Goudotostes Paulian, 1979

Ballerio, Alberto, 2021, The Ceratocanthinae of Madagascar and Comoro Islands: a revision of the genera Synarmostes and Goudotostes, and of the flightless Philharmostes, with description of 64 new species (Coleoptera: Scarabaeoidea, Hybosoridae), Fragmenta entomologica 53 (2), pp. 105-282 : 168-170

publication ID

https://doi.org/ 10.13133/2284-4880/545

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scientific name

Goudotostes Paulian, 1979
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Genus Goudotostes Paulian, 1979 View in CoL

Paulian 1979: 60 (key, description and distribution); Ballerio 2006: 299 (sexual dimorphism); Ocampo & Ballerio 2006: 190 (listing); Ballerio & Grebennikov 2016 (phylogeny).

Type species (by monotypy): Acanthocerus scabrosus Laporte, 1840 .

Taxonomic history. See under Synarmostes .

Diagnosis. The genus can be identified by the following combination of characters: 1) enrollment coaptations complete, 2) head with or without dorsal ocular area, sometimes vestigial, 3) when dorsal ocular area is present the genal canthus is complete (fused with the occipital area of head or almost reaching it), 4) ten antennomeres, 5) antennae with pedicellus strongly bent forward (many other Ceratocanthinae genera have a bent pedicellus, but only the genera Goudotostes and Cryptosphaeroides in such an extreme way), 6) pronotum with basal margin and fore margin raised (often forming monolobate or bilobate transverse carinae), 7) apical portion of elytra without several long carinae and furrows but often with tubercles and short carinae (in this cae carinae occupy at least the median third of elytra too), 8) parameres asymmetrical.

Description. Small to large Ceratocanthinae (TLR from 1.8 mm to 3.6 mm). Dorsum shiny or dull, colour variable, although the dorsum of the majority of species is dark-coloured, sometimes with metallic sheen (bronze to green), especially on tubercles and carinae. Head subpentagonal, clypeus triangle-shaped with sides almost rectilinear and obtuse apex (about 110-115°); genae unaligned with respect to clypeal sides, forming a right angle with genal canthus and slightly protruding outwards; genal canthus (when present) almost always fused with the occipital area of head; dorsal ocular area absent or small or vestigial, ventral ocular area small or medium-sized; head surface plane or tuberculate, with variable punctation, transversal striae and pubescence.

Pronotum: subtrapezoidal or subrectangular, as wide as maximum elytral width, unevenly convex; anterior angles distinctly but slightly protruding forward, gently or strongly truncate; base broad, the whole pronotal margins marked by a continuous distinct impressed line (circumnotal ridge), often invisible in dorsal view. Base and fore margin raised. Basal carina and anterior carina monolobate or bilobate. Surface with variable punctation and pubescence.

Scutellum : large, about as long as wide (W/L ratio = 1), sides distinctly notched by elytral articular process, then convergent to form a triangle with apex elongate and acute and sides slightly curved inward; apical portion of mesepisterna not visible from above.

Elytra: slightly longer than wide or as long as wide, apex in lateral view fairly re-entering inward; regularly convex, pseudoepipleuron absent; elytral suture raised sometimes only on distal third; sutural stria often not visible; inferior sutural stria present; striated articular area hardly visible in lateral view, relatively narrow and short; marginal area present, elytral articular process well developed, smooth and shiny.

