﻿Reconstitution of some tribes and genera of Lagriinae (Coleoptera, Tenebrionidae)

﻿Abstract 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 PrateiniNew 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: Prateusangulatus (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: Micropedinusceylonicus (Kaszab, 1939) and M.maderi (Kaszab, 1940). LorelopsisRevised Status is revalidated as a genus and eight species formerly in Lorelus are transferred to it resulting in the following six New Combinations: Lorelopsisbicolor (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). Lorelopsispilosa 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: Aemymonehansfranzi (Ferrer & Delatour, 2007), A.simplex (Fairmaire, 1889), A.striatipennis (Pic, 1934) and Restored Combinations: Aemymonecariosa (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 Gamaxushauxwelli Bates, 1868 is placed as a New Synonym of Phymatestesbrevicornis (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: Anaedusalbipes (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: Anaedusboliviensis (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 NamesAnaedusmaculipennis (for Spinolypropsmaculatus Kulzer, 1954), A.grimmi (for Aspisomaforticornis Pic, 1917), A.minimus (for Anaedusminutus Pic, 1938), A.merkli (for Anaedusdiversicollis Pic, 1938), A.ottomerkli (for Anaeduslateralis Pic, 1923), A.schawalleri (for Anaedusnepalicus 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.


Introduction
The family Tenebrionidae Latreille, 1802 presently contains 2,307 valid genera placed in 12 subfamilies (Bouchard et al. 2021). The subfamily Lagriinae Latreille, 1825 contains 273 genera which represent ca. 12% of the entire family. Currently, we estimate the number of tenebrionid species to be more than 30,000 (Bouchard et al. 2021). Of these, more than 3,600 are placed in the subfamily Lagriinae.
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 Watt (1974) who utilized both adult and larval morphology. Lagriinae can generally be diagnosed using the following characters: adults with only simple sensoria on antenna; labrum subquadrate to elongate; mandibular mola with few coarse ridges, not finely striate; procoxal cavities completely closed internally and externally; elytron with ten striae plus scutellary striole; hind wing without subcubital fleck; ovipositor coxite usually slender; gonostylus elongate, digitiform, situated apically; known larvae with 2-segmented antenna; body pubescent; antennal and mandibular bases separated by narrow strip of head capsule (Watt 1974;Tschinkel and Doyen 1980;Doyen and Tschinkel 1982;Doyen et al. 1990;Matthews and Bouchard 2008;Matthews et al. 2010).
Recent molecular studies have supported the monophyly of Lagriinae (Kergoat et al. 2014;Kanda et al. 2015;Aalbu et al. 2017), but also demonstrated that several tribes are not monophyletic. Based on the results of Kanda et al. (2015), Kanda (2016) transferred the South American genus Chaetyllus Pascoe, 1860 from Lupropini Lesne, 1926to Laenini Seidlitz, 1895, the latter being previously considered a strictly Old World group. Kanda (2016) also described the genus Grabulax Kanda, 2016 as a second Neotropical genus of Laenini. Aalbu et al. (2017) used molecular sequence data to transfer Eschatoporis Blaisdell, 1906 out of Goniaderini Lacordaire, 1859 and re-established the monogeneric tribe Eschatoporini Blaisdell, 1906. Although these studies contributed towards establishing monophyletic tribes, the phylogenetic trees presented by them showed additional taxonomic issues throughout Lagriinae which have yet to be resolved.
In the previously described studies, neither Goniaderini nor Lupropini were monophyletic, even after the taxonomic changes made in those papers (Fig. 1).
Both are globally distributed tribes of predominantly litter-inhabiting and subcortical lagriines, which tend to be more diverse in tropical regions. These tribes are historically poorly defined and finding shared diagnosable characters among the taxa presently included in each has not been possible. In the molecular phylogenies presented in Kanda et al. (2015) and Aalbu et al. (2017), four genera that were classified in Goniaderini (Lorelus Sharp, 1876, Phobelius Blanchard, 1842, Paratenetus Spinola, 1845, and Prateus LeConte, 1862 are recovered outside of the clade containing Goniadera Perty, 1832 and related genera. Phobelius was recovered in Lagriini Latreille, 1825, and the remaining genera were recovered in a clade with two genera currently classified in Lupropini (Enicmosoma Gebien, 1922 andAntennoluprops Schawaller, 2007). A second clade of Lupropini was also recovered containing Luprops Hope, 1833 andCoxelinus Fairmaire, 1869. Although these issues were evident in the phylogenetic tree, the authors refrained from making taxonomic changes until a more detailed study of morphological characters could be conducted.
During independent work on the West Indian tenebrionid fauna, discrepancies between historic determinations of Prateus and Lorelus, by Theodore J. Spilman  and John T. Doyen (Doyen 1993;Doyen and Poinar 1994), led to the discovery of problems with the placement of Prateus. It became clear that the North American Prateus and West Indian Lorelus were 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.
congeneric and comparison with the New Zealand type species indicated this was correct. These two genera are currently placed in different tribes (Goniaderini and Lupropini respectively), clearly a problem. The resulting conclusion that Prateus was morphologically mischaracterized in its tribal placement caused a cascade of taxonomic and nomenclatural discoveries and an ever-widening set of issues, eventually with global implications. This fits with the problems of tribal definitions exposed by the molecular work, and this paper and two smaller taxonomic works (Ivie et al. 2021;Johnston et al. 2022) are the outcome.
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.  Kanda (KKIC), and Andrew Johnston (MAJC). Dissections and study of defensive glands were performed using protocols described by Tschinkel and Doyen (1980). In the following treatments of tribes, genera we could not examine are indicated with (*).

