Two genera of Ceratocanthinae (Coleoptera Scarabaeoidea Hybosoridae), Melanophilharmostes Paulian, 1968 and Pseudopterorthochaetes Paulian, 1977, are evaluated through a cladistic morphological approach, resulting in the transfer of Melanophilharmostes demirei Paulian, 1977 to Pseudopterorthochaetes. All species are catalogued and all except two are keyed. Three new species are described: Melanophilharmostes poggii n. sp. from Annobon island (Ecuatorial Guinea), Pseudopterorthochaetes genierorum n. sp. and P. miomboincola n. sp., both from Mozambique. Melanophilharmostes zicsii (Paulian, 1968) is recorded for the first time for Burkina Faso, marking the northernmost record for African Ceratocanthinae. The records in Sudanian savannas and Miombo woodlands extend the ecological range of the subfamily.

The genera Melanophilharmostes Paulian, 1968 and Pseudopterorthochaetes Paulian, 1977 are two closely related genera of Ceratocanthinae (Coleoptera Scarabaeoidea Hybosoridae) (Ballerio & Grebennikov 2016) occurring in the Afrotropical region with respectively 17 and 6 species, thus representing an important portion of the diversity of the Ceratocanthinae recorded from continental Africa (56 species). Our knowledge of the diversity of these two genera is very poor, since their study is made difficult by many factors: few specimens are known for each species, some species are known by one sex only, some types are lost or inaccessible and, above all, the definition of these genera, as provided by Paulian (1977), is not satisfactory and has not been used consistently. Furthermore the examination of all known species demonstrates that, although the majority of species attributed to these two genera can be assigned to two discrete groups, there are species which show some intermediate characters. The first group includes dorsally setose species with male mesotibia ending with the inner apical spur bent inwards, roughly corresponding to Melanophilharmostes sensu Paulian, 1977; the second group includes dorsally glabrous species with male mesotibia ending with two straight apical spurs, roughly corresponding to Pseudopterorthochaetes sensu Paulian 1977. Intermediate characters between these two groups are observed in other species such as Melanophilharmostes ashantii (Paulian, 1974) and M. endroedyi (Paulian, 1968). The goals of the present work are therefore:

Methods and terminological conventions follow Ballerio & Grebennikov (2016) and references therein quoted. Label data are provided verbatim only for holotypes, with a slash to separate labels. In giving collecting data for holotypes, author’s comments are in square brackets, while depository collection acronyms are in parenthesis unless otherwise stated. Photographs were taken with a Canon Eos D5 MII with a macro objective MP 65 mm and Kenko extension tubes. Multi-layer images were then assembled using Zerene Stacker software and cleaned and unmasked using a photo processing software. SEMs were obtained with a Zeiss EVO 40 XVP Scanning Electron Microscope at MUSE (Trento, Italy) after gold coating.

To assess phylogenetic relationships of Melanophilharmostes and Pseudopterorthochaetes (= the ingroup), a parsimony phylogenetic analysis was perfomed using morphological characters of the adults. For this purpose, a matrix (Table 1) was created in Winclada (Nixon 2002) to include 33 terminals (the list of examined specimens besides types is in Table 3) and 18 parsimoniously informative characters (Table 2). The ingroup contained 25 valid species of both genera (including three newly described herein and excluding two from Angola). Eight additional species representing eight closely and more distantly related Ceratocanthinae genera as detected in Ballerio & Grebennikov (2016) were included as outgroups. All topologies were rooted on a representative of the most distantly related genus Ceratocanthus. Analysis was run with all characters as unweighted and unordered, except for character #14 (hind wing development), which was ordered. The matrix was spawned from Winclada to Hennig86 (Farris 1989) and the most parsimonious (=shortest) topologies were searched for by using the command <mhennig*> (constructing several trees and then applying branch-swapping to each) followed by the command <bb*> (branch-swapping to trees constructed by mhennig* and retaining the shortest trees up to the limits of the computer memory space).

The resulting shortest topologies were saved as a tree file by command <tsave>and then opened and further explored in Winclada by applying unambiguous character optimization to individual or by exporting the data to Nona (Goloboff 1999) for bootstrap analysis with 1000 replications.

This is not a complete revision of the involved genera, therefore in the catalogue all geographical records taken from literature have not been critically reviewed.

Abbreviations: EL maximum elytral length; EW maximum total elytral width; HL maximum head length; HW maximum head width; L length; PL maximum pronotal length at middle; PW maximum pronotal width at middle; W width.

