Philharmostes Kolbe, 1895

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 : 246-248

Genus Philharmostes Kolbe, 1895

Philharmostes Kolbe 1895: 344 (description and distribution); Fairmaire 1900: 471 (type species designation); Alluaud 1900: 245 (catalogue of Madagascan species); Arrow 1912: 45 (catalogue); Paulian 1936: 131 (catalogue, key and iconography); Boucomont 1937: 278 (key, description and distribution); Paulian 1979: 60 (key, description, distribution, iconography); Ballerio 2006: 299 (sexual dimorphism); Ocampo & Ballerio 2006: 190 (listing); Ballerio & Grebennikov, 2016 (phylogeny); Grebennikov, 2019 (phylogeny).

Type species (subsequent designation by Fairmaire, 1899): Philharmostes aeneoviridis Kolbe, 1895 .

Taxonomic history. The genus Philharmostes was established by Kolbe (1895) to accommodate two new species: Philharmostes aeneoviridis from Madagascar and Philharmostes integer from the Usambara mountains ( Tanzania). He characterized the genus by the number of antennomeres (nine), the rounded pronotal posterior angles, the absence of genae, the flattened metatibiae, the capability to form a perfect ball once rolled up and the presence of an elytral lateral carina. None of the aforementioned characters is among the autoapomorphies which currently define the genus (see Ballerio & Grebennikov 2016). In the year 1900 Fairmaire designated P. aeneoviridis as the type species, re-defined the genus, transferred the Madagascan Synarmostes perroti Was- mann, 1897 Synarmostes perrieri Fairmaire, 1898 and Synarmostes latericostatus Fairmaire, 1885 to the genus Philharmostes and described two new species from Madagascar: Philharmostes cupreolus and Philharmostes obscurus . Alluaud (1900) listed all the aforesaid species and transferred also Synarmostes basicollis Fairmaire, 1897 to the genus Philharmostes . Shortly after, Fairmaire (1903a; 1903b) described four more Madagascan species: Philharmostes corruscus , P. pilula , P. convexifrons and P. cribrarius . Arrow (1912) listed eleven species ( P. aeneoviridis , P. basicollis , P. convexifrons , P. corruscus , P. cribrarius , P. integer , P. latericostatus , P. obscuroaeneus , P. obscurus , P. perrieri and P. pilula ). In 1936 Paulian provided the first identification key for Madagascar (listing P. perrieri , P. latericostatus , P. obscuroaeneus , P. obscurus , P. cribrarius , P. aeneoviridis , P. basicollis , P. corruscus , P. convexifrons , P. pilula and P. cupreolus ). He mistakenly designated P. latericostatus as the type species. Almost contemporarily Boucomont (1937) provided another identification key and described three more new taxa ( P. bicolor , and P. latericostatus var. olsoufieffi ). After that date, similarly to what happened with the genus Synarmostes , a number of heterogeneous species from Africa, Asia and America were added to Philharmostes by various authors without a sound taxonomic basis (e. g., Basilewsky 1955; Petrovitz 1968, who, however, described also a true Madagascan Philharmostes , i.e., P. boucomonti ). Martínez & Pereira (1959) and Paulian (1977; 1978b) cleaned the matter up and circumscribed the genus Philharmostes to some species from Africa and Madagascar. With regard to Madagascar, Paulian (1979) provided the last treatment by recognizing 10 of the previously described species ( P. aeneoviridis , P. basicollis , P. cupreolus , P. perrieri , P. bicolor , P. corruscus , P. cribrarius , P. boucomonti , P. latericostatus , and P. olsoufieffi ) and by adding four more new species, i e., P. descarpentriesi Paulian, 1979 , P. criberrimus Paulian, 1979 , P. pseudobasicollis Paulian, 1979 , and P. vadonianus Paulian, 1979 . Paulian’s treatment has remained unchanged till 2008, when Ballerio transfrred P. perrieri to the genus Pseudosynarmostes . Ballerio (2000) placed the genus Philharmostes within the “ Philharmostes group of genera”, a group of genera from Afrotroipical region ( Carinophilharmostes Paulian, 1968 , Callophilharmostes Paulian, 1968 , Chaetophilharmostes Paulian, 1977 , Baloghianestes Paulian, 1968 , Petrovitzostes Paulian, 1977 , and Cryptophilharmostes Ballerio, 2000 ), whose monophyly was later cofirmed by Ballerio & Grebennikov (2016), using adult morphology, and by Grebennikov (2019), using molecular data. The latter, however, highlighted the presence of multiple paraphilies within the group of genera.

Diagnosis (valid only for the species dealt with in the present revision). The genus can be identified by the following combination of characters: 1) enrollment coaptations complete, 2) protibiae arquate, 3) genal canthus complete (fused with the occipital area of head), 4) dorsal ocular area small, 5) elytra with a lateral carina (sometimes interrupted more times), delimitating an inferior area (pseudoepipleure).

Distribution and habitat (valid only for the species dealt with in the present revision). All the new species here described have been found in forested habitats, all them in rainforests. Philharmostes centralis sp. nov. is currently the only Ceratocanthinae recorded from the Ambohitantely forest on the central highlands. The altitudinal range varies from sea level to 1400 m a.s.l. ( P. centralis p. nov.). Other montane species are P. schweikerti sp. nov. (940- 1250 m a.s.l.) P. metallicus sp. nov. (1200 m a.s.l.), and P. indri sp. nov. (1080 m a.s.l.), while P. nelsoni sp. nov. displays a wide altitudinal range, spanning from 200 to 900 m a. s. l.. All species have been collected by sifting leaf litter. One specimen of P. nelsoni has been collected in a nest of Nasutitermes (Andohahela, Managortry forest). All species are flightless (apterous or micropterous). Preimaginal stages are unknown.

Remarks. Madagascar is the main centre of diversity of the genus Philharmostes with 26 recorded species (out of a total of 44 described species), all them endemic. The other species of Philharmostes occur in three areas of continental Africa: the Guineo-Congolian rainforest block (7 species), the Eastern Arc rainforests and coastal rainforests of Tanzania and Kenya (8 species) and South Africa (3 species). The genus seems to be absent from the Comorian archipelago. The remaining Madagascan Philharmostes species not dealt with herein will be the subject of a subsequent paper. They all are volant, with a large dorsal ocular area and often have bright dorsal metallic colours (green, bronze, red).