Materials and methods
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 Matthews et al. (2010) and , though we prefer using the term "sternite" to homologously number the ventral sclerite of abdominal sections to "ventrite" which typically refers only to the externally visible abdominal sternites. We primarily employ external morphology along with internal characters of defensive glands and female ovipositors to diagnose the tribes below. Female genital tracts have shown great diagnostic utility across other tenebrionid groups but have not historically been used within Lagriinae (Watt 1974;Tschinkel and Doyen 1980;Doyen and Tschinkel 1982;Doyen et al. 1990;Matthews 1998). We only examined the internal tracts of a small fraction of taxa in this study but we were unable to discern any diagnostic trends or putative synapomorphies for the constituent groups. Bibliographic references are given for every species-and genus-group name treated in this study.

Note on thoracic morphology
One character system critical for the definition of the tribes involved herein requires explanation. The closure of the mesocoxal cavity is subject to misinterpretation . This character has four elements in this group (five in some non-focal taxa). They are the lateral arms of the meso-and metaventrites, the mesepimeron and the mesocoxa (the fifth being the mesanepisternum in some other taxa). The closed condition is usually defined by the lateral arms of the ventrites touching laterad the coxa (Fig. 2). The open condition is defined as the lateral arms not touching (Figs 3,4), but this does not fully explain the true situation. Under normal closed circumstances, the arms of the ventrites do clearly touch laterad the mesocoxa (Fig. 2). However, in the relatively soft-bodied taxa (for tenebrionids) involved here, the closure of the mesocoxal cavity may be subject to distortion, especially if the specimen has been dorso-ventrally compressed or the body distended in fluid preparation, resulting in the closure being "popped" open. The important part of this characteristic is not if the mesoventrite touches the metaventrite laterad the mesocoxa, but if the mesepimeron normally impinges into the space between them to reach the mesocoxae. In compressed and/or particularly soft specimens, the mesoventrite and metaventrite arms may be separated by a gap, but to be considered open, the mesepimeron requires a facing surface for contact with the ends of both arms, and the mesepimeron clearly reaches the mesocoxa. Some specimens, including some name-bearing type specimens, have this distorted condition, and this has led to historical misunderstanding and misplacement of taxa.
This "popped"-open condition is exemplified in Fig. 5 where there is a bead on the facing parts where the meso-and metaventrites normally touch. Though they are stretched apart, the mesepimeron does not extend between them to reach the mesocoxa. If you imagine the arms of the mesocoxae are moved back towards each other, the posterior face of the mesoventrite arm meets and conforms to the anterior face of the metaventrite arm, and the mesepimeron does not extend between them to reach the mesocoxa. Thus, the mesocoxal cavity is closed. Alternatively, when the meso-and metaventrite arms are clearly unable to meet because the tip of the mesepimeron touches the mesocoxa (Figs 3, 4), the mesocoxal cavity is truly considered open.
Description. 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. 2). Elytra usually confusedly punctured, rarely with well-defined striae. Metathoracic wings usually well developed, but reduced or absent in some species. Legs slender, not fossorial. Penultimate tarsomeres lobed or cupuliform.