The first Melanophilharmostes to be described was Philharmostes posthi Paulian, 1937 from Ivory Coast, described when the known continental African fauna of Ceratocanthinae scored only two species. In 1946 Paulian transferred P. posthi to the genus Pterorthochaetes Gestro, 1898 (a genus previously known only from the Indo-Malayan region) and three more species were added to the list (P. burgeoni Paulian, 1946 and P. puncticeps Paulian, 1946, both from the Democratic Republic of Congo, and P. elytratus from Cameroun). In 1955, Basilewsky added one more species, which, following Paulian’s new classification, was included in the genus Pterorthochaetes (P. donisi Basilevsky, 1955 from the Democratic Republic of Congo). Basilewsky added some more faunistic records for the other species. In 1968, Decelle added new faunistic data for Ivory Coast following Paulian’s classification. In the same year, thanks to the availability of more material, Paulian (1968) was able to decide that the placement within Pterorthochaetes was wrong and transferred all the African Pterorthochaetes back to the genus Philharmostes, at the same time isolating those species by erecting the subgenus Melanophilharmostes, which was then characterized by the shape of head, fore angles of pronotum rounded or truncate, body black and convex, and interrupted lateral carina of elytra. He included also four new species (P. zicsii, P. endroedyi, P. vincenti and P. ocellatus, all them described from the Republic of Congo) in the subgenus and designated P. zicsii as the type species of the subgenus. In the same year Petrovitz (1968) described one more species from the Democratic Republic of Congo, placed in the genus Philharmostes (P. palustris) withouth further discussion. In 1974, new findings from Ghana led Paulian to question his previous classification and to go back to the use of Pterorthochaetes for some species and of Philharmostes (without subgenus) for others. In particular, two new species were assigned to Pterorthochaetes (P. endroedyi and P. kumasii) and four new species were assigned to Philharmostes (P. ashantii, P. carinatus, P. bicarinatus and P. ghanae). In 1970, Martínez restricted the use of Pterorthochaetes to the Indo-Malayan and Australian species only and transferred P. puncticeps, P. posthi, P. burgeoni, P. elytratus (together with other species which do not belong to the taxa herein dealt with) to the genus Astaenomoechus Martínez & Pereira, a genus previously known only from the Neotropical region. He added new faunistic data for Angola and described three new species, two of them belonging to the taxa herein dealt with (A. carvalhoi and A. machadoi). Martínez’s paper sets a first obstacle to a complete revision of the involved taxa since the holotypes are deposited in the Museu do Dundo in Angola, currently inaccessible and whose collections’ fate is unknown. In 1977, in the framework of a complete revision of the continental African fauna of Ceratocanthinae, Paulian re-defined all known genera and distributed all the species above listed between two genera based mainly on shape and size of fore tibiae apical teeth and on the presence or absence of the carina delimiting pseudoepipleuron:

In an addenda to the revision the author provided a critical review of Martínez’s paper, although, not being able to examine the types, he simply suggested new combinations and synonymies. In the following years, only few data increased our knowledge of the two genera: a faunistic paper by Paulian (1979) added some data for Gabon, Ivory Coast, Liberia and the Democratic Republic of Congo; the description of Pseudopterorthochaetes cambeforti Paulian, 1981 from Ivory Coast; and the description of P. hystrix Paulian, 1991 from Madagascar. Ocampo & Ballerio (2006) listed all known species. Ballerio (2008) transferred P. hystrix to the Madagascan endemic genus Cryptosphaeroides Ballerio, 2008 (on the grounds of various morphological characters related to male genitalia, antennal morphology and sexual dimorphism). Ballerio et al. (2011) revised the Ceratocanthinae fauna of Cameroon and published some more faunistic data as well as some critical remarks on M. demirei. Finally, Ballerio & Grebennikov (2016) provided a phylogenetic framework to address relationships of Melanophilharmostes and Pseudopterorthochaetes, suggesting close relationship between these two genera, which clustered together in the same clade.

The genus Melanophilharmostes, as re-defined in the present paper, has its main center of distribution in the Guineo-Congolian rainforest block, occurring from Liberia in the West to the Kakamega forest in Kenya (Melanophilharmostes sp., Kenya, Kakamega Forest, light trap, 23.II.2002, ABCB) in the East. The northern borders of its distribution range are marked by two eccentric locations, i.e. Burkina Faso (see below under M. zicsii) and Ethiopia (see below under M. carinatus), while the southern borders reach Central Angola (Martínez 1970). There is also an insular record for Equatorial Guinea, Annobon island (see below under M. poggii n. sp.). Most records come from lowland and montane forests (up to 1600-1800 m, M. palustris; 1614 m, M. carinatus from Ethiopia), mostly gallery forests and rainforests, but we know at least one record from a Western Sudanian savanna in Burkina Faso (Fig. 2) (see below under M. zicsii).

The genus Pseudopterorthochaetes, as re-defined in the present paper, occupies a wide portion of the Guineo-Congolian rainforest block, from Ivory Coast in the West to Uganda in the East (Pseudopterorthochaetes sp., Uganda, Masindi District, Budongo Forest near Sonso, 1.45N, 31.35E, 19-30 June 1995, Thomas Wagner leg., ABCB). Several new eccentric records provided in this paper extend the distribution eastwards to Tanzania (a single female of a still undescribed species from Arusha-Chini, old collection, HNHM), and southwards to northern Mozambique (see below under P. genierorum n. sp. and P. miomboincola n. sp.) and to South Africa (a single female of a still undescribed species from Kruger National Park, CMN). Most records come from lowland and montane forests (up to 1550 m, P. endroedyi), mostly gallery forests and rainforests, but the two new species from Mozambique were collected in Miombo woodland (Fig. 3), i.e. from an ecologically quite different habitat, drier than rainforests and gallery forests and subject to relatively short seasonal rains.

Very little is known about the biology of these two genera. Most records have been done through sifting of leaf litter/humus or rotten wood in forest (Ballerio et al. 2011). Melanophilharmostes pseudoposthi was collected in a nest of Nasutitermes latifrons (Sjöstedt, 1896) (Paulian 1977). There are several collecting records of specimens attracted to light (Paulian 1977). There are some flightless species, some of them occurring through wide geographical areas, such as P. endroedyi occurring in Ghana, Ivory Coast, Cameroon and Gabon (Ballerio et al. 2011). Preimaginal stages are unknown. In Fig. 1, a map summarizes all known records for both the genera.

The phylogenetic analysis resulted in an overflow of 2807 shortest trees, each with the length of 54 steps, consistency index of 0.42 and retention index 0.79. A randomly selected tree with unambiguously optimized characters shown on internodes is depicted in Fig. 4, while the bootstrapping consensus topology is shown on Fig. 5. The ingroup was recovered as a moderately supported clade (bootstrap support 78%, Fig. 5). Both Pseudopterorthochaetes and Melanophilharmostes were recovered as monophyletic with moderate bootstrap support of 80% and 77%, respectively (except M. demirei, which was nested within Pseudopterorthochaetes). Although the two genera have a relatively uniform morphology, sharing several apomorphies and having at least two species which show some intermediate morphology between the two genera, the cladistic analysis did not provide enough evidence for synonymizing them. Therefore, also on the grounds that the examined material was fragmentary, I deem more prudent to leave them as separate genera, awaiting for more material and evidence before to synonymize them.

