Research Article Journal of Orthoptera Research 2024, 33(1): 127-146 New species of Pferoftiltus from Cameroon, and a new genus, Parapferotiltus (Orthoptera, Acrididae, Oxyinae) CHARLY OumAROU NcGoute!, C.H.F. ROWELL? 1 Zoology Unit, Department of Biology of Animal Organisms, Faculty of Science, University of Douala, Douala, Cameroon. 2 Zoologisches Institut der Universitat Basel, Basel, Switzerland. Corresponding author: Charly Oumarou Ngoute (coumaroungoute@yahoo.fr) Academic editor: Daniel Petit | Received 7 October 2022 | Accepted 2 February 2023 | Published 7 May 2024 https://zoobank. org/ED93B 11 2-FOB3-46D6-B536-62F099689C61 Citation: Oumarou Ngoute C, Rowell CHF (2024) New species of Pterotiltus from Cameroon, and a new genus, Parapterotiltus (Orthoptera, Acrididae, Oxyinae). Journal of Orthoptera Research 33(1): 127-146. https://doi.org/10.3897/jor.33.96043 Abstract The genus Pterotiltus Karsch, 1893 currently includes 12 described spe- cies from the equatorial forests of West and Central Africa. Here, we describe two new species from lowland forests of Southern Cameroon: Pterotiltus ngoylaensis sp. nov., and Pterotiltus campoensis sp. nov. We further describe two new species and the previously undescribed female of Pterotiltus mini- mus Ramme, 1929 from the highland areas of West Cameroon. These high- land species share a suite of characters that distinguish them from Pterotiltus; these characters include structures of both the male and female reproductive systems, tegmina not completely regressed, and generally smaller size. To accommodate these differences between the species, we erect the new genus Parapterotiltus gen. nov. with P. minimus as the type species and include in it Parapterotiltus minimoides sp. nov. and Parapterotiltus bamboutos sp. nov. Keywords Caelifera, Cameroon, grasshoppers, taxonomy, tropical forest Introduction The genus Pterotiltus Karsch, 1893 is an Afro-tropical taxon. To- gether with 37 other genera from Africa, Asia, and Australia, it is placed in the subfamily Oxyinae Brunner von Wattenwyl, 1893. The Oxyinae are currently divided into two recognized tribes: Oxyini Brunner von Wattenwyl, 1893 and Praxibulini Rehn, 1957 (Cigli- ano et al. 2022). Pterotiltus, and several other wet-forest oxyine gen- era of similar habitus [the African genera Badistica Karsch, 1891, Caryanda Stal, 1878, Cylindrotiltus Ramme, 1929, Digentia Stal, 1878, and Gerista Bolivar, 1905, and several Asian genera (Hollis 1975, Rowell 2005)] are currently excluded from both these tribes. However, Pterotiltus and the other excluded African genera seem to be close to the Oxyini, which includes other African grasshoppers, particularly Oxya Serville, 1831; the Praxibulini are confined to east- ern Australia and Tasmania (Rehn 1957, Rentz 1996). To date, the genus Prterotiltus contains 12 described species; these are all from the wet forests of West and Central Africa. The genus needs revision; several species are known only from single-sex types or ancient and sometimes poorly preserved material. Fifty years ago, Descamps (1972) included Pterotiltus in his list of African genera urgently requiring revision. Pterotiltus is rich in species but poor in museum specimens. Due to its relatively inaccessible habitat, the genus suffers from severe undercollection; it is considerably difficult to penetrate and collect specimens in the forest block, and the on- going reduction of tropical forest decreases the habitat of Pterotiltus, which has strict ecological requirements. Incomplete identifications of species have complicated the work of ecologists who have a ten- dency to either eliminate certain results and risk being incomplete or to include them and risk being inexact (Descamps 1972). Hollis (1975), reviewing the subfamily Oxyinae, redefined the genus and wrote that it seems to represent the culmination of adaptation to forest life in the Oxyinae. All its species are flightless, inhabitants of the herb layer of forests, and the ovipositor valves have become smooth and rodlike, obviously adapted to specialized oviposition sites (Hollis 1975). Rowell (2005) subsequently described epiphyl- lic oviposition on the foodplant of Pterotiltus hollisi. Pterotiltus species occur from Ghana in West Africa through Cameroon and the Congo Basin into Central Uganda. The genus lives in the equatorial forest belt and covers an altitudinal range from sea level to more than 2000 m. Here, we describe specimens recently collected in the lowland and highland forests of Cam- eroon. We describe four new species and the previously unknown female of P. minimus Ramme, 1929. We further erect a new genus, Parapterotiltus, to accommodate minimus and two of our new spe- cies. The present authors are preparing a review of the genus Ptero- tiltus, and the current paper is a preliminary to this. Materials and methods Field sites.