprops Fairmaire, 1882: 236. Type species: Pseudolyprops dilaticollis Fairmaire, 1882. syn. nov. (Fig.
=Spinolyprops Pic, 1917a: 12. Type species: Spinolyprops rufithorax Pic, 1917a. syn. nov. (Fig.
=Spinadaenus Pic, 1921: 18. Type species: Spinadaenus singularis Pic, 1921. syn. nov. (Fig.
=Sphingocorse Gebien, 1921: 110. Type species Sphingocorse angulicollis Gebien, 1921. syn. nov. (Fig.
=Trichulodes Carter, 1914: 223. Type species: Trichulodes punctatus Carter, 1914. Synonymized with Pseudolyprops by
Anaedus may be generally differentiated from other Goniaderini by the following combination of characters: (1) eyes reniform, not completely divided; (2) pronotum transverse, always wider than long, never divided by narrow waist; (3) femora lacking teeth and spines; (4) tarsal formula 5-5-4; (5) elytra with basal lateral margin distinctly serrate; (6) elytral striae in most species, at least confused basally, usually confused throughout entire length.
Anaedus is most similar to Aemymone, Lyprochelyda, and Ancylopoma. From Aemymone, it can be distinguished by the setae on the lateral margin of the elytra placed on the lateral carina (in Aemymone, the setae on the lateral margin of the elytra are placed dorsad to the lateral carina). In most species of Anaedus, elytral punctures are nearly always confused (punctures always in linear striae in Aemymone). In Anaedus, the basal lateral margin of the elytron is distinctly serrate (Figs
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
Distinguishing Anaedus and the newly synonymized genera has long been problematic. Characters initially used to distinguish these genera are here considered to be unreliable, especially when many species of this group were examined.
We examined 66 species of our broadened concept of Anaedus, including the type species of all newly synonymized genera except Pengaleganus. We examined the characters purported to distinguish these groups and discuss them below under specific synonymies. The updated diagnosis above delimits our broad concept of Anaedus from other members of Goniaderini.
Microanaedus (Fig.
Pengalenganus, known from the Indomalayan region, was also distinguished from Anaedus by the structure of the pronotum, which was described as short, strongly incised anteriorly in the middle, with anterior angles prominent, very constricted posteriorly to the middle, laterally margined and flattened, and laterally posteriorly incised (
Pseudanaedus (Fig.
Pseudolyprops (Fig.
Anaedus anaedoides (Gebien, 1921), comb. nov.
Anaedus albipes (Gebien, 1921), comb. nov.
Anaedus amboinensis (Kaszab, 1964), comb. nov.
Anaedus amplicollis (Fairmaire, 1896), comb. nov.
Anaedus australiae (Carter, 1930), comb. nov.
Anaedus beloni (Fairmaire, 1888), comb. nov.
Anaedus borneensis (Pic, 1917b), comb. nov.
Anaedus carinicollis (Gebien, 1921), comb. nov.
Anaedus gabonicus (Pic, 1917b), comb. nov.
Anaedus jacobsoni (Gebien, 1927), comb. nov.
Anaedus latus (Pic, 1917b), comb. nov.
Anaedus longeplicatus (Gebien, 1921), comb. nov.
Anaedus major (Pic, 1917b), comb. nov.
Anaedus nigrita (Gebien, 1927), comb. nov.
Anaedus pinguis (Gebien, 1927), comb. nov.
Anaedus punctatus (Carter, 1914), comb. nov.
Anaedus raffrayi (Pic, 1917b), comb. nov.
Anaedus rufus (Pic, 1917b), comb. nov.
Anaedus sumatrensis (Pic, 1917b), comb. nov.
Anaedus terminatus (Gebien, 1921), comb. nov.
Anaedus testaceipes (Pic, 1917b), comb. nov.
Spinolyprops (Fig.
Anaedus rufithorax (Pic, 1917a), comb. nov.
Anaedus maculipennis nom. nov. for Spinolyprops maculatus Kulzer, 1954: 21. Distribution: Sri Lanka. Secondary homonym of Anaedus maculatus Champion, 1886: 25. Distribution: Nicaragua and Panama.
Anaedus cribricollis (Schawaller, 2012), comb. nov.
Anaedus himalayicus (Kaszab, 1965), comb. nov.