Diagnosis.
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.

Taxonomic changes among Prateini genera
The synonymy was first proposed by Van Dyke (1953), who simultaneously described a second species, P. dentatus VanDyke, 1953, but was missed by Zoological Record, and remained unrecognized by all subsequent workers including recent catalogs (Bousquet et al. 2018;Bouchard et al. 2021). The synonymy was listed for the genus but no species-level combinations were proposed, which are now made explicitly below.  (Sharp, 1876), type species of Lorelus Sharp, 1876 17 Lorelopsis exilis (Champion, 1913) 18 L. trapeziderus (Champion, 1913). Scale bars: 1 mm. Kaszab, 1939: 108. Type species: Microlyprops ceylonicus Kaszab, 1939. syn. nov. Note. 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 (Gebien 1941) complemented with handwritten remarks "[Microlyprops] maderi Kaszab […] = Micropedinus (Phaleriini), p. 497". Moreover, he placed the Microlyprops specimens in the material of Micropedinus in the collection of the Hungarian Natural History Museum, Budapest. One of the co-authors, Otto Merkl, studied this material and confirmed the synonymy, but

= Microlyprops
had not yet taken images before his passing. This synonymy results in the following species-group changes: Micropedinus ceylonicus (Kaszab, 1939), comb. nov. and Micropedinus maderi (Kaszab, 1940), comb. nov.  Wolcott (1936Wolcott ( , 1951; see also Blackwelder 1945) mentioned an undescribed species from Yauco, Puerto Rico, determined by Chapin as belonging to this genus. Doyen (1993) described Lorelus wolcotti Doyen, 1993 and listed Wolcott's citation as a synonym but did not mention having actually seen the specimens cited by Wolcott. Doyen (1993) also stated that "Lorelopsis is probably not distinct from Lorelus." Bouchard et al. (2021) record the two genera as synonyms, listing Doyen (1993) as a first synonymy ignoring the provisional nature of the statement. 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 Champion (1913), Doyen (1993), and Doyen and Poinar (1994) to Lorelopsis. The new concept of this genus includes the species given in the checklist below, though a number of undescribed species are also known from the West Indies. Note that Champion (1896: 15) considered this genus to be masculine with his single described species ending in -us and this was followed by all subsequent workers through Bousquet et al. (2018). However, following ICZN Article 30.1.2, Bouchard et al. (2021) appropriately treated this genus as feminine and the species epithets are emended accordingly here.

