Anaedus Blanchard, 1842

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

Genus Anaedus Blanchard, 1842

Figs 30-41 , 42-45

Anaedus Blanchard, 1842: pl. 14. Type species: Anaedus punctatissimus Blanchard, 1842 (Fig. 27 ).

= Aspisoma Duponchel & Chevrolat, 1841: 240. Type species: Aspisoma fulvipenne Duponchel & Chevrolat, 1841. Synonymy by Lacordaire 1859: 396, junior homonym of Aspisoma Laporte, 1833 ( Coleoptera : Lampyridae ).

= Anaedes Agassiz, 1846: 20. Type species: Anaedus punctatissimus Blanchard, 1842. Unjustified emendation, not in prevailing usage ( Bouchard et al. 2021).

= Aspidosoma Agassiz, 1846: 36. Type species: Aspisoma fulcipenne Duponchel & Chevrolat, 1841. Unjustified emendation, not in prevailing usage ( Bouchard et al. 2021).

= Microanaedus Pic, 1923: 16. Type species: Microanaedus notatus Pic, 1923. syn. nov. (Fig. 36 ).

= 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. 37 ).

= Pseudolyprops Fairmaire, 1882: 236. Type species: Pseudolyprops dilaticollis Fairmaire, 1882. syn. nov. (Fig. 38 ).

= Spinolyprops Pic, 1917a: 12. Type species: Spinolyprops rufithorax Pic, 1917a. syn. nov. (Fig. 39 ).

= Spinadaenus Pic, 1921: 18. Type species: Spinadaenus singularis Pic, 1921. syn. nov. (Fig. 40 ).

= Sphingocorse Gebien, 1921: 110. Type species Sphingocorse angulicollis Gebien, 1921. syn. nov. (Fig. 41 ).

= Trichulodes Carter, 1914: 223. Type species: Trichulodes punctatus Carter, 1914. Synonymized with Pseudolyprops by Doyen et al. (1990: 231).

Diagnosis.

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 42-44 ) whereas in Aemymone , the basal lateral margin of the elytron is smooth and never serrate (Fig. 45 ). Lyprochelyda possesses a wide, 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 Anaedus 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.

Pseudolyprops (Fig. 38 ), distributed in the Australasian and Indomalayan regions, is also distinguished by the shape of the pronotum ( Fairmaire 1882; Wei and Ren 2020). Again, this is not a diagnostic character and Pseudolyprops is placed in synonymy with Anaedus , resulting in the following new combinations:

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. 39 ), known from Asia, was also considered to have a unique prothorax ( Pic 1917a) and was characterized by the color patterning of the elytra. Again, the pronotum falls within the diversity seen in Anaedus . Patterned elytra are also seen in Anaedus species from multiple biogeographic realms as well as Microanaedus (Fig. 33 ) and thus is not a reliable character to distinguish genera. We place Spinolyprops as a synonym of Anaedus , resulting in the following new combinations, and necessitating two new replacement names.

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. 40 ), a monotypic genus known from Borneo, is unique with its extreme laterally spinose pronotum. Many species of Anaedus have a pronotum with undulate margins, and a few species, including Anaedus serratus Wei & Ren, 2020 have denticulate lateral margins. Although the pronotum of Spinadaenus is very spinose, we believe it represents a single extreme example of the range found within Anaedus and therefore the genus is placed as a synonym of Anaedus resulting in Anaedus serrimargo (Gebien, 1914), comb. nov. Note that the type species Spinadaenus singularis Pic, 1921 is considered a junior subjective synonym of Lyprops [sic] Anaedus serrimargo Gebien, 1914 ( Grimm and Schawaller 2021).

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.

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 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 (1917b, 1917c, 1932, 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.

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 Gebien’s (1941) inclusion within Anaedus and results in the following nomenclatural acts:

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 Grimm and Schawaller 2021.

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.

Figures

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

Figures 42 - 45. Elytral humerus of Anaedus and Aemymone. 42 Anaedus lateralis (Pic, 1917), formerly in Spinolyprops 43 Anaedus brunneus Ziegler, 1844 44 Anaedus punctatissimus Blanchard, 1842 45 Aemymone sp.

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