Anaedus lateralis (Pic, 1917a), comb. nov.
Anaedus ottomerkli nom. nov., for Anaedus lateralis Pic, 1923: 16. Distribution: Vietnam. Secondary homonym of Anaedus lateralis (Pic, 1917a: 12) [Spinolyprops]. Distribution: Myanmar, Thailand, Laos, Malaysia, and Indonesia.
Anaedus pakistanicus (Schawaller, 1996), comb. nov.
Anaedus thailandicus (Schawaller, 2012), comb. nov.
Anaedus trautneri (Schawaller, 1994), comb. nov.
Spinadaenus (Fig.
Sphingocorse (Fig.
Anaedus angulicollis (Gebien, 1921), comb. nov.
Anaedus nepalicus (Kaszab, 1975), comb. nov.
Anaedus maculipennis (Schawaller, 2011), comb. nov.
Anaedus schawalleri nom. nov. for Anaedus nepalicus Schawaller, 1994: 267. Distribution: Nepal. Secondary homonym of Anaedus nepalicus (Kaszab, 1975) [Sphingocorse]. Distribution: Nepal.
Aspisoma Duponchel & Chevrolat, 1841 (Coleoptera: Tenebrionidae) has a complicated taxonomic history, and although its synonymy with Anaedus was established by
Several papers by
All four of Pic’s works indicate that the species were meant to be placed in Tenebrionidae. In each paper, the species are described between Anaedus and other genera which we here treat as synonyms (e.g., Pseudolyprops). Furthermore,
We recognize the following species as members of Tenebrionidae: Lagriinae which leaves no western hemisphere species described by Pic remaining in Aspisoma Laporte (Lampyridae). The combinations are restored to
Anaedus boliviensis (Pic, 1934: 36), comb. rest.
Anaedus claveri (Pic, 1917c: 13), comb. rest.
Anaedus diversicollis (Pic, 1917b: 22), comb. rest.
Anaedus elongatus (Pic, 1934: 36), comb. rest.
Anaedus grimmi nom. nov. for Aspisoma forticornis Pic, 1917b: 23. Distribution: Brazil. Secondary homonym of Anaedus forticornis (Fairmaire, 1883: 35) [Lyprops]. Distribution: Indonesia. See
Anaedus guyanensis (Pic, 1917b: 22), comb. rest.
Anaedus holtzi (Pic, 1934: 36), comb. rest.
Anaedus inangulatus (Pic, 1934: 35), comb. rest.
Anaedus inhumeralis (Pic, 1917b: 24), comb. rest.
Anaedus mendesensis (Pic, 1917b: 23), comb. rest.
Anaedus minutus (Pic, 1917b: 24), comb. rest.
Anaedus rufimembris (Pic, 1932: 17), comb. rest.
Anaedus rufipennis (Pic, 1917b: 23), comb. rest.
Anaedus subelongatus (Pic, 1932: 17), comb. rest.
Anaedus minutus (Pic, 1917b: 24), comb. rest.
Anaedus minimus nom. nov. for Anaedus minutus Pic, 1938: 16. Distribution: Vietnam. Secondary homonym of Anaedus minutus (Pic, 1917b) [Aspisoma] Distribution: Brazil.
Anaedus merkli nom. nov. for Anaedus diversicollis Pic, 1938: 17. Distribution: Vietnam. Secondary homonym of Anaedus diversicollis (Pic, 1917b: 22) [Aspisoma]. Distribution: Guyana.