Clypeopleuron short, not furrowed. Mouthparts (based on Goudotostes orangeanus sp. nov., G. laevis sp. nov., G. mayottensis sp. nov., and G. randrianirinai sp. nov.): labrum wide and short, distally depressed at middle, bearing a row of long, erect, fine setae. Distal epipharynx ( Fig. 37 h View Fig , 38 h View Fig ) with semicircular or subtrapezoidal shape, longitudinally divided by a sharp anterior median process, distally raised; median brush and corypha absent; apical fringe made of long fine setae, absent in the middle; lateral combs made of long fine setae. Mentum ( Fig. 37 e View Fig , 38 e View Fig ) ventrally flat, deeply emarginated in the middle of anterior edge, emargination regularly wide-U-shaped; labial palpi (including palpiger) four segmented, fourth palp subconical, shorter than the preceding one which is plump- er. Maxillae ( Fig. 37 f View Fig , 38 f View Fig ) with a short single lacinia, covered with fine long setae, monolobed galea proximally sclerotized and distally clothed with very coarse long thin setae ending with a pectinate apex (galeal brush) ( Fig. 37 g View Fig , 38 g View Fig ); maxillary palpi (including palpiger) four segmented, fourth palpus long and subconical, slightly longer than the preceding three together, apically bearing some short sensilla. Mandibles ( Fig. 37 View Fig a-d, 38 a-d) slightly asymmetrical, regularly curved, apicalis with long pointed apical tooth, protruding over mesal brush, mesal brush well developed. Antennae ( Fig. 8 d View Fig ) with ten antennomeres scape distally subcarinate (securiform), distally bearing some setae, funicle short with pedicellus bent at about a right angle with respect to the longitudinal axis of scape, the remaining antennomeres of funicle very short, distinctly wider than long, antennal club three-jointed, joints hairy, relatively short, narrow; club small, about as long as wide although longer than the length of funicle.

Protibiae almost straight, outer edge regularly finely serrate, denticles short, ending with two distinct outer teeth distinctly more protruding than denticles; apical spur relatively long, sharp, very gently and feebly curved downward (sexually dimorphic: in males of G. fisheri sp. nov. strongly enlarged and blunt). Protarsi with first article about as long as or longer than the following three together, articles two and four slightly dilated distally, article five slightly longer than the former; each tarsomere, with the exception of the last one, ventrally bearing tufts of dense fine short setae, which, however, do not conceal the tarsal surface; at least in G. mayottensis sp. nov. some peg-like short setae are present on the first protarsomere mixedto the normally longer setae ( Fig. 7 c View Fig ). Claws small. Mesotibiae subrectangular, long, inner angle of apex with two apical spurs. Mesotarsi inserted near the inner angle of apical edge, about twice the length of apical edge of tibia; each tarsomere, with the exception of the last one, ventrally bearing a tuft of dense setae. Metatibiae triangular ( Fig. 9 m View Fig ), elongate, ending with two straight sharp fine paired spurs. Metatarsi almost as long as the apical edge of tibia; each tarsomere, with the exception of the last one, ventrally bearing a tuft of dense setae.

Wings: flightless (apterous or micropterous).

Sexual dimorphism: apex of clypeus elongate in the females (apical projection. The development of this apical projection varies depending on species, e.g., in G. pittinoi sp. nov. the female projection is extremely long and sharp, conversely in G. simplicipennis sp. nov. is much weaker) (e.g., Fig. 5 d View Fig , 48 d View Fig , 59 a View Fig ), apex of female protibiae ( Fig. 9 f View Fig ) elongate and with apical tooth more elongte than in males, forked, with the inner fork smaller, sharper, shorter and subject to wear, whereas in males ( Fig. 9 g View Fig ) the apical tooth is not forked, shorter and normally at a right angle; male mesotibiae with the inner apical spur bent inwards at a right angle ( Fig. 9 l View Fig ) while in the females both apical spurs are straight.

Male genitalia: spiculum gastrale variously shaped, fairly sclerotized; phallobase of aedeagus distinctly twist- ed; temones present; parameres strongly asymmetrical, short or long.

Female genitalia: bursa copulatrix without sclerites, spermatheca weakly sclerotized.

Etymology. Not indicated by Paulian, but clearly referring to Charles Prosper Goudot (see under Synarmostes tibialis for more information), the collector of the holotype of the type species. Gender masculine.

Distribution, habitat and bionomics. Endemic to Madagascar and the Comoros. In the Comoros records are limited to Moheli and Mayotte (see under Synarmostes for more details on Brian Fisher’s collecting in the archipelago). In Madagascar all species have been found in forested habitats, the majority of them (26 species) in rainforests and seven species in dry deciduous forests. The altitudinal range varies from sea level to 1325 m a.s.l. ( Goudotostes parvus sp. nov.). Other montane species are G. angelii sp. nov. (1200 m a.s.l.) G. elegans sp. nov. (1100 m a.s.l.), G. rafanomezantsoai sp. nov. (1200 m a.s.l.), G. ramamonjisoae sp. nov. (1300 m a.s.l.), and G. simplicipennis sp. nov. (1280 m a.s.l.). The majority of species have been collected between 100 and 500 meters above sea level. All species have been collected by sifting leaf litter (although the holotype of G. scabrosus was found in a termite nest). All species are flightless (apterous or micropterous). Preimaginal stages are unknown.

