A revision of Syngalepsus Beier, with the description of two new species from the Central African Republic and Malawi (Mantodea, Tarachodidae)

Abstract The praying mantis subgenus Syngalepsus Beier, 1954 occurs in sub-Saharan region and represents the least diverse subgenus of Galepsus in Africa (Ehrmann 2002). All species included within the subgenus Syngalepsus are comprehensively revised with a distribution stretching from North of Congo Basin to South Africa. Two new species of Galepsus (Syngalepsus) Beier, 1954 (Mantodea, Tarachodidae) from the Central African Republic (CAR) and Malawi are described. Among several Galepsus specimens collected during scientific expeditions of SANGHA, Biodiversité en Terre Pygmée, in CAR’s Dzanga-Ndoki National Park, a specimen differed by genitalia conformation and other morphological characters. Two specimens collected in Malawi proved to differ by genitalia morphology. Galepsus (Syngalepsus) buchetisp. n. and Galepsus (Syngalepsus) dudleyisp. n. are described. An identification key for the six species of the subgenus is provided.


Introduction
Subgenus Syngalepsus was erected by Max Beier in 1954 to organise some species from the genus Galepsus Stål, 1876: G. (S.) bipunctatus described in 1931 and G. (S.) denigratus Beier 1954. Beier also described G. (S.) birkenmeierae in 1969. Kaltenbach described a fourth species, G. (S.) beieri, from South Africa in 1996. In total, four species are currently known (Ehrmann 2002, Otte andSpearman 2005). All species of the subgenus have the vertex, straight or slightly arched, without bump or incision near the eyes and a right phallomere with the posterior apical region rounded and toothless, unlike Onychogalepsus. The prosternum has two circular black spots, more or less covered by a black patch. Three of the species are present in Southern Africa: South Africa (Natal) for G. (S.) beieri Kaltenbach (1996: 233); Mozambique and South Africa for G. (S.) bipunctatus Beier (1931: 3) (Type species) and Malawi for G. (S.) birkenmeierae Beier (1969b: 201). The fourth species, G. (S.) denigratus Beier (1954: 19), was collected in Angola (Beier 1969a), Gabon, Republic of Congo, Democratic Republic of Congo and Uganda (Beier 1957, Roy 1968. The discovery of a specimen in CAR that seems close to Syngalepsus led me to gather all the existing information concerning this subgenus (Moulin et al. 2017). The old descriptions have been revised and the figures of the genitalia grouped. Examination of its morphology and that of other specimens preserved in the collection has highlighted a second new species, native to Malawi. This paper aims to provide additional knowledge about Mantodea from Africa. Here we provide details about morphology and distribution of known species of the subgenus Syngalepsus and describe two new species.

Taxonomy
After the examination of several specimens of Galepsus genus in the Muséum national d'Histoire naturelle, Paris (MNHN) and a bibliographic survey (Beier 1931(Beier , 1954(Beier , 1957(Beier , 1969a(Beier , 1969bRoy 1968;Kaltenbach 1996Kaltenbach , 1998Ehrmann 2002;Otte and Spearman 2005), the genus Galepsus (Syngalepsus) appears to include six species. Two of these are unknown and are described in this subgenus revision. The genitalia were illustrated from types for all existing species, which served as the central distinguishing feature for the treated species.

Materials and methods
During the scientific expedition named "SANGHA, Biodiversité en terre Pygmée" (2012) in CAR, some Galepsus individuals were collected. Two species were identified: G. (Galepsus) globiceps Beier, 1942 and G. (Galepsus) laticeps Werner, 1907. But one specimen was atypical of the others and its genital characters were similar to those of Syngalepsus. In my comparisons to collection specimens, two undetermined specimens from Malawy were found in the material of the Muséum national d'Histoire naturelle, Paris, France. The specimens exhibit the diagnostic characters of the subgenus Syngalepsus (sensu Beier 1954) and differ from the RCA specimen and the four known species.
The specimens were photographed with a Dynax 5D Konica Minolta camera and a Leica MC 120 HD camera mounted on a S8APO Leica stereomicroscope. The freeware Combine ZP (Hadley 2008) was used to process images.