1 | Pronotum divided into two clear sections by a narrow “waist”, at least anterior section bearing a large elongate horn-like spine laterally [Afrotropical] (Fig. |
Spinolagriella |
– | Pronotum not divided as above | 2 |
2 | Eyes completely divided [Tropical America] (Fig. |
Xanthicles |
– | Eyes not divided, typically reniform | 3 |
3 | At least some femora with teeth | 4 |
– | All femora without teeth | 5 |
4 | Middle and hind femora with large tooth, pronotum more or less explanate laterally, dorsum reddish or patterned yellow and black, not tuberculate [Afrotropical] (Fig. |
Lyprochelyda |
– | Profemur, sometimes other femora armed in males, surface metallic, tuberculate [tropical America] (Fig. |
Phymatestes |
5 | Anterior lateral angles of pronotum greatly extended forming posterior angled arcs with anterior margins with spinose extensions [tropical America] (Fig. |
Ancylopoma |
– | Anterior lateral angles of pronotum not greatly extended forming posterior angled arcs | 6 |
6 | Tarsal formula 5-5-4. Pronotum with lateral margin slightly concave before hind angles, hind angles variable | 7 |
– | Tarsal formula 4-4-4. Pronotum with lateral angles evenly rounded without concave aspect near hind angles, hind angles obtuse, never spinose [tropical America] (see |
Myrmecopeltoides |
7 | Basal hind tarsomere distinctly longer than length of tarsomeres 2+4 or nearly as long as the other tarsomeres together | 8 |
– | Basal hind tarsomere equal or subequal in size to terminal tarsomere | 9 |
8 | Lateral margin of elytra near humeral angle with setae placed in the marginal carina, rendering the margin interrupted and the outline variably serrate, if these serrations weak, punctures on elytral disc and apex always confused; elytral disc occasionally with punctures in distinct linear series (striate), but in those cases the lateral elytral margin distinctly serrate; widespread including Tropical America] (Figs |
Anaedus |
– | Lateral margin of elytra with setae dorsad the lateral carina, not interrupting it; elytral disc and apex with punctures in linear series [tropical America] (Figs |
Aemymone |
9 | Lateral margins of pronotum at most with one or two obtuse angles, slightly explanate in some species, not strongly explanate on anterior section, surface usually smooth but some species with small tubercles. Elytra striate to costate, some species with costae forming elongate or short tubercles, without large tubercles. Lateral aspect of elytra smooth, not strongly dentate [tropical America] (Fig. |
Goniadera |
– | Lateral margins of pronotum and elytra strongly dentate, pronotum strongly explanate on anterior two-thirds. Both pronotum and elytra with numerous large tubercles [tropical America] (Fig. |
Opatresthes |
*Acropachia Mäklin, 1875 [Tropical America] One species, pronotum with lateral pits. We would have to see type to confirm tribe and key placement.
*Microgoniadera Pic, 1917a [Tropical America] One species, possibly another striate form of Anaedus.
Luprops Hope, 1833.
Body length: 5.2–11.2 mm; stout to elongate, glabrous or setose. Most species are unicolored but some are bicolored (e.g., pronotum and elytra with different coloration).
Head : Eyes reniform, anteriorly notched by canthus, rarely completely divided. Antennae moderately long, usually reaching or slightly extending past base of pronotum; antennomeres obconical to moniliform.
Thorax : Pronotum shape variable, quadrate to cordate, usually narrower than width of elytra. Lateral margins complete. Procoxae clearly separated by prosternal process. Mesocoxal cavity laterally closed, at least partially, by mesepimeron. Elytra striate. Metathoracic wings well developed or absent. Legs slender, not fossorial, penultimate tarsomeres lobed or cupuliform.
Abdomen
: Intersegmental membranes visible between sternites V–VII, abdominal hinging tenebrionoid. Defensive glands present (Fig.
Lupropini can be distinguished from Goniaderini and Prateini by having the mesocoxal cavity open and abdominal defensive glands present.
In Lagriinae, this character combination is shared with Adeliini Kirby, 1828, Pycnocerini Lacordaire, 1859, and Lagriini. Lupropini can be distinguished from these tribes as follows:
Both Adeliini and Pycnocerini possess abdominal defensive glands, but their configuration is different from Lupropini. Adeliini defensive gland reservoirs open between sternites VIII and IX (Fig.
Lagriini also possesses paired defensive gland reservoirs between sternites VII and VIII (Figs
Coxelinus Fairmaire, 1869, Curtolyprops Pic, 1917d, Dichastops Gerstaecker, 1871 and Luprops Hope, 1833.