Remarks. With this revision, the genus Goudotostes increases its composition from one species to 33 species. The species here ascribed to the genus Goudotostes can be divided into at least two groups plus some incertae sedis species (i.e., species which do not form a group and cannot be assigned to any of the two groups above). The first group includes the type species G. scabrosus plus G. andreonei sp. nov., G. antsahabensis sp. nov, G. lapidisilvae sp. nov., G. litoralis sp. nov., G. montanellus sp. nov., G. orangeanus sp. nov., G. rasoamananae sp. nov., G. siccaesilvae sp. nov., G. similis sp. nov., G. trapeticollis sp. nov., G. electrimontis sp. nov. and G. insularis sp. nov., and is characterized by the combination of: a) short strongly asymmetrical parameres of aedeagus, b) monolobate anterior and posteri- or carinae, c) presence of a clypeal tubercle (absent or very weak in some species) and c) the sculpturing of elytral base having three to five tubercles or short carinae. The second group (hereinafter indicated as the laevis group of species) includes G. laevis sp. nov., G. masoalae sp. nov., G. parvus sp. nov., G. rafanomezantoai sp. nov., G. rajemisonae sp. nov., G. rakotonirinai sp. nov., G. ramamonjisonae sp. nov., and G. ranaivoi sp. nov. and is characterized by the combination of: a) small size, b) presence of a weakly raised fronto-clypeal tubercle, c) bilobate anterior and posterior carinae, d) elytral base with only two tubercles (parasutural and humeral tubercle), and e) left paramere with ventral margin invaginated ( Fig. 8 f View Fig ). This group seems to be restricted to the rainforests of the northern half of eastern Madagascar. As regards the remaining species, G. angelii sp. nov. and G. elegans sp. nov. are similar to each other and show an outer morphology and aedeagal morphology similar to the one found in the laevis group of species with some resemblance also with G. andohahelae sp. nov. due to the tubercles of elytral base and the posterior and anterior pronotal lobes. Goudotostes mayottensis sp. nov. and G. hirtellus sp. nov. show an aedeagal morphology similar to the one found in the scabrosus group of species, although their dorsal sculpturing is completely different. Goudotostes fisheri sp. nov., G. lokobensis sp. nov. and G. pittinoi sp. nov., group together due to the strong dorsal sculpturing and the strongly asymmetrical and relatively elongate parameres, which are quite peculiar and strongly diverge from the one found in the scabrosus group of species. Finally, G. rugatulus sp. nov., G. andohahelae sp. nov., G. simplicipennis sp. nov., G. phantasticus sp. nov. and G. randrianirinai sp. nov. form each one an isolate combination of external morphology and aedeagal features, which makes it difficult to assign them to any of the aforementioned groups. The diverse morphology found in the genus Goudotostes , resulting from this revision, chal- lenges the separation of the genus Cryptosphaeroides Ballerio, 2008 from Goudotostes . Similarities between Cryptosphaeroides and Goudotostes were already suggested by Ballerio (2008) due to the same basic morphology of mouthparts, the same shape and orientation of antennal pedicellus, the same pattern of sexual dimorphism and the strongly asymmetrical parameres. Then, the main differences between the two genera were found in the sculpturing of head, pronotum and elytra: Goudotostes, sensu Ballerio (2008) , was characterized by a glabrous and strongly sculptured dorsal body surface, with carinae, tubercles and deep, dense punctation and the pronotum with a prominently raised posterior margin, while Cryptosphaeroides was characterized by smooth dorsal surface with sparse large horseshoe-shaped punctures and erect setae and the pronotum evenly convex, without any raised posterior margin. With this revision, the only character separating the two genera is the strongly raised posterior margin of pronotum in Goudotostes . Only a cladistic analysis would reveal whether Cryptosphaeroides is not paraphyletic within Goudotostes or still forms a separate clade. Finally, like in the case of the genus Synarmostes , also for many species of Goudotostes , especially belonging to the laevis and scabrosus groups of species, to reach a reliable species-level identification, the examination of the aedeagus is mandatory.

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Coleoptera

Family

Hybosoridae

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