Taxonomy relied on Mantodea Species File website (Otte et al. 2018) as well as recent works on praying mantis molecular phylogenetics (Svenson and Whiting 2009, Wieland 2013, Roy 2014, Svenson et al. 2015. All morphological descriptions and measurements (in millimeters) refer to the material studied here unless explicitly stated otherwise. Terminology follows Brannoch et al. (2017) and Shcherbakov (2017). A total of 12 measurement classes were captured including: 1 Body length = length of body from central ocelli to posterior tip of wing or abdomen (intraspecifically variable measurement, primarily for general size estimation). 2 Forewing length = from proximal margin of axillary sclerites to distal tip of the discoidal region. 3 Pronotum length = from anterior margin to posterior margin. 4 Pronotum width = from lateral margins at the widest point, the supra-coxal bulge. 5 Head width = from lateral margins of the eyes at widest point. 6 Prothoracic coxae length = from proximal margin abutting pronotum to trochanter. 7 Prothoracic femur length = from proximal margin abutting trochanter to distal margin of genicular lobe. 8 Prothoracic tibiae length = from distal margin of genicular lobe to distal terminal spur. 9 Prothoracic femur width = at the widest point. 10 Anteroventral femoral spine count = all inner marginal ridge spines but excluding the genicular spine. 11 Anteroventral tibial spine count = all inner marginal ridge spines but excluding the distal terminal spur. 12 Posteroventral tibial spine count = all outer marginal ridge spines but excluding the distal terminal spur.
A map was created in QGIS 3.0.0. with administrative areas from GADM, the database of Global Administrative Areas website (https://gadm.org/).
As part of the revision of Syngalepsus, the descriptions of the four known species are repeated here, according to the original texts, translated into English. The questioning of the identity of the female allotype referred to G. (S.) denigratus by Beier between 1957 and1969 is re-evaluated here with the examination of two females, conserved at the MNHN, from the Republic of the Congo. The genitalia illustrated in the original documents are reproduced here (Figs. 1, 2). Beier, 1931 Figures 1A, 6A, 7A, 8 Galepsus bipunctatus : Beier 1931: 3;Beier 1954: 19;Kaltenbach 1996: 233;Kaltenbach 1998: 40;Ehrmann 2002: 154;Otte and Spearman 2005: 336. Holotype. Male deposited in ZMUH Hamburg; Type locality: Quilimane, Mozambique. Paratype: Male deposited in NHM Wien. Diagnosis. Pronotum with a blackened median line. Right phallomere without a process at the apex of the main lobe (fda); apical process on left phallomere (paa) rounded at the apex, having a small lateral process unlike other species.
The female of this species, of which there is one specimen (allotype) collected at Kasai, Shamba, was previously unknown. It is dorsally almost uniformly brown, only the vertex is blackened and metazone carries two large black spots. Dorsal margin transverse, lower frons almost straight. Vertex flat arched. Forewings blackish-reddish brown, blackened ventrally along anterior radius. Hindwings mostly blackish, with reddish brown edges. Prosternum as in the male. Forecoxae with blackish pre-apical bandage. Trochanter mostly black. Forefemora half black in ventral face, with black thorns. Fore tibiae half brownish, tarsus blackened. Meso-and metathoracic legs lateral and dorsal brown dotted. Supra-anal plate wide, triangular and keeled. -Body length 27 mm; width of head 3,3 mm; pronotum length 7 mm; width of pronotum 2,7 mm; forewings length 4 mm. It is very similar in colour to the female of Galepsus (Onychogalepsus) femoratus Giglio-Tos collected from Bechuana land (Botswana), but differs from it in terms of its bright fore tibiae and body mass, above all by a slimmer pronotum and relatively shorter forewings." The female symbol in the description of Beier, in 1957, is a mistake. He wanted to indicate the measurements for the males.
The species is new to Angola. It was previously known only from the Congo." Note: The female described in 1957 from Shamba in Kasai (Republic of the Congo) is confirmed to be G. (S.) denigratus, after examination.
Redescription of female. Fine body, brown, length 33.0 mm from head to apex of abdomen. Wings reduced, brown or more or less bicoloured according to examined specimens.
Head. Vertex slightly convex, brownish at the top. Parietal sulcus well marked. A brown-black spot on the vertex, against the parietal sulcus, near the juxtaocular bulge. Lower frons almost square, slightly wider than long. Eyes slightly prominent on the side, almost square head, front view. Antennae broken in both examined females. Maxillary palps with the base of the last segment brown-black.
Thorax. Pronotum with prozone almost as wide as metazone. Two small depressions, stained brown, on each side of the midline of the metazone, behind the supracoxal sulcus. Two parallel black spots in the middle of the metazone; closer to the edge than the median line. Two small black-brown bands, parallel to the median line, near the posterior border. Margin of pronotum serrated along its entire length. Prosternum with a large black-brown spot, not reaching the edges of the pronotum. Two depressions characteristic of the subgenus, not very visible because of this dark spot.