Philharmostes (Melanophilharmostes) Paulian, 1968

Type species (by original designation): Philharmostes (Melanophilharmostes) zicsii Paulian, 1968.

Adults display a relatively uniform morphology. Small Ceratocanthinae, reddish-brown, brown or black, with complete enrollment coaptations. Dorsum setose (40x). Dorsal ocular area always visible and genal canthus complete. Antennae with ten antennomeres. Mouthparts with mandibles with sharp apex (Figs 6 a, b), mesal brush narrow and well-developed, molar lobe very strong and labium with U-shaped emargination and thin palpi (Fig. 6 c), the last palpomere being longer than the preceding palpomeres. Epipharynx as in Fig. 6 d. Maxilla as in Figs 6 e, f. Wings (Fig. 11 j), when fully developed (there are some flightless species) have a distal fork at the end of vein CuA, and a distal expansion of vein MP 1+2. Dorsal sculpture consists of a variety of mostly shallow punctures and/or lines. Several species have a distinct pseudoepipleure delimited by a carina (Fig. 11 l), often interrupted or incomplete. Sexual dimorphism involves the following traits: the apex of clypeus, which in the females of some species is elongate (Fig. 11 b); the apex of female protibiae, which are elongate and have apical teeth more developed than in males; male mesotibiae which have the inner apical spur bent inwards at a right angle (Fig. 11 i). Spiculum gastrale and parameres have varied shapes and are useful for species-level differentiation (parameres can be flattened dorsally or laterally, glabrous or setose, some species have parameres which remain separated making visible a median lobe and are useful for species distinction). Besides male genitalia other characters useful for species differentiation are found in the punctation of head, pronotum and elytra, setation and shape of carina delimiting pseudoepipleure.

Philharmostes ashantii Paulian 1974: 206 (description, distribution); Melanophilharmostes ashantii: Paulian 1977 (key, description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, ♀ (HNHM), Ghana: / Ghana: Ashanti region, Ofinso, 259 m, N 6 54 – W 1 39, Dr. S. Endrody-Younga / Nr. 37, sifting, 18.VII.1965 / Holotypus 1974, Philharmostes ashanti Paulian /Holotype / Philharmostes ashanti n. sp. R. Paulian det. / Melanophilharmostes ashantii (Paulian) det. A. Ballerio ‘97 /

Distribution. Ghana (Paulian 1974).

Remarks. Very similar to M. endroedyi, of which it could be a synonym. The only noticeable difference is the punctation of dorsum which in M. ashantii is more impressed. Both species share the following combination of characters: dorsum setose, lack of fine punctation, presence of nine pairs of longitudinal fine lines on elytral distal third, lateral carina of elytra almost invisible and limited to distal third. The holotype of M. ashantii, the only known specimen of that species, has the clypeal apex elongate, probably a sexually dimorphic character of females, typically found in Pseudopterorthochaetes and not in Melanophilharmostes.

Philharmostes bicarinatus Paulian 1974: 208 (description, distribution); Melanophilharmostes bicarinatus: Paulian 1977 (key, description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, ♂ (HNHM), Ghana: / Ghana: Ashanti region, Bobiri forest res., 320 m, N 6 40 – W 1 15, Dr. S. Endrody-Younga / Nr. 142, sifting, 27.III.1966 / Holotypus 1974, Philharmostes bicarinuatus Paulian /Holotype / Philharmostes bicarinatus n. sp. R. Paulian det. / Melanophilharmostes bicarinatus (Paulian) det. A. Ballerio ‘97 /

Distribution. Ghana (Paulian 1974).

Remarks. Easily separable from all other known species by the interrupted elytral carina delimiting pseudoepipleuron. The carina starts at the medial third, it is interrupted at the beginning of the distal third, and then continues along the distal third (this third portion of carina is not aligned with the medial one) in combination with ocellate punctation on pronotum. The closest species to it is M. ocellatus, which however has a continuous carina (although slightly sinuate in correspondence of the interruption in M. bicarinatus). Aedeagus and spiculum gastrale as in Figs 12 g-h.

Pterorthochaetes burgeoni: Paulian 1946: 201 (description, distribution); Basilewsky 1955 (distribution); Decelle 1968 (distribution); Astaenomoechus burgeoni: Martínez 1970 (iconography, key, description, distribution); Melanophilharmostes burgeoni: Paulian 1977 (key, iconography, description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, sex unknown (MRAC), Democratic Republic of Congo: / Typus Pe. Burgeoni Paul. / Musée du Congo / Haut-Uele: Yebo Moto III-1926 L. Burgeon / R. det. E. 5093 / Pterorthochaetes burgeoni n. sp. R. Paulian det. / Melanophilharmostes burgeoni (R Paul) R. Paulian det. /

Distribution. Democratic Republic of Congo (Paulian 1946, 1977; Basilewsky 1955), Cameroon (Paulian 1977), Angola (?) (Martínez 1970), Ivory Coast (?) (Decelle 1968).

Remarks. The ocellate punctation of pronotal disc combined with sparse horseshoe-shaped punctures on elytra (mixed to very few simple punctures) and the shape of elytral carina delimiting pseudoepipleuron (limited to distal half) allow the identification of this species. The species most similar to it is M. donisi, which however has denser elytral punctation and a larger number of simple punctures mixed to horseshoe-shaped punctures on elytra.