Lorelopsis pilosa
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. 19) but becomes less distinct when more dense and longer setae are present, blending with others ( Fig. 20). Since not all species assigned here have been examined (specifically several of the Champion species from the mainland), and since this character has not previously been mentioned, it is possible that it does not occur in all the mainland species, but it is there in the species we have seen. Some, but not all, species have the fourth tarsomere lobed beneath for a variable length. Champion used this as a primary character when he described the genus, but it has proved to be a species-level character. 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.
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 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.
Note. 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;Ferrer and Delatour 2007), the original descriptions suggest that each author formulated their description based on different specimens. To deal with the homonymy, Aemymone striatipennis (Pic, 1934), which was synonymized by Ferrer and Delatour (2007) with Aemymone cariosa (Fairmaire, 1873) is considered the valid name. Type specimens of both species must be examined before a decision can be made about whether Fairmaire's A. cariosa is a subjective synonym of Bates'.
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 42-44) whereas in Aemymone, the basal lateral margin of the elytron is smooth and never serrate (Fig. 45). Lyprochelyda possesses a wide,  (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). transverse pronotum and elytra with confused punctures similar to Anaedus but can be distinguished by the presence of a large tooth on the middle and hind femora. Anyclopoma possesses elytra with confused punctures like Anaedus, but the width of the base of the pronotum is shorter than the length of pronotum (see Johnston et al. 2022 for additional discussion). These three genera fall close to our expanded concept of Anaedus but seem to us recognizably distinct in the specimens at hand and are here retained as valid genera, though more data are desired to help clarify these relationships in the future.
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. Schawaller (2011) stated this problem, saying "the separation of the genera Pseudolyprops Fairmaire, 1882, Sphingocorse Gebien, 1921, and Spinolyprops Pic, 1917 within the tribe Lupropini[sic] is still in a preliminary state and not yet based on discriminating characters." At that time, Anaedus was placed in Goniaderini and therefore Schawaller did not include it or other similar genera within Goniaderini in his discussion and analysis. With our newly updated tribal concepts, the delimitation of these genera required additional investigation.
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. 36), known from Sumatra and Gabon, was distinguished from Anaedus by its small size (roughly 5 mm) and the structure of the prothorax, which is described as transverse, laterally crenulate, regularly arched, with posterior corners prominent (Pic 1923). Both the size and the structure of the prothorax fall clearly within the range of Anaedus. Prominent hind angles are used as a character to distinguish other synonymized genera including Spinolyprops. Numerous examined Anaedus species also have this character, and thus it is not reliable for distinguishing genera in this complex. Microanaedus is placed as a synonym resulting in Anaedus notatus (Pic, 1923), comb. nov. and Anaedus bartolozzii (Ferrer, 2002), comb. nov.
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 (Pic 1917a). Although we have not examined specimens attributed to this genus, the description of the pronotum falls within the diversity seen in Anaedus. Additionally, the synonym was first suggested by Kaszab in his unpublished annotations in his physical copy of the Gebien (1941) Catalog complemented with handwritten remarks "Pengalenganus Pic = Anaedus!" and his comment about the type species "9731 inaequalis Pic. Mel. Ent, 23, 1917, 10 Java = Anaedus 9759A." He spent considerable time studying tenebrionid material deposited in the Muséum national d'Histoire Naturelle, Paris, and very likely saw Pic's types. This synonymy results in Anaedus inaequalis (Pic, 1917a), comb. nov., Anaedus angustatus (Pic, 1921), comb. nov., and Anaedus testaceicornis (Pic, 1921), comb. nov.
Pseudanaedus (Fig. 37), with two species known from Cameroon, is characterized mainly by what Gebien considered to be a deep groove around the dorsal lobe of the eye. However, other species of Anaedus have grooves around the eyes to varying degrees. In South America, this feature seems more prevalent in species with a pronotum with spinose posterior angles. Pseudanaedus was also characterized by being hairy. This character state is also present in numerous Anaedus species, as well as newly synonymized genera (e.g., Pseudolyprops and Spinolyprops), and is not diagnostic. Therefore, Pseudanaedus Gebien, 1921 is placed as a synonym of Anaedus resulting in: Anaedus biangulatus (Gebien, 1921), comb. nov. and Anaedus conradti (Gebien, 1921), comb. nov.
Sphingocorse (Fig. 41), known from Central Africa and Asia, was considered by Gebien to be very close to Pseudanaedus, differing in the shape of the penultimate hind tarsomere and absence of a deep groove around the top of the eye (Gebien 1921). In his key to African 'Heterotarsinae' (an old subfamily concept, which contained Anaedus, Luprops, and other genera considered to be similar), Sphingocorse and Pseudanaedus are distinguished from Anaedus by the shape of the pronotum. Again, the shape of the pronotum is not distinct and falls within the diversity of forms seen In Anaedus. We place Sphingocorse in synonymy with Anaedus, resulting in the following new combinations, and necessitating one new replacement name.
Aspisoma Duponchel & Chevrolat, 1841 (Coleoptera: Tenebrionidae) has a complicated taxonomic history, and although its synonymy with Anaedus was established by Lacordaire (1859) and has persisted to this day, we uncovered additional taxonomic issues concerning this name. The name 'Aspisoma' was published in Dejean's (1834) second catalog as a genus belonging to Hétéromères: Ténébrionites but included no available species and thus is not available from that publication (Bousquet and Bouchard 2013). The name was validated by Duponchel and Chevrolat (1841) who, referring the name to Dejean, diagnosed the genus and included the type species Aspisoma fulvipenne Duponchel & Chevrolat, 1841. However, the authors failed to realize that the name "Aspisoma" had already been published by Laporte (1833) for a genus of Lampyridae (Coleoptera), and thus Aspisoma Duponchel & Chevrolat, 1841 is a junior homonym of Aspisoma Laporte, 1833.
Several papers by Pic (1917bPic ( , 1917cPic ( , 1932Pic ( , 1934 described 14 new species in the genus "Aspisoma" which have long been confused in catalogs and checklists. Gebien (1941) recognized these species as belonging to Aspisoma Duponchel & Chevrolat (Tenebrionidae) and therefore included them within the genus Anaedus following Lacordaire's synonymy. However, Blackwelder (1945) listed all 14 Pic species in Aspisoma Laporte (Lampyridae) where they have continued to be listed (McDermott 1966). One of the species described by Pic (Aspisoma inangulata Pic, 1934) was included as a member of Anaedus by Bousquet et al. (2018).
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, Pic (1917b) compares one of his species to a species of Anaedus. We have not seen any of these types but from the descriptions and arrangement in his works we are confident that Pic meant to place these species in Aspisoma Duponchel & Chevrolat (Tenebrionidae), though it is not clear if he merely missed Lacordaire's synonymy or truly intended to return the group to genus rank.