Fore legs. Femora with 12 anteroventral femoral spines, 4 posteroventral femoral spines, 4 discoidal spines; tibiae with 11-12 anteroventral tibial spines, 11-12 posteroventral tibial spines. Coxae with a big black-brown spot at the apex, in the posterior margin and a smaller one in a more anterior position. Trochanter with a smaller, blackbrown spot, very close to the junction with the femora. Anteroventral face of femora with several dark brown spots of various sizes and shapes forming a band along the ventral half. Dark brown spots at the base of the big spines. Apex of anteroventral femoral spines dark brown. Tibiae with a darker longitudinal line inside. Meso-and metathoracic legs stained with dark-brown little spots as forelegs. Wings. Forewings and hindwings, short, not reaching the beginning of the 1 st abdominal segment. Forewings light brownish red, with a dark spot taking all the discoidal area. Costal area of the same colour as the first part of the discoidal area. Hind wings brownish. Abdomen. Stained with brown markings. Triangular supra-anal plate, as wide as the abdomen. Cerci relatively long, 4.5 mm, flattened, the last two distal cercomeri longer than wide.
Measurements (mm). Body length 30.2-33.0, pronotum length 7.2-8.2, fore wings length 4.1-5.0, fore coxae length 4.0-4.1, fore femora length 4.9-5.4, fore tibiae length 3.4-3.8; width of pronotum 2.6-2.7, width of head 3.4-3.9, width of fore femora 1.4-1.6.  Translation. "Galepsus (Syngalepsus) birkenmeierae n. sp. ♂. Relatively small. General colour yellowish brown, dark brown speckled. Head relatively wide, wider than pronotum. Vertex almost straight. Eyes relatively bulging laterally, dorsally rounded, without notching vertex. Lower frons wider than high. Base of antennae (flagellum) brilliant, gradually becoming darker towards the apex. Pronotum almost 3 times longer than broad, dark brown spotted or with a dark median line, metazone slightly narrower than prozone. Prosternum with a large oval black spot, which occasionally reveals the paired black spots characteristic of the subgenus. Forewings not completely covering the abdomen, just as Alae, slightly smoked. Costal field bright. Discoidal field with brown veins, quite rough, crossing anterior cubitus veins. Media and radius veins very clear and therefore interrupted. Forelegs light brown on ventral surface, with a little black spot, laterally marbled brown, trochanter with three black spots. Fore coxae almost reaching the posterior border of the prosternum. Meso-and metathoracic legs speckled brown. Cerci flat. Genitalia: ventral phallomere with short and more medial end claw, smaller subapical tooth (pda) and widely rounded corner pointing to the right. Ventral phallomere of rectangular general shape. Right phallomere, blunt, slightly thickened and with strong bristles at the apex; ventral plate (pia) with a series of granules, ventral process (pva) with a blunt basal tooth and an apical lobe. Apical process (paa) of left phallomere, slender, with the terminal part in the shape of an obtuse fork; posterior process of phalloid apophysis (apa) smooth, pointed, with a large hairy area.
Diagnosis. Close to Galepsus (Syngalepsus) denigratus Beier and even more to Galepsus (Syngalepsus) birkenmeierae Beier with the large black patch on prosternum but different by genitalia conformation. Head wider than the pronotum; vertex slightly convex; prosternum with a large black patch.
Description male. Fine body, brown, length 32.7 mm from head to subgenital plate. Hind wings with venation brown. Head. Vertex straight or slightly convex with the region between the parietal sutures and the eyes convex. Two black spots near of the eyes, on the posterior face of the head. Lower frons transverse. Maxillary and labial palps with base of penultimate and last segment with black patch. Below of the last segment black. Antennae brown-black. Thorax. Pronotum. Smooth lateral margins. Surface smooth. Prozone broader than metazone. Pronotum 2.8 times longer than broad. Covered by small dark-brown spots, diffuses. Prosternum with a large black patch covering the posterior ¾ of the surface. Place of the spot within the other species visible. Forelegs. Femora with 12 anteroventral spines, four posteroventral spines, four discoidal spines; tibiae with eleven anteroventral spines. ten-eleven posteroventral spines. Legs globally of same colour of pronotum. Coxae without coloured patch. Trochanter with a black spot. Base of femora with a black patch, in extension of the one of trochanter. Tibiae with an elongated black patch at base of anteroventral tibial spines, from the 3 rd to 11 th . All tarsomere black below. Meso-and metathoracic legs stained with dark-brown. All tarsomere black at the distal end. Wings. Forewings and hindwings are uniformly translucent pale brown with brown veins. Reaching tip of abdomen. Abdomen. Flattened. Supra-anal plate: two times wider than length, distal margin rounded, pubescent. Cerci: relatively longs, flattened, the last three distal cercomeri longer than wide. Subgenital plate: pubescent, posterior edge almost straight. Styles: long, relatively thin, hairy.