—Collections were made by Oumarou Ngoute C. in six localities in highland and lowland rainforests in Cameroon. Two of these are located in the lowland forests of the southern Cameroon: Campo (in Campo National Park) in Ocean division and Ngoyla (in Nki National Park) in High-Nyong division (Fig. 1). Copyright Charly Oumarou Ngoute & C.H.F. Rowell. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) 128 Of the other sites, three (Banganté in Nde division, Mount Bam- boutos in Bamboutos division, and Bamenda in Mezam division) are located in the highland rainforests of Western Cameroon. Makénéné is in the Mbam and Inoubou division in the foothill of these mountains (Fig. 1). Morphological study.—Specimens were studied using a Wild stereo microscope. Drawings were made using the same microscope with a drawing tube and refined in Photoshop (Adobe Corp.). Phallic prep- arations were dissected from pinned specimens relaxed in water, then macerated in 8% KOH, cleaned manually, neutralized in 5% acetic acid, and stained in a weakly acidic solution of acid fuchsin. The ter- minology used for genitalia description is that of Hollis (1971, 1975). Measurements.—Measurements were made under the microscope using a graticule eyepiece and a digital stage (Mitutoyo) reading to 0.01 mm. The following measurements were made: P, length of the pronotum in the midline; L, overall length from the tip of the fastigium to the most posterior part of the abdomen, measured in lateral view; Ant, length of antenna from the scape to the tip of an- tenna; IOS, inter-ocular space measured dorsally; E-E, total ocular width measured dorsally; F maximum length of the hind femur; FD, depth of hind femur measured as the distance between the two lines drawn parallel to the long axis of the femur and touching the dorsal and ventral extremities of the femur; Tal, Ta2, Ta3, lengths of each of the three hind tarsal segments; Sp Ext, number of external spines on the hind tibia; Sp Int, number of internal spines on the hind tibia (Rowell 2005, Oumarou-Ngoute and Kekeunou 2017). Measurements and ratios are given as the range and average of in- dividual measurements and are also presented after normalization [obtained by dividing the measured value by P (the length of the pronotum)]. The normalized values allow comparison of species of different sizes. The foot formula shows the relative lengths of the three tarsal segments within the total length of the foot and again allows for comparison between different species (Rowell 2005). The foot formula values and the ratios describing sexual dimor- phism (i.e., male/female) were calculated from the average values of the sample and not from individual measurements. Number (N) of specimens examined/measured.—Pterotiltus ngoylaensis sp. nov., 13 males 2 females; Pterotiltus campoensis sp. nov., 5 males 2 females; Parapterotiltus minimus (Ramme, 1929), 2 males 2 females; Parapterotiltus minimoides sp. nov., 11 males 6 females; Parapterotiltus bamboutos sp. nov., 3 males 5 females. Abbreviation of depositories. — Mf£N Museum ftir Naturkunde, Berlin. Results Taxonomy Order ORTHOPTERA Olivier, 1789 Superfamily ACRIDOIDEA MacLeay, 1821 Family ACRIDIDAE MacLeay, 1821 Subfamily OXYINAE Brunner von Wattenwyl, 1893 Genus Pterotiltus Karsch, 1893 =Pygostolus Karsch, 1891 (syn. Karsch, 1893) Type species. —Pterotiltus impennis (Karsh, 1891). C. OUMAROU NGOUTE AND C.H.FE ROWELL Subsequent literature. —Ramme 1929: 313. Johnston 1956: 258. Jago 1964: 190-204. Dirsh 1965: 579. Johnston 1968: 173. Dirsh 1970: 119. Hollis 1975: 226. Rowell 2005: 33-43. Revised description of the genus.