Philharmostes carinatus Paulian 1974: 207 (description, distribution); Melanophilharmostes carinatus: Paulian 1977 (key, description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, ♂ (HNHM), Ghana: / Ghana: Ashanti region, Kumasi, 330 m, N 6 43 – W 1 36, Dr. S. Endrody-Younga / Nr. 39, sifting, 2.VII.1965 / Holotypus 1974, Philharmostes carinatus Paulian /Holotype / Philharmostes carinatus n. sp. R. Paulian det. / Melanophilharmostes carinatus (Paulian) ♂ /

Distribution. Ghana (Paulian 1974), Ethiopia (?) (see below under remarks).

Remarks. Melanophilharmostes carinatus is extremely similar to M. posthi and M. puncticeps. Thanks to the courtesy of V. V. Grebennikov (Ottawa, Canada) I have also observed a single female from Ethiopia (Bonga, 7.1954N 36.2812E, 1614 m, 10 November 2014, sift 29, local collector, CNC) which shows the same outer morphology of M. carinatus. Examination of more material and dissection of males is needed in order to better evaluate the status of these species and of the Ethiopian population. All them belong to the group of species characterized by the presence of fine punctation on pronotum and elytra and formed by M. zicsii, M. carinatus, M. ghanae, M. palustris, M. posthi, M. pseudoposthi, M. puncticeps and M. vincenti. Aedeagus and spiculum gastrale as in Figs 12 i-j.

Astaenomoechus carvalhoi: Martínez 1970: 20 (iconography, key, description, distribution); Melanophilharmostes carvalhoi: Paulian 1977 (discussion); Ocampo & Ballerio 2006 (listing).

Holotype, ♂ (according to original publication), in coll. Museo do Dundo, Dundo, Angola: “Angola, Moxico: detritos vegetales del suelo de la selva en galería de la margen izquierda del río Lumeje (11°40’S, 20°36’E), cerca de una picada que sale de la estación de Sandando, 17-I-1955 (A. De Barros Machado-coll.)”.

Distribution. Angola (Martínez 1970).

Remarks. Based on the original description, Paulian (1977) stressed close similarity with M. vincenti and I concur with him, however, since the fate of the collections of the Museo do Dundo in Angola is unknown and nobody has ever examined the types of M. carvalhoi and P. machadoi after the description, it is not possible to draw any final decision on the synonymy. The identity of M. carvalhoi and P. machadoi remains therefore doubtful.

Pterorthochaetes donisi: Basilewsky 1955:17 (description, distribution); Astaenomoechus donisi: Martínez 1970 (key, distribution); Melanophilharmostes donisi: Paulian 1977 (iconography, key, description, distribution); Paulian 1979 (distribution); Ocampo & Ballerio 2006 (listing).

Holotype, sex unknown (MRAC), Democratic Republic of Congo: / Holotypus / Yangambi. 1953 C. Donis z A / coll. R. Mayné com. Èt. Bois Congo R. 2467 / coll. – Mus. Congo Don R. Mayné / A. Jannssens det. 1954 Pterorthochaetes Donisi n. sp. / Pterorthochaetes Donisi n. sp. P. Basilewsky det., 1955 / Melanophilharmostes donisi (Basilew) det. R. Paulian 1976 /

Distribution. Democratic Republic of Congo (Basilewsky 1955; Paulian 1977, 1979).

Remarks. See under M. burgeoni.

Philharmostes (Melanophilharmostes) endroedyi: Paulian 1968: 93 (iconography, description, distribution); Melanophilharmostes endroedyi: Paulian 1977 (key, description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, ♂ (HNHM), Republic of Congo: / Soil-Zoological Exp., Congo-Brazzaville, Kindamba, Meya, Louolo river / 2.11.1963. No. 78, sifted litter, leg. Endrödy-Younga / Type / Holotypus Philharmostes endroedyi Paul. / Philharmostes endroedyi Paulian n. sp. / Type /

Distribution. Republic of Congo (Paulian 1968, 1977).

Remarks. See under M. ashantii. Aedeagus and spiculum gastrale as in Figs 12 e-f.

Philharmostes ghanae: Paulian 1974: 208 (description, distribution); Melanophilharmostes ghanae: Paulian 1977 (key, description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, ♀ (HNHM), Ghana: / Ghana: Ashanti region, Kumasi, 330 m, N 6 43 – W 1 36, Dr. S. Endrody-Younga / Nr. 39, sifting, 2.VII.1965 / Holotypus 1974, Philharmostes ghanae Paulian /Holotype / Philharmostes ghanae n. sp. R. Paulian det. / Melanophilharmostes ghanae (Paulian) ♀ /

Distribution. Ghana (Paulian 1974, 1977), Togo (Paulian 1977).

Remarks. Due to the sparse horseshoe-shaped elytral punctures, this species is similar to M. vincenti from which can be easily separated by the different punctation of elytral base. The correct type locality is Kumasi, the type locality reported in Paulian (1977), i.e. Bobiri, is wrong.

Philharmostes (Melanophilharmostes) ocellatus: Paulian 1968: 94 (iconography, description, distribution); Melanophilharmostes ocellatus: Paulian 1977 (iconography, key, description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, ♀ (HNHM), Republic of Congo: / Soil-Zoological Exp., Congo-Brazzaville, Bouenza, catarract / 30.11.1963. No. 308, sifted, in float, leg. Endrödy-Younga / Holotypus, Philharmostes ocellatus Paul. / Philharmostes ocellatus n. sp. / type / Melanophilharmostes ocellatus (Paulian), det. A. Ballerio ‘97 /

Distribution. Republic of Congo (Paulian 1968, 1977).

Remarks. See under M. bicarinatus.

Philharmostes palustris: Petrovitz 1968: 255 (description, distribution); Melanophilharmostes palustris: Paulian 1977 (key, description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, sex unknown (MRAC), Democratic Republic of Congo: / Holotypus / Biot. N. 101 Bord petit marais en foret (Mus. Roy. Afr. Centr. Tshuapa: Terr. Ikela, riv. Gombe, IX-1959 B. 101 N. leleup / Typus / Philharmostes palustris nov. 1963 Petrovitz / Philharmostes palustris Petr. Det. R. Paulian 1976/

Distribution. Democratic Republic of Congo (Petrovitz 1968; Paulian 1977).