1 | Eyes completely divided by a broad epistomal canthus. [eastern and southern Africa] (monotypic D. subaeneus Gerstaecker, 1871 (Fig. |
Dichastops |
– | Eyes not divided (but sometimes narrowed) by a broad epistomal canthus | 2 |
2 | Pronotum with posterior margin notched subapically [Madagascar] (Fig. |
Coxelinus |
– | Pronotum with posterior margin entire | 3 |
3 | Body relatively narrow, elongate; temples rounded, shorter than eye length in dorsal view; pronotum with lateral carinae thin, not ending with prominent process; pronotal and elytral surface regularly convex or flattened, not vermiculate [Afrotropical, Palearctic, Indo-Malaysian, Australian] (Fig. |
Luprops |
– | Body very wide, elytra short; temples parallel-sided, rectangular posteriorly, longer than eye length in dorsal view; pronotum with lateral carinae thick, ending subanteriorly with prominent tooth-like process; pronotal and elytral surface coarsely uneven, vermiculate [Afrotropical] (Fig. |
Curtolyprops |
Capeluprops Schawaller, 2011: 271. Type species: Capeluproprs laenoides Schawaller, 2011.
Capeluprops Schawaller, 2011 is provisionally moved from Lupropini to Laenini. Capeluprops contains six species of small, litter-inhabiting, flightless tenebrionids restricted to southern South Africa (
Paratypes and recently collected specimens of the type species were examined (Figs
Plastica Waterhouse, 1903: 563. Type species: Plastica polita Waterhouse, 1903.
Plastica Waterhouse, 1903 is transferred from Apocryphini Lacordaire, 1859 (Tenebrioninae) to Laenini (Lagriinae). This genus contains a single species which occurs in high elevation arid regions around Lake Titicaca in Bolivia.
Capeluprops and Plastica, two genera transferred to Laenini. 50 Dorsal habitus of Capeluprops laenoides Schawaller, 2011, paratype 51 Dorsal habitus of Plastica polita Waterhouse, 1903, specimen compared with holotype 52 lateral view of head of C. laenoides, non-type specimen 53 lateral view of head of P. polita, same specimen as Fig.
Examination of images of the holotype stored in the
Phobelius Blanchard, 1842.
Phobelius (Fig.
In molecular phylogenetic studies that included Phobelius (
Before this study, two subtribes of Lagriini were recognized, Lagriina and Statirina. The two subtribes can be distinguished based on differences in the prothorax. In Lagriina, the lobes of the hypomera meet behind the procoxae (Fig.
As
We reinstate Phobeliina Ardoin, 1961 as a valid subtribe of Lagriini based upon the previous molecular phylogenetic analyses and morphological discussion presented above. We propose the following diagnosis of this lineage of Lagriini: body form stout; antennomeres stout, terminal antennomere not distinctly elongated in either sex; pronotum lacking lateral margin; procoxae separated by distinct prosternal process; hypomera extending mesally behind procoxae and both joined to prosternal process, not meeting each other; mesocoxae open; paired defensive glands present between abdominal sternites VII and VIII, glands large, conical.
Based upon our updated recognition of Phobeliina, we also tentatively include within it the genus Rhosaces Champion, 1889 (Figs
Paralorelopsis Marcuzzi, 1994: 117. Type species: Paralorelopsis bordoni Marcuzzi, 1994.
Pseudesarcus Champion, 1913: 115. Type species: Pseudesarcus villosus Champion, 1913.
Pseudesarcus is placed incertae sedis within Diaperinae. Pseudesarcus was described in the family Mycetophagidae and transferred to Lagriinae incertae sedis by
Defensive gland reservoirs of Lagriinae. 62 Lagria villosa (Fabricius, 1781), Lagriini: Lagriina 63 Statira sp., Lagriini: Statirina 64 Phobelius sp., Lagriini: Phobeliina 65 Luprops sp., Lupropini 66 Aediotorix sp., Pycnocerini 67 Cardiothorax sp., Adeliini. Abbreviation: dgr = defensive gland reservoir.
Pseudesarcus can be characterized by: stellate antennal sensoria present on antennomeres 5–11 (Fig.
Based on the above observations, Pseudesarcus is clearly not a lagriine (possesses stellate sensoria, lacks internal ridge of sternite VII) and seems to fall within the circumscription of Diaperinae (see
Falsotithassa Pic, 1934: 18. Type species: Falsotithassa sumatrana Pic, 1934.
