A catalogue of the tribe Sepidiini Eschscholtz, 1829 (Tenebrionidae, Pimeliinae) of the world

Abstract This catalogue includes all valid family-group (six subtribes), genus-group (55 genera, 33 subgenera), and species-group names (1009 species and subspecies) of Sepidiini darkling beetles (Coleoptera: Tenebrionidae: Pimeliinae), and their available synonyms. For each name, the author, year, and page number of the description are provided, with additional information (e.g., type species for genus-group names, author of synonymies for invalid taxa, notes) depending on the taxon rank. Verified distributional records (loci typici and data acquired from revisionary publications) for all the species are gathered. Distribution of the subtribes is illustrated and discussed. Several new nomenclatural acts are included. The generic names Phanerotomea Koch, 1958 [= Ocnodes Fåhraeus, 1870] and Parmularia Koch, 1955 [= Psammodes Kirby, 1819] are new synonyms (valid names in square brackets). The following new combinations are proposed: Ocnodesacuductusacuductus (Ancey, 1883), O. acuductusufipanus (Koch, 1952), O. adamantinus (Koch, 1952), O. argenteofasciatus (Koch, 1953), O. arnoldiarnoldi (Koch, 1952), O. arnoldisabianus (Koch, 1952), O.barbosai (Koch, 1952), O.basilewskyi (Koch, 1952), O.bellmarleyi (Koch, 1952), O. benguelensis (Koch, 1952), O. bertolonii (Guérin-Méneville, 1844), O. blandus (Koch, 1952), O. brevicornis (Haag-Rutenberg, 1875), O. brunnescensbrunnescens (Haag-Rutenberg, 1871), O. brunnescensmolestus (Haag-Rutenberg, 1875), O. buccinator (Koch, 1952), O. bushmanicus (Koch, 1952), O. carbonarius (Gerstaecker, 1854), O. cardiopterus (Fairmaire, 1888), O. cataractus (Koch, 1952), O. cinerarius (Koch, 1952), O. complanatus (Koch, 1952), O. confertus (Koch, 1952), O. congruens (Péringuey, 1899), O. cordiventris (Haag-Rutenberg, 1871), O. crocodilinus (Koch, 1952), O. dimorphus (Koch, 1952), O. distinctus (Haag-Rutenberg, 1871), O. dolosus (Péringuey, 1899), O. dorsocostatus (Gebien, 1910), O. dubiosus (Péringuey, 1899), O. ejectus (Koch, 1952), O. epronoticus (Koch, 1952), O. erichsoni (Haag-Rutenberg, 1871), O. ferreiraeferreirae (Koch, 1952), O. ferreiraezulu (Koch, 1952), O. fettingi (Haag-Rutenberg, 1875), O. fistucans (Koch, 1952), O. fraternus (Haag-Rutenberg, 1875), O. freyi (Koch, 1952), O. freudei (Koch, 1952), O. fulgidus (Koch, 1952), O. funestus (Haag-Rutenberg, 1871), O. gemmeulus (Koch, 1952), O. gibberosulus (Péringuey, 1908), O. gibbus (Haag-Rutenberg, 1879), O. globosus (Haag-Rutenberg, 1871), O. granisterna (Koch, 1952), O. granulosicollis (Haag-Rutenberg, 1871), O.gridellii (Koch, 1960), O. gueriniguerini (Haag-Rutenberg, 1871), O. guerinilawrencii (Koch, 1954), O. guerinimancus (Koch 1954), O. haemorrhoidalishaemorrhoidalis (Koch, 1952), O. haemorrhoidalissalubris (Koch, 1952), O. heydeni (Haag-Rutenberg, 1871), O. humeralis (Haag-Rutenberg, 1871), O. humerangula (Koch, 1952), O. imbricatus (Koch, 1952), O.imitatorimitator (Péringuey, 1899), O. imitatorinvadens (Koch, 1952), O. inflatus (Koch, 1952), O. janssensi (Koch, 1952), O. javeti (Haag-Rutenberg, 1871), O. junodi (Péringuey, 1899), O. kulzeri (Koch, 1952), O. lacustris (Koch, 1952), O. laevigatus (Olivier, 1795), O. lanceolatus (Koch, 1953), O. licitus (Peringey, 1899), O. luctuosus (Haag-Rutenberg, 1871), O. luxurosus (Koch, 1952), O. maputoensis (Koch, 1952), O. marginicollis (Koch, 1952), O. martinsi (Koch, 1952), O. melleus (Koch, 1952), O. mendicusestermanni (Koch, 1952), O. mendicusmendicus (Péringuey, 1899), O. miles (Péringuey, 1908), O. mimeticus (Koch, 1952), O. misolampoides (Fairmaire, 1888), O. mixtus (Haag-Rutenberg, 1871), O. monacha (Koch, 1952), O. montanus (Koch, 1952), O. mozambicus (Koch, 1952), O. muliebriscurtus (Koch, 1952), O. muliebrismuliebris (Koch, 1952), O. muliebrissilvestris (Koch, 1952), O. nervosus (Haag-Rutenberg, 1871), O.notatum (Thunberg, 1787), O. notaticollis (Koch, 1952), O. odorans (Koch, 1952), O. opacus (Solier, 1843), O. osbecki (Billberg, 1815), O. overlaeti (Koch, 1952), O. ovulus (Haag-Rutenberg, 1871), O. pachysomaornata (Koch, 1952), O. pachysomapachysoma (Péringuey, 1892), O. papillosus (Koch, 1952), O. pedator (Fairmaire, 1888), O. perlucidus (Koch, 1952), O. planus (Koch, 1952), O. pretorianus (Koch, 1952), O. procursus (Péringuey, 1899), O. protectus (Koch, 1952), O. punctatissimus (Koch, 1952), O. puncticollis (Koch, 1952), O. punctipennisplanisculptus (Koch, 1952), O. punctipennispunctipennis (Harold, 1878), O. punctipleura (Koch, 1952), O. rhodesianus (Koch, 1952), O. roriferus (Koch, 1952), O. rufipes (Harold, 1878), O. saltuarius (Koch, 1952), O.scabricollis (Gerstaecker, 1854), O. scopulipes (Koch, 1952), O. scrobicollisgriqua (Koch, 1952), O. scrobicollissimulans (Koch, 1952), O. semirasus (Koch, 1952), O. semiscabrum (Haag-Rutenberg, 1871), O. sericicollis (Koch, 1952), O.similis (Péringuey, 1899), O. sjoestedti (Gebien, 1910), O. spatulipes (Koch, 1952), O. specularis (Péringuey, 1899), O. spinigerus (Koch, 1952), O. stevensoni (Koch, 1952), O. tarsocnoides (Koch, 1952), O. temulentus (Koch, 1952), O. tenebrosusmelanarius (Haag-Rutenberg, 1871), O. tenebrosustenebrosus (Erichson, 1843), O. tibialis (Haag-Rutenberg, 1871), O. torosus (Koch, 1952), O. transversicollis (Haag-Rutenberg, 1879), O. tumidus (Haag-Rutenberg, 1871), O. umvumanus (Koch, 1952), O. vagus (Péringuey, 1899), O. vaticinus (Péringuey, 1899), O. verecundus (Péringuey, 1899), O. vetustus (Koch, 1952), O. vexator (Péringuey, 1899), O. virago (Koch, 1952), O. warmeloi (Koch, 1953), O. zanzibaricus (Haag-Rutenberg, 1875), Psammophanesantinorii (Gridelli, 1939), and P.mirei (Pierre, 1979). The type species [placed in square brackets] of the following genus-group taxa are designated for the first time, Ocnodes Fåhraeus, 1870 [Ocnodesscrobicollis Fåhraeus, 1870], Psammodophysis Péringuey, 1899 [Psammodophysisprobes Péringuey, 1899], and Trachynotidus Péringuey, 1899 [Psammodesthoreyi Haag-Rutenberg, 1871]. A lectotype is designated for Histrionotusomercooperi Koch, 1955 in order to fix its taxonomic status. Ulamus Kamiński is introduced here as a replacement name for Echinotus Marwick, 1935 [Type species.Aviculaechinata Smith, 1817] (Mollusca: Pteriidae) to avoid homonymy with Echinotus Solier, 1843 (Coleoptera: Tenebrionidae).