Anaedus minutus
Provisional key to the genera of Goniaderini 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. 22) (Fig. 25)  Description. 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).

Diagnosis. 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. 67) and Pycnocerini possesses just a single rectangular reservoir located medially between sternites VII and VIII (Fig. 66). In contrast, Lupropini has paired reservoirs that open between sternites VII and VIII (Fig. 65).
Lagriini also possesses paired defensive gland reservoirs between sternites VII and VIII (Figs 62-64). This tribe is the most speciose in Lagriinae, and is currently divided into three subtribes: Lagriina Latreille, 1825, Statirina Blanchard, 1825, and Phobeliina Ardoin, 1961 (see below for justification of the inclusion of Phobeliina in Lagriini). The subtribes can be distinguished from Lupropini as follows. Lagriina is characterized by the terminal antennomere elongate in most species; prosternal process greatly reduced, resulting in the procoxae appearing to be nearly contiguous; pronotum lacks lateral carinae. Statirina is characterized by the terminal antennomere elongate in all species; prosternal process narrow or wide, clearly separating procoxae; pronotum has complete lateral carinae. Phobeliina is characterized by the terminal antennomere subequal to penultimate antennomere; prosternal process wide, clearly separating procoxae; pronotum lacks lateral carina. In contrast, Lupropini has terminal antennomere subequal to penultimate antennomere; prosternal process wide, clearly separating procoxae, pronotum with lateral carinae clearly developed, at least in anterior fourth.