Falsotithassa Pic, 1934 is transferred from Lupropini (Lagriinae) to Leiochrinini Lewis, 1894 (Diaperinae). Falsotithassa contains ten species of small Tenebrionidae distributed across the Indo-Malayan biogeographic region. In the original description of this genus, Pic noted its similarity to Tithassa, which in this present paper is classified in Prateini. Based on the ordering of the descriptions in
Dorsal habitus and structures of Pseudesarcus Champion, 1913. 68 Dorsal habitus of Pseudesarcus villosus Champion, 1913, holotype 69 antennae of Pseudesarcus sp. 70 defensive gland reservoirs of P. villosus 71 ovipositor of P. villosus 72 female internal reproductive tract of Pseudesarcus sp. 73 details of spermatheca/spermathecal accessory gland complex of Pseudesarcus sp. Abbreviations: cas = compound antennal sensoria, dgr = defensive gland reservoir, par = paraprocts, goc = gonocoxites, gos = gonostyli, od = oviduct, bc = bursa copulatrix, cs = capsular spermatheca, sag = spermathecal accessory gland. Fig.
Examination of character states not discussed by
Dorsal habitus and structures of Falsotithassa sumatrana Pic, 1934. 74 Dorsal habitus of holotype 75 antennae of non-type specimen 76 defensive gland reservoirs 77 ovipositor 78 portion of spermatheca. Abbreviations: cas = compound antennal sensoria, dgr = defensive gland reservoir, lc = lateral commissure, goc4 = fourth gonocoxite, gos = gonostylus, sp = spermatheca, sag = spermathecal accessory gland. Fig.
We transfer Falsotithassa to Leiochrinini based upon the three characters discussed above: (1) antennae with complex sensoria on antennomeres 4–11; (2) abdominal defensive gland reservoirs joined by lateral commissure; (3) female internal reproductive tract with large thin-walled spermathecae. Further support of this conclusion is provided by characters mentioned by
Mimocellus Wasmann, 1904: 11. Type species: Mimocellus trechoides Wasmann, 1904: 12.
Mimocellus is placed incertae sedis in Tenebrionidae belonging in either Diaperinae or Stenochiinae. Mimocellus contains seven sub-Saharan African species of Tenebrionidae, including several species that are associated with termite nests. In the original description,
Dorsal habitus and structures of Mimocellus Wasmann, 1904. 79 Dorsal habitus of Mimocellus trechoides Wasmann, 1904 80 antennae of M. trechoides 81 defensive gland reservoir of M. trechoides, one reservoir was damaged during dissection. Abbreviations: dgr = defensive gland reservoir. Scale bar: 1 mm.
As with Falsotithassa, examination of antennae clearly excludes Mimocellus from Lagriinae. Mimocellus has distinct patches of complex antennal sensoria (Fig.
Archaeoluprops Nabozhenko, Perkovsky & Nazarenko, 2023.
The tribe Archaeolupropini Nabozhenko, Perkovsky & Nazarenko, 2023 is transferred from Lagriinae to Tetratomidae: Tetratominae Billberg, 1820. This tribe was recently described for a single beetle preserved in Eocene amber (
Archaeoluprops groehni Nabozhenko, Perkovsky & Nazarenko, 2023 possesses the following characters consistent with Tetratomidae: the basal two ventrites connate with 3–5 articulated; antennal insertions visible from above; elongate and linear terminal maxillary palpomeres; vertical lateral aspect of the abdominal ventrites which fit beneath the elytra; paired depressions near the posterior pronotal margin; hind coxae elongate, not bounded laterally by the sides of the first abdominal ventrite. The images do not clearly show the procoxal closure, but it appears they could be open externally. The lack of elytral striae, shape of the scutellar shield, and general facies indicate that this species belongs in the nominate subfamily Tetratominae, though the available specimen does not allow for examination of antennal clubs or male genitalia which are the primary features currently used to separate tetratomid subfamilies (