Introduction
The Sepidiini Eschscholtz, 1829 are a diverse tribe of ground-dwelling darkling beetles (Tenebrionidae) of the subfamily Pimeliinae Latreille, 1802 (Figs 1-45). The tribe is widely distributed throughout the Afrotropical Realm, with several species reaching the southern part of the Western Palaearctic . Some Sepidiini (mainly Ocnodes Fåhraeus, 1870 and Psammodes Kirby, 1819) are commonly known for their tapping behaviour (sexual communication), which accounts for their vernacular name, the "toktokkies" (Lighton 1987, Matthews et al. 2010. The group also includes many large and/or morphologically remarkable species, e.g., Stridulomus sulcicollis (Péringuey, 1885), the largest (~ 80.0 mm) currently known tenebrionid species , Matthews et al. 2010. From the morphological perspective, Sepidiini are defined by the following combination of characters : (i) cardo and stipes of maxillae and prelabium not covered by mentum (Fig. 46), (ii) anterior margin of postgenae with a maxillary ridge or emargination (Fig. 46), (iii) antennae with eleven segments (Fig. 47), (iv) mesocoxae, in vast majority of cases, with visible trochantin (reduced in Sepidiina and a few Molurina Solier, 1843) ( Fig. 48), (v) large scutellum, extending across entire width of mesothoracic peduncle (Fig. 49),and (vi) elytral base without vertical articulation face (the pronotum consequently freely movable on scutellum). Doyen (1994) also noted that in many Sepidiini the abdominal-sternal interlocking mechanism is different from all other Pimeliinae with the epipleural edge of the elytron overlapping the expanded sternite edge, rather than dovetailing into a groove.
The last comprehensive checklist of the species currently classified within Sepidiini was published by Gebien (1937a). At that time, 581 species were listed and divided over two separate tribes, Molurini and Sepidiini. No subtribal classification was proposed. After Gebien's catalogue (1937a) more than 50 contributions were published on the taxonomy, nomenclature, and classification of today's Sepidiini (see references). This includes descriptions of more than 400 species, reinterpretation of extremely diverse genera (e.g., Ocnodes, Psammodes, Somaticus , fusion of the former tribes Molurini and Sepidiini (see , Doyen 1994, and finally, designation of the six currently recognised subtribes (see Bouchard et al. 2005Bouchard et al. , 2011.
From the strictly formal point of view, the validity of many names introduced after 1937 remained questionable (e.g., Histrionotus omercooperi , since their unusual "descriptions" were incorporated in remarks concerning other taxa. Furthermore, the taxonomic affiliation of many genera and species is uncertain because of ambiguous remarks made by the contributors, see notes in the catalogue below.
The main aim of this work is to synthesise available nomenclatural, taxonomic, and distributional information concerning Sepidiini.

Nomenclatural data
All nomenclaturally available family-, genus-, and species-group names are included. The author, year, and page of the original description are provided for each scientific name. The type genus for each family-group name and the type species and type fixation for each genus-group name are included. Type species fixed by original designation were only accepted when an explicit statement (e.g., "Type species.") was used in the original publication (see ICZN 1999, Articles 67.5, 68.2). The reference in which a given generic or specific name is first placed in synonymy with the current valid name is listed [e.g., "syn. by Penrith (1986: 11)"]. For every species-group name which was subsequently transferred to another genus, the original genus-group name is provided. The author which transferred a particular species-group name to a currently accepted subgenus is listed in square parentheses (e.g., vagecostatus (Fairmaire, 1882b) Psammodes [Koch 1953a] listed under Psammophanes (Psammophanes)): in this example, Koch (1953a) was the one to include this species under the subgenus Psammophanes. The subtribal classification follows Bouchard et al. (2011).
Type deposition data and the status of the name bearing types was primarily taken from the original publications. However, when authors provided the name of the entomological collection for the type deposition without referring to any public institutions the following publications were used in order to locate those collections: ralearthdata.com). The list of localities was built by consulting available literature and is available as an Suppl. material to this publication (Suppl. material 1). Because of the uncertain status of most of the listed species, and extreme difficulties with identification of the majority of Sepidiini representatives, only records acquired from original species descriptions or revisionary papers were included. Geographic data with low degrees of accuracy (e.g., countries or states) were not georeferenced, and therefore are absent on the maps and distributional sections of particular subtribes.