—Body of small to medium size. Integument rugose and pitted on head, thoracic and proximal ab- dominal tergites, but otherwise smooth and shiny. Head. Antennae filiform, longer than head and pronotum to- gether, especially in males. Fastigium of vertex roughly triangular or pentagonal in dorsal view, short, wider than long, sloping for- wards, slightly concave, sometimes with a weak medial carinula; the apex runs smoothly into the frontal ridge. Frons oblique and sometimes slightly incurved in male, in female almost vertical or only slightly oblique; frontal ridge clearly defined in its upper half, with shallow medial sulcus, but weak or obliterated in lower half. Lateral facial carinae complete. Eyes small or large, almost round or oval, strongly convex; interocular space in males narrower, in females equal to or slightly wider than antennal scape. Thorax. Dorsum of pronotum cylindrical, crossed by three deep transverse sulci, medial carina absent or very weak, lateral carinae absent; mesosternal interspace as long as wide or longer than wide. Metasternal interspace open or nearly closed. Pros- ternal tubercle simple, conical, vertical, with acute or slightly rounded apex. Tegmina and hind wings absent or extremely re- duced, usually represented only by small ridges of integument on mesothorax and metathorax. In some species, minute tegmina are discernable, not extending beyond the hind margin of the meso- thoracic segment. Hind femur slender (4-5 times as long as wide), its tip exceeding the end of the abdomen. Lower lobe of hind knee spined, as is typical for the subfamily. Hind tibia moderately ex- panded distally, with 9 internal and 7 external spines. External api- cal spine present. Internal tibial spurs much larger than external spurs. Arolium large. Second tarsal segment very short (ca. 15% of foot), third segment long (50-60%). Abdomen. Tympanum oval or circular. Last abdominal tergite of male partially divided with a minute furcula, the paired projections in some species fusing to form a single medial process or a short medial tubercle. Male supra-anal plate widely triangular, very short. Female supra-anal plate somewhat longer and more lingulate than in male. Male and female cerci laterally compressed, triangular in lateral view, wide at base, narrowing to an acute or spine-like apex, straight or slightly incurved in dorsal view. Male subgenital plate very short, rounded. Ventral surface of female subgenital plate flat or smoothly rounded, its posterior margin straight or concave, never with posteriorly directed processes (vide Parapterotiltus gen. nov.). Dorsal valves of ovipositor laterally compressed, with round- ed apices; ventral valves smooth, slender, straight, rod-like, some- times dorsally flattened or slightly grooved. Egg guide prolonged horizontally rearwards between the ventral valves, up to half the length of the latter, straight or weakly curved, sharply pointed. Spermathecal vestibule large, sac-like; spermatheca with small api- cal diverticulum and a large curved subapical diverticulum. Epiphallus with divided bridge, ancorae absent or extremely reduced, and one or two pairs of lophi; in most species, the outer lophi are long, slender, blade-like, and curved forwards. Ectophal- lic membrane well developed, including a ventrolateral sclerite en- circling the ventral half of the ectophallus. Cingular valves fused together with the arch sclerite and elaborated into a valvular plate (Hollis 1971) covering the aedeagus dorsally and often laterally. In many species, this plate is foliose, being extensively ornamented with cuticular folds and lamellae, often of species-specific form. Endophallus with a slender flexure leading to short processes that JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) C. OUMAROU NGOUTE AND C.H.E ROWELL do not themselves form aedeagal valves. Aedeagus composed of the valvular plate and of ventral aedeagal sclerites on which the endophallic processes terminate. The known species of Pterotiltus are morphologically extreme- ly homogenous, outwardly differing principally in size and col- oration. Their phallic structures are more diverse, especially the epiphallus and the valvular plate. Where both sets of data are available, phallic diversity confirms the species boundaries origi- nally drawn on the basis of coloration. Pterotiltus ngoylaensis sp. nov. https://zoobank.org/6A8C75F6-8559-4998-AC05-F630140FCE25 Figs 1, 2A, B, 3-5, Table 1 Material examined.—Holotype: 3; CAMEROON e High-Nyong Di- vision, Ngoyla, Nki National Park; altitude 510 m; 10 May 2021; C. Oumarou Ngoute leg.; specimen no. 2021007; MfN. Paratype: CAMEROON ° 19; same data as holotype; specimen no. 2021006; MfN. Other material examined: CAMEROON e 12 4, 1 9; same data as holotype; personal collection of C. Oumarou Ngoute. Type locality. —Cameroon, High-Nyong Division, Ngoyla, Nki Na- tional Park. Etymology.