Note. 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 (Schawaller 2011). The genus was included in Lupropini without morphological discussion and the original description of the genus did not discuss the closure of the mesocoxal cavity nor presence or absence of defensive glands. Paratypes and recently collected specimens of the type species were examined (Figs 4, 43, 45). The mesocoxal cavities of this species are open, as in Lupropini, but abdominal defensive glands are absent. Therefore, this genus is excluded from Lupropini. Five lagriine tribes share these two character states: Belopini, Chaerodini, Eschatoporini, Laenini, and Goniaderini. In Capeluprops, the presence of well-developed eyes, tenebrionoid abdominal hinging, and lack of highly modified adaptions for psammophily exclude it from the first three tribes. However, Capeluprops cannot be definitively placed in Laenini nor Goniaderini. As in all other known Laenini, Capeluprops lacks hind wings, and the elytra are fused. Although the eyes of Capeluprops (Fig. 45) are more developed than typical members of the tribe, the current definition of Laenini based on molecular and morphological data (Kanda 2016) includes species in South America with slightly reniform eyes (e.g., some species of Chaetyllus Pascoe, 1860 and Grabulax darlingtoni Kanda 2016). The ovipositor is very similar to those in Goniaderini, being very stout with long digitate gonocoxites. However, this character state is also present in a few Lupropini. Based on the absence of wings, and overall body form, we provisionally move Capeluprops to Laenini. Further data and a comprehensive review of Laenini are needed to confirm this placement.
Genus Plastica Waterhouse, 1903Figs 51, 53 Plastica Waterhouse, 1903. Type species: Plastica polita Waterhouse, 1903. Note. 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. Waterhouse (1903) placed Plastica in Apocryphini based on its apparent similarity to species in the genus Apocrypha Eschscholtz, 1831, but separated the two genera based on differences in the femora and tarsi. Apocrypha, as with all other members of Tenebrioninae, possess abdominal defensive glands that open between abdominal sternites VII and VIII.
Examination of images of the holotype stored in the NHMUK provided by Dmitry Telnov and dissection of specimens matched with the holotype clearly place Plastica in Laenini. Plastica polita does not possess abdominal defensive glands, excluding it from any lineages of Tenebrioninae. The following characters support its placement in Laenini: eyes small and round, not emarginate anteriorly (Fig. 53); mesocoxal cavity open; hind wings absent; elytral humeri rounded; abdomen with visible intersegmental membranes between abdominal sternites V-VII, lateral (tenebrionoid) hinging between these segments; abdomen lacking defensive glands. Ardoin, 1961, stat. rev. Figs 56, 57, 60, 61, 64 Type genus. Phobelius Blanchard, 1842. Note. Phobelius (Fig. 56) contains 13 Neotropical species. The genus was included in the group "Phobéliides" by Lacordaire (1859) within the tribe Hétérotarsides along with Phymatestes, Anaedus, and Luprops. Subsequently, Ardoin (1961) included Phobeliina as a subtribe of Adeliini and transferred all genera except Phobelius to other groups. Matthews (1998), in his comprehensive review of Adeliini, noted that Phobelius exhibits characters consistent with Lagriini and that the only difference between Phobelius and other members of the tribe was that Phobelius did not have the elongate terminal antennomere typically found in Lagriini. Matthews further concluded that Phobelius should be included in a third subtribe (separate from Lagriina and Statirina) in Lagriini. However, subsequent catalogus did not follow Matthews' assessment, and Phobelius is currently included within Goniaderini with Phobeliina similarly synonymized under this tribe (Bouchard et al. 2005(Bouchard et al. , 2011(Bouchard et al. , 2021Bousquet et al. 2018).

Subtribe Phobeliina
In molecular phylogenetic studies that included Phobelius (Kanda et al. 2015;Aalbu et al. 2017), the genus was recovered in a clade with Lagriini ( Fig. 1), supporting  can be distinguished from other Lagriinae by the following characters: presence of abdominal defensive glands that open between abdominal sternites VII and VIII; pronotum with lateral margins absent or weakly impressed; antennae usually with terminal antennomeres elongate. Although Phobelius does not have elongate terminal antennomeres, they do have abdominal defensive glands and lack lateral pronotal margins.
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. 58) and, in Statirina, the lobes of the hypomera do not meet and are separated by the prosternum (Fig. 59). The prosternal process in Lagriina is thin and recessed between strongly projecting procoxae, sometimes resulting in the procoxal cavities appearing to be contiguous. In Statirina, the prosternal process forms a complete strip of cuticle, approximately ¼ the width of the procoxa, and clearly separates the coxae throughout their entire length. In addition to prothoracic characters, Lagriina tend to be broader bodied while Statirina tend to be more slender. The elongation of the terminal antennomere tends to be much more pronounced in Statirina, and in some Lagriina the terminal antennomere is nearly the same length as the penultimate antennomere.
As Matthews (1998) noted, Phobelius does not neatly fit within either of the two subtribes. Its prothorax (Fig. 60) resembles Statirina; the lobes of the hypomera do not meet posterior to the coxae and the prosternal process is wide and not recessed as in Lagriina. The stout body (Fig. 56) is more like body forms seen in Lagriina. The terminal antennomere is also not particularly elongate in either males or females, at most only 1.5 times longer than the preceding one. The shape of the abdominal defensive gland reservoirs differs from both Lagriina and Statirina as well. In Phobelius, the gland reservoirs are large and conical, with wide openings (Fig. 64). In Lagriina and Statirina, the gland reservoirs are small, sometimes inconspicuous, and are widely separated (Figs 62, 63).
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 57, 61). This monotypic genus was erected for Rhosaces clavipes Champion, 1889 and placed within Statirina where it has been treated ever since (Blackwelder 1945;Bousquet et al. 2018;Bouchard et al. 2021), although Champion (1889) pointed out the strong differences in antennae (lacking an elongate terminal antennomere), a short epistoma, and a broadly rounded intercoxal process of the abdomen. All of the characters mentioned by Champion (1889) are shared with Phobelius, and the defensive glands, mesocoxal openings, and prothoracic characters similarly seem to unite these two genera. It is clear that Rhosaces does not fit within our concept of Lagriina, and it does adhere to our diagnosis of Phobeliina, and we look forward to future phylogenetic investigations that can more rigorously test the monophyly of this assemblage.