Type series. Holotype, ♀ in coll. MCSN, Equatorial Guinea: / Is. Annobon 400-500 m. IV-V 1902 L. Fea / Melanophilharmostes poggii sp. n. A. Ballerio det. 2015 ♀ / Museo Civico di Genova /

Diagnosis. Melanophilharmostes poggii n. sp. is a small Melanophilharmostes which belongs to the group of species characterized by the presence of fine punctation on pronotum and elytra. It differs from all other species belonging to the aforesaid group by the following combination of characters: a) shiny dorsum, b) lateral elytral carina complete, c) presence of a longitudinal wide smooth elytral stria without any punctures, d) the lack of longitudinal rows of lines, which are replaced by irregular rows of horseshoe-shaped punctures and e) flightlessness.

Description. Size: HL = 0.63 mm; HW = 1.13 mm; PL = 1.10 mm; PW = 1,86 mm; EL = 2.13 mm; EW = 2.00 mm. Overall morphology as in generic diagnosis. Small Melanophilharmostes. Light brown, shiny, setate, fore margins of clypeus, sternum, tarsi and antennae reddish-brown. Flightless.

Head: interocular distance about ten times the maximum width of dorsal ocular area, frons smooth, clypeus completely covered by comma-shaped punctures with opening oriented forwards, each one having one or two simple punctures inside, sometimes bearing a long fine erect seta, fore margin with some irregular deep transverse lines. Pronotum: setate, setae relatively long, thin, erect, clavate. Margin complete, fore angles subtruncate, completely covered by long, transverse irregular shallow lines, sometimes anastomizing, mixed to dense simple fine punctures. Scutellum: covered by horseshoe-shaped punctures with opening backwards mixed to dense fine punctures.

Elytra: setate (setae erect, sparse, relatively long, thin, clavate), ovate, humeral callus indistinct, periscutellar area with at first some transverse lines and then large horseshoe-shaped punctures. Elytra covered by longitudinal irregular rows of large horseshoe-shaped punctures, mixed to sparse fine punctures, at middle a longitudinal smooth stria without any punctures. Lateral carina complete. Pseudoepipleura with some irregular longitudinally oriented straight shallow lines.

Flightless (brachypterous).

Sexual dimorphism: unknown. The female holotype however lacks any clypeal projection. Fore tibiae are as in other Melanophilharmostes females.

Etymology. Named after Roberto Poggi, former director of MCSN, who lent me the new species for study. Noun in the masculine genitive singular case.

Distribution and habitat. Annobon Island (Equatorial Guinea). According to a letter that Leonardo Fea sent to Raffaello Gestro dated 5 April 1902 (R. Poggi pers. comm.), when Fea collected the holotype the island was partly covered by degraded secondary lowland rainforest.

Remarks. This is the first insular record for Melanophilharmostes.

Pterorthochaetes posthi: Paulian 1937: 431 (key, description, distribution); Paulian 1946 (key, description, distribution), Decelle 1968 (distribution); Astaenomoechus posthi: Martínez 1970 (key, distribution); Melanophilharmostes posthi: Paulian 1977 (iconography, key, description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, ♀ (MNHN), Ivory Coast: / Museum Paris, Cote d’Ivoire, env. de Dimbokro, Capitaine Posth 1911/ Type / Pterorthochaetes posthi n. sp. Det. R. Paulian 1937/

Distribution. Ivory Coast (Paulian 1937, 1946, 1977; Decelle 1968), Liberia (Paulian 1979), Togo (Paulian 1977).

Remarks. See under M. carinatus.

Melanophilharmostes pseudoposthi: Paulian 1977: 297 (key, description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, ♀ (FMNH), Democratic Republic of Congo: / Camp Putnam, Epulu, Belg. Congo May 15: 1948 AE Emerson leg. / Host: Nasutitermes latifrons Siöst / Holotype / Melanophilharmostes pseudoposthi n. sp. R. Paulian det. /

Distribution. Democratic Republic of Congo (Paulian 1977).

Pterorthochaetes puncticeps: Paulian 1946: 200 (key, description, distribution); Basilewsky 1955 (distribution); Astaenomoechus puncticeps: Martínez 1970 (key, distribution); Melanophilharmostes puncticeps: Paulian 1977 (iconography, key, description, distribution); Paulian 1979 (distribution); Ocampo & Ballerio 2006 (listing).

Holotype, sex unknown (MRAC), Democratic Republic of Congo: / Holotypus / Musée du Congo Equateur: Flandria XI-1929 R. P. Hulstaert / R. det. C. 5093 / Pterorthochaetes puncticeps n. sp. R. Paulian det. / Melanophilharmostes puncticeps (R. Paul.) R. Paulian det. /

Distribution. Democratic Republic of Congo (Paulian 1946, 1977, 1979; Basilewsky 1955), Liberia (Paulian 1977).

Remarks. See under M. carinatus.

Pilharmostes pygmaeus: Petrovitz 1968: 256 (description, distribution); Melanophilharmostes pygmaeus: Paulian 1977 (key, description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, ♀ (MHNG), Democratic Republic of Congo: / Yangambi / Congo Belge / Typus / Philharmostes pygmaeus nov. 1968 Petrovitz / coll. Petrovitz /

Distribution. Democratic Republic of Congo (Petrovitz 1968; Paulian 1977).

Philharmostes (Melanophilharmostes) vincenti: Paulian 1968: 92 (iconography, description, distribution); Melanophilharmostes vincenti: Paulian 1977 (iconography, key, description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, ♀ (HNHM), Republic of Congo: / Soil-Zoological Exp., Congo-Brazzaville, Kindamba, Meya, Louolo river / 2.11.1963. No. 78, sifted litter, leg. Endrödy-Younga / Type / Holotypus Philharmostes vincenti Paul. / Philharmostes vincenti Paulian n. sp. / Melanophilharmostes vincenti (Paulian) ♀/

Distribution. Republic of Congo (Paulian 1968, 1977).