Genitalia. Right phallomere with ventral process (pva) and ventral plate (pia) sclerotised; posterior process of phalloid apophysis (apa) long, with sclerotised denticles, ventral plate (pia) sclerotised with a rectangular form in anterior part and a hairs tuft at the posterior part. Apex of right phallomere rectangular covered on apex and left side with thick bristles. Ventral phallomere elongated, with a tooth (pda) on posterior margin, distant of the left margin. Left phallomere with a posterior process of phalloid apophysis (apa) pointed shorter than apical process (paa) and sclerotised at apex. Apical process (paa) large, hammer form at apex. Thick bristles on posterior process of phalloid apophysis (apa) apex and main lobe (fda) of the right phallomere apex.
Female unknown. Measurements (mm). Holotype: body length 32.7, pronotum length 7.1, fore wings length 23.2, fore coxae length 4.3, fore femora length 4.9, fore tibiae length 2.9; width of pronotum 2.5, width of head 4.3, width of fore femora 1.5.  Diagnosis. Close to Galepsus (Syngalepsus) beieri Kaltenbach and Galepsus (Syngalepsus) bipunctatus Beier with the two black spots on prosternum but different by genitalia conformation: ventral process (pva), of right phallomere, end with several pointy teeth; left phallomere with apical process (paa) without a bump that gives a spoon or mallet appearance. Head as wide as the pronotum; vertex straight; prosternum with two black spots.

Galepsus
Description male. Fine body, light brown, length 32-34 mm from head to subgenital plate. Hindwings with venation light brown. Head. Slightly wider than pronotum. Vertex: straight, slightly convex. Frontal shield transversal, wider than high. Labial palps with the last two segments stained black at the base on both segments, and below for the apical segment. Maxillary palps broken. Antennae light brown. Thorax. Pronotum, three times longer than its largest width, prozone broader than metazone, lateral margins and surface smooth; median line of the pronotum blackened. Prosternum, same colour of the pronotum, with two black spots closed to centre of prosternum, on both sides of the median line. Forelegs. Coxae without coloured patch, two-three very small tubercles on the anteroventral border. The dark spot on the left trochanter is not a spot of colour on the cuticle but blackened material inside the tegument. Femora: with 11-12 anteroventral spines, four discoidal spines, four posteroventral spines, all spines with black-brown  apex. Tibiae: with 11-12 anteroventral spines, eleven posteroventral spines, tarsus with blackened underside. Wings. Forewings and hindwings are uniformly translucent pale brown with light-brown veins. Reaching tip of abdomen.
Abdomen. Flattened. Supraanal plate: almost wider than length, distal margin rounded, pubescent. Cerci: relatively longs, flattened, the last two cercomeri longer than wide. Subgenital plate: triangular shape, pubescent. Styles: relatively short, thick, hairy. Genitalia. Right phallomere with ventral process (pva) and ventral plate (pia) sclerotised; ventral process (pva) margin with large denticles. Apex of the right phallomere rounded, covered on apex and left side with thick bristles; hairs tuft at the posterior part. Ventral phallomere large, with a sharp process on posterior margin, towards the right. Left phallomere with a posterior process of phalloid apophysis (apa) long and sharp, slightly sclerotised at apex. Apical process (paa) long and round at apex, covered at apex by thick bristles; left margin of left phallomere covered by thick bristles.

Discussion
The present study led to the description of two new species, Galepsus (Syngalepsus) bucheti sp. n. and Galepus (Syngalepsus) dudleyi sp. n. Males are easily identifiable using the characters of both habitus and male genitalia. However, females are unknown, which is common for mantises because studies with light trap sampling fail to attract females. Here, only females of G. (S.) denigratus are known (Beier 1969a). They are brachypterous and likely live on tree trunks and branches. Sweep nets and visual searching should be used more broadly in Mantodea studies to ensure more females can be collected. Furthermore, this method will also reveal mantises' micro-habitat preferences. In CAR, no female of G. (S.) bucheti sp. n. were collected, although all known modern sampling techniques were implemented (sweeping, beating, tree climbing, light trap). With brachypterous females it a likely the distribution of each species could be restricted to small geographic ranges, which could indicate many species remain to be discovered in remote areas of Africa and other parts of the world.