1 | Mesocoxal cavities closed (i.e., meso- and metaventrites fully enclosing mesocoxal cavity) | 2 |
– | Mesocoxal cavities open (i.e., laterally at least partially closed by mesepimeron) | 3 |
2 | Pronotum strongly flanged, covering head; abdominal membranes not exposed [Afrotropical, Palearctic, Indo-Malaysian] | Cossyphini |
– | Head always visible from above, pronotum without flanges covering the head; abdomen with exposed membranes between sternites V–VII [worldwide] | Prateini |
3 | All species with eyes, eyes typically reniform | 5 |
– | Few or all species without eyes, if eyes present, eyes rounded, globose; lacking sternal defensive glands | 4 |
4 | All species completely lacking eyes [Nearctic, associated with subterranean streams] | Eschatoporini |
– | Almost all species with rounded eyes, which may be reduced in size and sometimes absent [Palearctic, Neotropical, Afrotropical, Indo-Malayan] | Laenini |
5 | Penultimate protarsomeres simple, not ventrally prolonged into lobes | 6 |
– | Penultimate protarsomeres prolonged ventrally into lobes | 7 |
6 | Intersegmental membrane not visible between abdominal sternites; defensive glands absent [Nearctic, Palearctic, Neotropical and Australia] | Belopini |
– | Intersegmental membrane clearly visible between abdominal sternites; single defensive gland between sternites VII and VIII [Afrotropical and Indo-Malayan] | Pycnocerini |
7 | Body globose; legs fossorial [Australasian, Australia, New Zealand, on beaches] | Chaerodini |
– | Body not globose; legs not fossorial | 8 |
8 | Prosternal process very narrow, procoxae nearly contiguous [worldwide except Nearctic] | Lagriini: Lagriina |
– | Prosternal process always visible between procoxae | 9 |
9 | Terminal antennomere very elongate especially in males [worldwide except Europe] | Lagriini: Statirina |
– | Terminal antennomere normal length in both sexes | 10 |
10 | Pronotum lacking carina separating pronotum from epipleura [Neotropical] | Lagriini: Phobeliina |
– | Pronotum always with carina separating pronotum from epipleura | 11 |
11 | Abdominal defensive glands present | 12 |
– | Abdominal defensive glands absent [worldwide] | Goniaderini |
12 | Abdominal defensive glands paired between sternites VIII and IX [Neotropical and Australasian] | Adeliini |
– | Abdominal defensive glands paired between sternites VII and VIII [Afrotropical, Palearctic, Indo-Malayan, Australasian, Oceanic] | Lupropini |
A list of the proposed changes from the current positions of pertinent genera within Lagriinae are summarized in Table
Genera treated or figured in this study with previous classification, proposed taxonomic changes, and figure number.
Genus | Previous classification | Taxonomic changes | Figure(s) |
---|---|---|---|
Acropachia Mäklin, 1875 | Goniaderini | – | – |
Aemymone Bates, 1868 | Goniaderini (subgenus of Goniadera) | Goniaderini (valid genus) | 21, 45 |
Aediotorix Bates, 1868 | Pycnocerini | – | 66 |
Anaedus Blanchard, 1842 | Goniaderini | – | 30–44 |
Ancylopoma Pascoe, 1871 | Goniaderini | – | 28 |
Antennoluprops Schawaller, 2007 | Lupropini | Prateini | – |
Archaeoluprops Tetratominae Nabozhenko, Perkovsky & Nazarenko, 2023 | Archaeolupropini (Tenebrionidae: Lagriinae) | Archaeolupropini (Tetratomidae: Tetratominae) | – |
Ardoiniellus Schawaller, 2013 | Lupropini | Prateini | – |
Bolitrium Gebien, 1914 | Lupropini | Prateini | 6 |
Capeluprops Schawaller, 2011 | Lupropini | Laenini | 4, 50, 52 |
Cardiothorax Motschulsky, 1860 | Adeliini | – | 67 |
Coxelinus Fairmaire, 1869 | Lupropini | – | 46 |
Curtolyprops Pic, 1917 | Lupropini | – | 47 |