Remarks. See under M. ghanae. Aedeagus and spiculum gastrale as in Figs 12 k-l.

Philharmostes (Melanophilharmostes) zicsii: Paulian 1968: 91 (iconography, description, distribution); Melanophilharmostes zicsii: Paulian 1977 (iconography, key, description, distribution); Paulian 1979 (distribution); Ocampo & Ballerio 2006 (listing); Ballerio et al. 2011 (iconography, key, distribution).

Holotype, ♀ (HNHM), Republic of Congo: / Soil-Zoological Exp., Congo-Brazzaville, galery [sic] forest 20 km W from Loudima / 9.12.1963. No. 422, sifted fungous trunks, leg. Endrödy-Younga / Holotypus Philharmostes zicsii Paul. / Type / Philharmostes zicsii Paulian n. sp. / Melanophilharmostes zicsii (Paulian) ♂/

Distribution. Republic of Congo (Paulian 1968, 1977), Gabon (Paulian 1979), Cameroon (Paulian 1979; Ballerio et al. 2011). I also know two specimens recently collected in Burkina Faso, which are tentatively assigned to M. zicsii (new country record): they are 1 male and 1 female, labeled as follows: Burkina Faso: Comoé, Koflandé (village), 290m 10.14.42 N 004.27.50 W, 4 July 2006, “zone soudanienne, savane boisée, piége lumineux”, F. & S. Génier 2006-01 (FGIC).

Remarks. Aedeagus and spiculum gastrale as in Figs 12 m-o. The new record from Burkina Faso is particularly interesting: it marks the northernmost record of Ceratocanthinae for Africa and comes from an unusual habitat, i.e. the West Sudanian savanna, consisting mainly of woodlands (Fig. 2). All other known records of Melanophilharmostes zicsii comes from moister habitats, i. e. rainforests or gallery forests. The two specimens from Burkina Faso however differ from all other M. zicsii known to the author because of the sculpturing of elytra, which has larger and denser horsesehoe-shaped punctures, aligned on longitudinal rows, much denser and deeper transverse lines on pronotum and, overall, a less convex dorsum; conversely, male genitalia (parameres and spiculum gastrale) are almost identical to the ones of the holotype of M. zicsii from Congo and of the specimens from Cameroon. I therefore prudentially assign the Burkina Faso specimens to M. zicsii, awaiting for more evidence before to separate them from M. zicsii.

Type species (by original designation): Pterorthochaetes elytratus Paulian, 1946.

Adults display a relatively uniform morphology. Small Ceratocanthinae, dark-brown to black, with complete enrollment coaptations. Dorsum glabrous (40x). Dorsal ocular area always visible and genal canthus complete. Antennae with ten antennomeres. Mouthparts similar to the ones of Melanophilharmostes, with mandibles with sharp apex, mesal brush narrow and well-developed, molar lobe very strong and labium with U-shaped emargination and thin palpi, the last palpomere being longer than the preceding palpomeres. Wings (Fig. 11 j), when fully developed (there are some flightless species) have a distal fork at the end of vein CuA, and a distal expansion of vein MP 1+2. Dorsal sculpture consists of a variety of mostly shallow punctures and/or lines. All known species lack the carina delimiting the pseudoepipleure, which is therefore hardly discernable (Fig. 11 k). Sexual dimorphism involves the apex of clypeus which in the females of all known species is elongate (Fig. 11 b), female protibiae, which are elongate and have apical teeth more developed than in males, male mesotibiae which have two straight apical spurs (a very unusual feature). Spiculum gastrale and parameres have varied shapes and are useful for species differentiation. Besides male genitalia, other characters useful for species differentiation are found in the punctation of head, pronotum and elytra.

Pseudopterorthochaetes cambeforti: Paulian 1981: 329 (description, distribution); Ocampo & Ballerio 2006 (listing).

Holotype, ♂ (MNHN), Ivory Coast: / Lamto (Toumodi), Cote d’Ivoire, XI.1980, Y. Cambefort leg. / tamisage litière de forêt galerie / holotype / Pseudopterorthochaetes cambeforti n. sp. R. Paulian det. /

Distribution. Ivory Coast (Paulian 1981).

Remarks. This species is very similar to P. elytratus and P. kumasii. Only examination of more material will allow a better evaluation of the status of these three species since the holotypes of the other two species are females. Aedeagus and spiculum gastrale in Figs 12 a-b.

Pseudopterorthochaetes criberrimus: Paulian 1977 (iconography, key, description, distribution); Paulian 1979 (distribution); Ocampo & Ballerio 2006 (listing).

Holotypus, ♀ (MRAC), Democratic Republic of Congo: / Holotypus / Biot. N° 72 Foret marécageuse tete de source / Holotype / I.R.S.A.C. Mus. Congo Kwango: Feshi, source de la Kikoli III-1959 B. 72 Mme J. Leleup / Pseudopterorthochaetes criberrimus n. sp. R. Paulian det. /

Distribution. Democratic Republic of Congo (Paulian 1977), Gabon (Paulian 1979).

Remarks. Because of its dense and small elytral punctation, this species can be easily differentiated from all other known species of the genus.

Melanophilharmostes demirei: Paulian 1977: 294 (iconography, key, description, distribution); Ocampo & Ballerio 2006 (listing); Ballerio et al. 2011 (iconography, key, distribution, discussion).

Holotype, ♂ (MNHN), Cameroon: / Mt. Febé 29-VI-66 / Muséum Paris, Cameroun, B. de Miré / Holotype / Melanophilharmostes demiréi n. sp. R. Paulian det. /

Distribution. Cameroon (Paulian 1977; Ballerio et al. 2011).