Results and remarks
A total of 1009 valid species and subspecies divided over 55 genera (33 subgenera) and six subtribes is listed in this catalogue. The subtribe Molurina is the most diverse with 382 valid species-group taxa, followed by the Trachynotina (218), Phanerotomeina (177), Sepidiina (124), Oxurina (63), and Hypomelina (45). In most cases, the species and subspecies diversity is not equally divided over the available genera (Fig. 50). This is most evident in the case of Phanerotomeina, where a single genus, Ocnodes, groups over 83% of currently accepted species and subspecies. A similar trend is seen in Molurina and Trachynotina, while in Sepidiina over 80% of known species and subspecies diversity is divided between Sepidium Fabricius, 1775 and Vieta Hope, 1840 (Fig. 50). This tendency was not reported for the most recently revised subtribes (Louw 1979, Penrith 1986), i.e., Hypomelina and Oxurina. In total, 11 monotypic genera are listed.
Although this publication focuses on the nomenclature and classification of Sepidiini, the examined references enabled to reveal the most urgent taxonomic problems within the tribe. According to Penrith (1986Penrith ( , 1987, the status of the majority of currently recognised subtribes should be tested. This strictly relies on the verification of monophyly of the most speciose genera, such as Ocnodes and Psammodes. The taxonomic history of theses taxa is complex (see catalogue below), resulting in taxonomic ambiguities at the higher classification levels. The other urgent taxonomic problem within Sepidiini concerns the verification of the status of many genera of Molurina. The available contributions to the taxonomy of molurines presented in several different publications (e.g., Gebien 1910a, Wilke 1921, Koch 1951, 1952, 1953b, 1954a, 1956, 1960, 1962a. The lack of a comprehensive revisions may cause taxonomic inflation, especially when alphataxonomic contributions prevail. Future efforts concerning Sepidiini should include phylogenetic and revisionary studies. A database containing 2523 distributional records (857 not georeferenced) was created during this study (Fig. 51,Suppl. material 1). It needs to be highlighted that this list exclusively concerns loci typici and taxonomically revised data (acquired from generic revisions). However, because most Sepidiini species are known only from the type series, some basic remarks concerning distributional patterns of this tribe might be made.
According to the acquired data, Sepidiini are widely distributed throughout the Afrotropical Realm, except its northwestern parts (Fig. 51). Only Sepidiina has a distri- bution that extends into the Western Palaearctic (mainly Mediterranean Basin). However, the majority of the species of this subtribe were described from Somalia. Within the remaining subtribes, the presence of only Molurina and Phanerotomeina was revealed north of the equator. The former seems to be especially speciose in the Horn of Africa. Furthermore, Molurina is the only subtribe within Sepidiini with Malagasy representatives. The distribution of Hypomelina, Oxurina, and Trachynotina is limited to the southern part of the African continent. The majority of the species representing the Hypomelina were described from the Namibian coast (Fig. 51). Distribution. Western Palaearctic (mainly Mediterranean Basin) and Afrotropical Realm (with exception of western Africa) (Fig. 51). Distribution. Southern part of the Afrotropical Realm. Majority of species were described from the Namibian coast. A single genus, Bombocnodulus, reaching north to Central Africa (Fig. 51).
Genus originally described under Phanerotomeina, and subsequently transferred to Hypomelina by Louw (1979).
Notes. In 1904, Péringuey described a new species named damarinus under "Gen. Trachynotideus Péring". The spelling "Trachynotideus" was generally treated as an incorrect subsequent spelling of Trachynotidus by subsequent authors (e.g., Gebien 1910bGebien , 1937a and is not in prevailing usage. This view is also adopted here. Judging from the context, Péringuey (1904) did not intend to describe "Trachynotideus" as a new genus as he stated "genus … nov" or "n. gen." near the other newly introduced generic names, i.e., "DIESTESOMA, n. gen.".  provided a drawing of a species which he referred to as "Trachynotidus XXI-lineatus". However, this description does not meet the criteria of Art. 13.1.1 of the ICZN (1999).

Genus Microphligra Koch, 1955: 47
Type species. Phligra minuta Péringuey, 1904: (by original designation) Notes. In caption of plate 9 in his paper from 1955 Koch introduced the second representative of Microphligra: "Microphligra (Paraphligra) succulentium (subg. nov. in litt. sp. nov. in litt.)". A habitus of this beetle is presented in the preceding page. However, Koch did not specify any characters separating this new entity from the Microphligra minuta, therefore this cannot be treated as a valid description according to the regulations of the ICZN (1999).