Dichastops Gerstaecker, 1871 | Lupropini | – | 48 |
Enicmosoma Gebien, 1922 | Lupropini | Prateini | 7 |
Falsotithassa Pic, 1934 | Lupropini | Leiochrinini (Diaperinae) | 74–78 |
Gamaxus Bates, 1868 | Goniaderini | Goniaderini (synonym of Phymatestes) | 29 |
Goniadera Perty, 1830 | Goniaderini | – | 22 |
Indenicmosoma Ardoin, 1964 | Lupropini | Prateini | 8 |
Iscanus Fauvel, 1904 | Lupropini | Prateini | 9 |
Kuschelus Kaszab, 1982 | Lupropini | Prateini | – |
Lagria Fabricius, 1775 | Lagriini: Lagriina | – | 54, 58, 62 |
Lorelopsis Champion, 1896 | Lupropini (synonym of Lorelus) | Prateini (valid genus) | 17–20 |
Lorelus Sharp, 1876 | Lupropini | Prateini (synonym of Prateus) | – |
Luprops Hope, 1833 | Lupropini | – | 49, 65 |
Lyprochelyda Fairmaire, 1899 | Goniaderini | – | 27 |
Mesotretis Bates, 1872 | Lupropini | Prateini | 10 |
Microanaedus Pic, 1923 | Goniaderini | Goniaderini (synonym of Anaedus) | 36 |
Microcalcar Pic, 1925 | Lupropini | Prateini | 11 |
Microgoniadera Pic, 1917a | Goniaderini | – | – |
Microlyprops Kaszab, 1939 | Goniaderini | Prateini (synonym of Micropedinus) | – |
Micropedinus Lewis, 1894 | Lupropini | Prateini | 5, 12 |
Mimocellus Wasmann, 1904 | Lupropini | Tenebrionidae incertae sedis | 79–81 |
Opatresthes Gebien, 1928 | Goniaderini (subgenus of Goniadera) | Goniaderini (valid genus) | 25 |
Paralorelopsis Marcuzzi, 1994 | Lupropini | Lagriinae incertae sedis | – |
Paratenetus Spinola, 1845 | Goniaderini | Prateini | 13 |
Pengalenganus Pic, 1917 | Goniaderini | Goniaderini (synonym of Anaedus) | – |
Phobelius Blanchard, 1842 | Goniaderini | Lagriini: Phobeliina | 56, 60, 64 |
Phymatestes Pascoe, 1866 | Goniaderini | – | 3, 23, 29 |
Plastica C.O. Waterhouse, 1903 | Apocryphini | Laenini | 51, 53 |
Prateus LeConte, 1862 | Goniaderini | Prateini | 2, 15, 16 |
Pseudanaedus Gebien, 1921 | Goniaderini | Goniaderini (synonym of Anaedus) | 37 |
Pseudesarcus Champion, 1913 | Lagriinae incertae sedis | Diaperinae incertae sedis | 68–73 |
Pseudolyprops Fairmaire, 1882 | Goniaderini | Goniaderini (synonym of Anaedus) | 38 |
Rhosaces Champion, 1889 | Lagriini: Statirina | Lagriini: Phobeliina | 57, 61 |
Sphingocorse Gebien, 1921 | Lupropini | Goniaderini (synonym of Anaedus) | 41 |
Spinadaenus Pic, 1921 | Goniaderini | Goniaderini (synonym of Anaedus) | 40 |
Spinolagriella Pic, 1955 | Lupropini | Goniaderini | 24 |
Spinolyprops Pic, 1917 | Lupropini | Goniaderini (synonym of Anaedus) | 39, 42 |
Statira Lepeletier & Audinet-Serville, 1828 | Lagriini: Statirina | Lagriini: Statirina | 55, 59, 63 |
Terametus Motschulsky, 1869 | Lupropini | Prateini | – |
Tithassa Pascoe, 1860 | Goniaderini | Prateini | 14 |
Xanthicles Champion, 1886 | Goniaderini | – | 26 |
All curators and persons responsible who made contributions to this study are gratefully acknowledged for their help. Although we made a strong effort to examine specimens of all genera treated, this was greatly hindered by museum closures and staff shortages and by the travel and shipping restrictions during the COVID pandemic. For this reason, Christophe Rivier (MNHN), Dmitry Telnov (
The authors have declared that no competing interests exist.
No ethical statement was reported.
No funding was reported.
Conceptualization: RLA, KK, MAI, MAJ. Investigation: RLA, KK, MAI, OM, MAJ. Resources: RLA, KK, MAI, OM, MAJ. Visualization: RLA, KK, MAI, OM, MAJ. Writing - original draft: RLA, KK, MAI, MAJ. Writing - review and editing: RLA, KK, MAI, MAJ.
Kojun Kanda https://orcid.org/0000-0001-5561-8471
Ottó Merkl https://orcid.org/0000-0003-3301-273X
Michael A. Ivie https://orcid.org/0000-0003-0996-2946
M. Andrew Johnston https://orcid.org/0000-0002-0166-6985
All of the data that support the findings of this study are available in the main text.