Remarks. Ballerio et al. (2011) suggested that the correct placement of this species should have been within the genus Pseudopterorthochaetes, mainly on the grounds of sexual dimorphism. The cladistic analysis herein performed confirms this hypothesis and therefore the species is formally transferred to Pseudopterorthochaetes. Ballerio et al. (2011) remarked that there are slight differences between the two known populations (Mt. Kupé and Mt. Febé, both in Cameroon): the differences are mainly in the punctures of pronotum and elytra, which are larger and sparser in the Mt. Kupé population, while the aedeagus does not show appreciable differences. The lack of additional material from the type locality (only the holotype is known from there) makes it impossible to draw any conclusion about any possible specific separation of these two populations. The species more similar to M. demirei is M. endroedyi. Both species are flightless. Besides the characters indicated in the key, M. demirei has more parallel sides of elytra. Aedeagus as in Fig. 12 c.

Pterorthochaetes elytratus: Paulian 1946: 201 (key, description, distribution); Basilewsky 1955 (distribution); Astaenomoechus elytratus: Martínez 1970 (distribution); Pseudopterorthochaetes elytratus: Paulian 1977 (key, description, distribution); Paulian 1979 (distribution); Ocampo & Ballerio 2006 (listing); Ballerio et al. 2011 (key, distribution).

Holotypus, sex unknown (MRAC), Cameroon: / Musée du Congo Kamerun: Joko Coll. Clavareau / R. det. D 5093 / Pseudopterorthochaetes elytratus (Paul) R. Paulian det. / Pterorthochaetes elytratus n. sp. R. Paulian det. / Typus Pt. Elytratus Paul /

Distribution. Cameroon (Paulian 1946, 1977; Ballerio et al. 2011), Democratic Republic of Congo (Paulian 1946, 1977).

Remarks. See under P. cambeforti.

Pterorthochaetes endroedyi: Paulian 1974: 205 (description, distribution); Pseudopterorthochaetes endroedyi: Paulian 1977 (key, description, distribution); Paulian 1979 (distribution); Ocampo & Ballerio 2006 (listing); Ballerio et al. 2011 (iconography, key, distribution).

Holotype, ♂ (HNHM), Ghana: / Ghana, Central region, Kibi, 274 m, N 6 10 – W 0 34, Dr. S. Endrody-Younga / Nr. 351, sifting, 11.V.1969 / Holotypus 1974, Pterorthochaetes endroedyi Paulian / Pterorthochaetes endroedyi R. Paulian det. / Holotype / Pseudopterorthochaetes endroedyi (Paulian) det. A. Ballerio ‘97 /

Distribution. Ghana (Paulian 1974, 1977), Ivory Coast (Paulian 1979), Cameroon and Gabon (Ballerio et al. 2011).

Remarks. For morphological comparison see under P. demirei. Aedeagus as in Fig. 12 d.

Type series. Holotype, ♂ in coll. CMN, Mozambique: / Moz.: Cabo Delgado, Mareja (site 1), P. N. Quirimbas 100m 12°50’36”S 40°10’53”E 23.12.2012, degraded eastern Miombo woodlands, light trap, F. & S. Génier & M. Denja, 2012-05 / Pseudopterorthochaetes genierorum sp. n. A. Ballerio det. 2015 ♂/ [distended, glued on card, dissected, male genitalia glued on a separate card, same pin] Paratypes: 3 ♀♀ in coll. FGCI (2) and ABCB (1): Moz.: Cabo Delgado, Namaluco (site 6), P. N. Quirimbas 230m 12.17.40S 40.13.19E 23 december 2012, degraded eastern Miombo woodlands, light trap, F. & S. Génier & M. Denja, 2012-22.

Diagnosis. Pseudopterorthochaetes genierorum n sp. due to the presence of longitudinal lines on elytra can be confused only with P. miomboincola n. sp. from which however differs not only by the lack of deep wrinkles on pronotum but also because of the small size of horseshoe-shaped punctures on elytra and the much sparser simple and comma-shaped punctures between lines. The shape of aedeagus is also similar to the one of P. genierorum and possibly to the one of P. machadoi, which, however, according to the original description, has a completely different pattern of punctation on pronotum and elytra.

Description. Size: HL = 1.14 mm; HW = 1.40 mm; PL = 1.22 mm; PW = 2.71 mm; EL = 2.28 mm; EW = 2.37 mm. Overall morphology as in generic diagnosis. Large Pseudopterorthochaetes. Black, shiny, glabrous (40X), fore margins of clypeus, sternum, tarsi and antennae reddish-brown. Volant.

Head: interocular distance about ten times the maximum width of dorsal ocular area, disc with very fine spare punctation, sides with some deep larger and denser simple punctures, becoming comma-shaped distally, fore margin with some irregular deep transverse lines.

Pronotum: margin complete, fore angles subtruncate, surface, with the exception of sides and base, completely covered by sparse small comma-shaped punctures, with opening oriented forwards, some long transverse lines are present at each side, mainly basally, and on disc (but shorter than the ones at sides).

Scutellum: covered by relatively deep small comma-shaped punctures with opening backwards.

Elytra: humeral callus indistinct, proximal third with at first some transverse short shallow lines or comma-shaped punctures becoming small irregularly distributed horseshoe-shaped punctures distally, medial and distal third characterized by the presence of eight longitudinal pairs of lines (excluding the sutural one) with very rare simple punctures in the middle. Lateral carina absent. Pseudoepipleura with longitudinally oriented straight dense irregular lines.

Metathoracic wings fully developed.

Sexual dimorphism: male mesotibia ending with two straight apical spurs, female clypeus ending with a distinct long projection slightly bent upwards.