Genus Paralorelopsis Marcuzzi, 1994
Paralorelopsis Marcuzzi, 1994: 117. Type species: Paralorelopsis bordoni Marcuzzi, 1994. Note. Marcuzzi (1994), in his very limited description based on a single example, described Paralorelopsis as agreeing with Champion's description of Lorelopsis except for a single difference being the lack of a lamina on the subapical tarsomere. His new species, P. bordoni, is also much larger in size than species of either Lorelopsis or Prateus. In both the tarsi and size, it agrees more with some American genera belonging to Belopini. We were unable to examine specimens of this genus and therefore place it as incertae sedis in Lagriinae for now. Note. Pseudesarcus is placed incertae sedis within Diaperinae. Pseudesarcus was described in the family Mycetophagidae and transferred to Lagriinae incertae sedis by Lawrence and Newton (1995: 886) (Bousquet et al. 2018). Pseudesarcus villosus was described from two Panamanian specimens, one of which was photographed by Keita Matsumodo (Fig. 68) and the other examined for us by Maxwell Barclay (both from NHMUK). A third specimen from Costa Rica (Figs 70-73) was identified as this genus based upon the images and description of the types and was dissected to examine internal structures. We also identified a seemingly undescribed species from Ecuador that possessed internal and external characters used to diagnose Pseudesarcus (Fig. 69). Pseudesarcus is clearly a member of Tenebrionidae and part of the 'tenebrionoid-branch' (sensu Doyen and Tschinkel 1982; see Matthews and Bouchard 2008). Pseudesarcus can be characterized by: stellate antennal sensoria present on antennomeres 5-11 (Fig. 69); labrum concealed beneath epistoma with symmetrical epistomal tormae; lacinia lacking uncus; procoxal cavities closed internally and externally; mesocoxae closed laterally by mesoventrite and metaventrite; paired defensive glands present, lacking common volume, not pleated (Fig. 70); female genital tract with secondary bursa copulatrix, spermatheca forming annulated sclerotized capsule at end of spermathecal gland; ovipositor reduced .
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 Doyen and Tschinkel 1982;Matthews and Bouchard 2008;Johnston et al. 2020), but lacks any clear relationships with the established tribes (see Johnston et al. 2020). We place it as incertae sedis within Diaperinae until such time as its constituent tribes are better understood. Pic, 1934Figs 74-78 Falsotithassa Pic, 1934: 18. Type species: Falsotithassa sumatrana Pic, 1934 Note. 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 Pic's (1934) manuscript, and the placement of the descriptions of Falsotithassa between species of Anaedus (Goniaderini) and Tithassa, it can be inferred that Pic considered this genus to be closely related to these taxa, and therefore included in Lagriinae. Schawaller (2000) revised Falsotithassa, providing a detailed diagnosis for this genus, and synonymized Derispiolina Kaszab, 1979 which was originally described as a tentative member of the tribe Leiochrinini (Kaszab 1979). In that study, Schawaller suggested that Falsotithassa belonged in Diaperini (Diaperinae), but in a later paper (Schawaller 2007c) reinterpreted the same characters studied in his initial 2000 revision as supporting the placement of Falsotithassa in Lupropini. This placement is adopted in the recent generic catalog of Tenebrionidae (Bouchard et al. 2021).