Male genitalia: aedeagus with parameres relatively long, dorsally flattened, as in Figs 10 a-b, spiculum gastrale with distinct manubrium, as in Fig. 10 c.

Etymology. Named after François and Simon Génier (Ottawa, Canada), collectors of the type series. Noun in the masculine genitive plural case.

Distribution and habitat. Known from the type locality only, the Quirimbas national park, a coastal protected area in northern Mozambique. The area is characterized by the presence of wide expanses of Miombo woodland (e.g. Fig. 3). The whole type series was collected at light.

Pterorthochaetes kumasii: Paulian 1974: 206 (description, distribution); Pseudopterorthochaetes kumasii: Paulian 1977 (key, description, distribution); Paulian 1979 (distribution); Ocampo & Ballerio 2006 (listing).

Holotypus, ♀ (HMNH), Ghana: / Ghana: Ashanti region, Bobiri forest res., 320 m, N 6 40 – W 1 15, Dr. S. Endrody-Younga / Nr. 142, sifting, 27.III.1966 / Holotypus 1974, Pterorthochaetes kumasii Paulian / Pterorthochaetes kumasii R. Paulian det. / Holotype / Pseudopterorthochaetes endroedyi (Paulian) det. A. Ballerio ‘97 /

Distribution. Ghana (Paulian 1974, 1977), Gabon (Paulian 1979).

Remarks. See under P. cambeforti.

Astaenomoechus machadoi: Martínez 1970: 31 (iconography, key, description, distribution); Pseudoptrorthochaetes machadoi: Paulian 1977 (discussion); Ocampo & Ballerio 2006 (listing).

Holotype (according to original publication), ♂ in coll. Museo do Dundo, Dundo, Angola: “Angola, Dundo: naciente del río Dundo (07°25’S, 20°47’E), en los detritos del suelo de la selva en galería, 16-III-1954 (A. De Barros Machado y Ed. Luna de Carvalho-coll.).”

Distribution. Angola (Martínez 1970).

Remarks. Based on the original description, Paulian (1977) suggested close similarity with P. elytratus. See also under M. carvalhoi.

Type series. Holotypus, ♂ in coll. CMN, Mozambique: / Mozambique: Cabo Delgado, Ravia (site 6), Parque Nacional das Quirimbas 380 m 12°31’02”S 39°18’38”E 3.1.2013, eastern Miombo woodlands, light trap, F. & S. Génier & M. Denja, 2013-35 / Pseudopterorthochaetes miomboincola sp. n, A. Ballerio det. 2015 ♂/ [distended, glued on card, dissected, male genitalia glued on a separate card, same pin]. Paratypes: 2 ♀♀ in coll. FGIC (1) and AB-CB (1): Mozambique: Cabo Delgado, Namaluco (site 6), P. N. Quirimbas 230 m 12.17.40S 40.13.19E 23 December 2012, degraded eastern Miombo woodlands, light trap, F. & S. Génier & M. Denja, 2012-22. 1 ♀ (FGIC): Mozambique: Cabo Delgado, Taratibu (site 7), P. N. Quirimbas, 310 m, 12.48.25S 39.42.20E, 10 January 2013, eastern Miombo woodlands, light trap prairie edge, F. & S. Génier & M.Denja, 2013-68.

Diagnosis. Pseudopterorthochaetes miomboincola n. sp. can be easily distinguished from all other known Pseudopterorthochaetes by the deeply wrinkled pronotum. The species closest to it is P. genierorum n. sp. with which it shares the presence of longitudinal lines on elytra, although in P. miomboincola these are denser, the simple punctures between lines are denser too. The shape of aedeagus is also similar to the one of P. genierorum and possibly to the one of P. machadoi, which, however, according to the original description, has a completely different pattern of punctation on pronotum and elytra.

Description. Size: HL = 1.00 mm; HW = 1.28 mm; PL = 1.14 mm; PW = 2.50 mm; EL = 2.22 mm; EW = 2.14 mm. Overall morphology as in generic diagnosis. Large Pseudopterorthochaetes. Black, shiny, glabrous (40X), fore margins of clypeus, sternum, tarsi and antennae reddish-brown. Volant.

Head: interocular distance about ten times the maximum width of dorsal ocular area, disc with very fine spare punctation, sides with deep larger and denser comma-shaped punctures centrifugally oriented, fore margin with some irregular deep transverse lines.

Pronotum: margin complete, fore angles subtruncate, completely covered by deep long, sometimes anastomizing, transverse lines, occupying most of the surface with the exception of sides and base. Surface between lines with very few sparse small shallow simple punctures.

Scutellum: covered by relatively deep wide comma-shaped punctures with opening backwards.

Elytra: humeral callus indistinct, proximal third with at first some transverse short lines or comma-shaped punctures becoming small irregularly distributed horseshoe-shaped punctures distally, medial and distal third characterized by the presence of eight longitudinal pairs of lines (excluding the sutural one) with simple punctures or very short comma-shaped punctures in the middle. Lateral carina absent. Pseudoepipleura with longitudinally oriented straight dense irregular lines.

Metathoracic wings fully developed.

Sexual dimorphism: male mesotibia ending with two straight apical spurs, female clypeus ending with a distinct long projection slightly bent upwards.

Male genitalia: aedeagus with parameres relatively long, dorsally flattened, as in Figs 10 d-e, spiculum gastrale with distinct manubrium, as in Fig. 10 f.

Etymology. Named after its peculiar habitat, the Miombo woodland (Miombo, the local name of Brachystegia spp., plus Latin noun “incola” meaning “dweller”). Noun in apposition (hence not subject to gender agreement).

Distribution and habitat. Known from the type locality only, the Quirimbas National Park in Northern Mozambique, for details see under P. genierorum sp. n.

Pseudopterorthochaetes machadoi and Melanophilharmostes carvalhoi are excluded from the key, due to lack of specimens available for comparison.