Genus Falsotithassa
Examination of character states not discussed by Schawaller (2000Schawaller ( , 2007c and reinterpretation of female reproductive structures described in these papers support the exclusion of Falsotithassa from Lagriinae and supports its inclusion in Leiochrinini. The most evident character for excluding Falsotithassa from Lagriinae is the presence of complex sensoria on antennomeres 4-11 (Fig. 75); all Lagriinae have only simple antennal sensoria. Additionally, the abdominal defensive gland reservoirs have a lateral commissure joining the base of the left and right reservoir (Fig. 76). This arrangement is only known to occur in Leiochrinini and Nilionini (Doyen et al. 1990;Matthews and Bouchard 2008). The main characters used by Schawaller (2000Schawaller ( , 2007c in placing Falsotithassa in Diaperini was the presence of a capsular spermatheca ("check valve"), a character that is only known to occur in Diaperini and Nilionini (Tschinkel and Doyen 1980;Aloquio and Lopes-Andrade 2016). New dissections of specimens identified as Falsotithassa sumatrana by Schawaller, and matched with the holotype, show a large thin-walled balloon-like spermathecae (Figs 77, 78). These are not the same as the capsular spermathecae present in Diaperini and Nilionini but are very similar to spermathecae illustrated for other species of Leiochrinini (Doyen et al. 1990;Matthews and Bouchard 2008). 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 Schawaller (2000), including internally open procoxal cavities and the female reproductive tract lacking a bursa copulatrix. However, Falsotithassa departs from the coccinellid-like appearance of all other current members of Leiochrinini and indeed is externally similar to members of Scaphidemini Reitter, 1922. The latter is presently defined by a strongly sclerotized T-shaped spermatheca and the defensive glands lacking a commissure (Doyen et al. 1990;Matthews and Bouchard 2008) which preclude the placement of Falsotithassa therein. We hypothesize that the tribes Leiochrinini and Scaphidemini are likely closely related and should be reevaluated with respect to each other in future studies. treatments of the genus and taxonomic catalogs place Mimocellus in Lupropini (Schawaller 2005;Robiche et al. 2002;Bouchard et al. 2021).

Genus
As with Falsotithassa, examination of antennae clearly excludes Mimocellus from Lagriinae. Mimocellus has distinct patches of complex antennal sensoria (Fig. 80). Additionally, the defensive gland reservoirs differ from those found in Lupropini. While lupropine gland reservoirs tend to be stout with wide openings (Fig. 65), Mimocellus possesses elongate gland reservoirs (Fig. 81) similar to the type found in Diaperini (Diaperinae) and Cnodalonini (Stenochiini) (Tschinkel and Doyen 1980). Due to limited availability of specimens, we were unable to examine female internal morphology. For this reason, we currently place this genus as incertae sedis within Tenebrionidae until its placement within the tenebrionoid-branch subfamilies can be elucidated.

Tribe Archaeolupropini Nabozhenko, Perkovsky & Nazarenko, 2023
Type genus. Archaeoluprops Nabozhenko, Perkovsky & Nazarenko, 2023. Note. 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 (Nabozhenko et al. 2023). This beetle was compared to several tribes of Lagriinae where it was found to not belong to any of them, and therefore was placed into a new tribe in the subfamily. Examination of the descriptions and excellent photographs in that paper demonstrate that this taxon clearly belongs within the family Tetratomidae.
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 (Nikitsky 1998). Based upon the preponderance of evidence, we hereby transfer Archaeolupropini to Tetratomidae: Tetratominae and leave it there as a valid tribe in that subfamily pending further revision.