—Toponymic, referring to the locality of Ngoyla, Cam- eroon. Diagnosis. —Color of body as in Fig. 2A, B: multi-colored, predomi- nantly green with black and yellow markings, eyes noticeably violet ( oO wd 0 i; Bamenda Mbengwi ome wy $428 Mamte wi 129 in life. Average size of body: 20.3 mm (males) and 24.4 mm (fe- males) (Table 1). Metazona of pronotum short, about one-fourth the length of prozona (Fig. 3C, D). Tegmina extremely reduced, narrow, and straight, not reaching the posterior margin of meso- thoracic segment (Fig. 5). Furcula of the last abdominal tergite in male with acute corners either side of the midline (Fig. 3A), separa- tion of tips 0.53 mm. Epiphallus as in Fig. 4G, H: large forwardly curved blade-like outer; inner lophi absent; anterior processes of epiphallus well developed, forward pointing; ancorae absent; oval sclerites present, with two small tubercles on their ventral surfaces. Description.—(Figs 2-5, Table 1). Male. Body of medium size (average L = 20.3 mm). Fastigium of vertex pentagonal, about three times wider than long; frons slightly oblique; eyes large, oval, protuberant; interocular space equal to or slightly wider than antennal scape. Pronotum: me- dial carinae absent (Fig. 3C, D); the space between sulci 2 and 3 almost equal to the length of metazona; metazona short, about one-fourth of the length of prozona, and only 19% of the total pronotal length; posterior margin of metazona slightly notched in the midline, otherwise straight; anterior margin of pronotum slightly convex, minutely notched in midline; prosternal process conical, pointed; mesosternal space as wide as long; metasternal space open, about half the length of mesosternal space (Fig. 3B). Tegmina extremely reduced: in male, minute, narrow and straight, not reaching the posterior margin of mesothoracic segment, in female even shorter, rounded (Fig. 5). Posterior ventral angle of pronotum rounded, only slightly produced rearwards. Hind tibia slightly expanded apically. Tympanum oval; last abdominal tergite Tignére> “cSambolabbo ) ja ere’. < i Ber AP XH A OUR O > J | Collecting sites of studied species 150km Jr: B® Pterotiltus ngoylaensis sp. nov. i Pterotiltus campoensis sp. nov. Parapterotiltus minimoides sp. nov. i Parapterotiltus bamboutos sp. nov. §) Parapterotiltus minimus (Ramme, 1929). Fig. 1. Collecting sites of the studied species. (Source : UN Cartographic Section https://www.nationsonline.org/oneworld/map/cam- eroon-administrative-map.htm) JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) 130 C. OUMAROU NGOUTE AND C.H.F ROWELL B * ™. a 7 : ' 44 ‘a - m\ Ai iC af db ae e ¥ + Pe SES Fig. 2. Pterotiltus ngoylaensis sp. nov. and Pterotiltus campoensis sp. nov. habitus. A. P. ngoylaensis male; B. P. ngoylaensis female; C. P. cam- poensis male; D. P. campoensis female. divided with acute corners either side of the midline (Fig. 3A), the hind margin with very small lobiform processes forming a minute furcula; cercus straight, strongly compressed laterally, with acute apex, shorter than the subgenital plate but exceeding the supra- anal plate. Supra-anal plate (Fig. 3A) triangular in dorsal view, somewhat elongated distally, with a rounded apex; proximally, there is a short medial longitudinal groove bounded posteriorly by a curved transverse ridge that extends across the width of the plate. Phallic complex (Fig. 4) of large size for the genus, epiphal- lus with large forwardly curved blade-like outer lophi (Fig. 4G, H); inner lophi absent; anterior processes of epiphallus well de- veloped, ancorae absent (Fig. 4H). The oval sclerites have two small tubercles on their ventral surfaces. Lobular plate apparently formed from the midline fusion of a pair of bilaterally symmetri- cal structures, in dorsal view showing three main lobes, each of which has a concave inner surface (Fig. 4B-E). Female. In general, similar to the male. Female particularities: medium size (average L = 24.4 mm), fastigium of vertex pentago- nal and concave in dorsal view; last abdominal tergite divided but furcula absent; cercus straight in dorsal view, equal in length to the supra-anal plate or slightly shorter. Egg guide rodlike, pointed, produced rearwards, about 1/3 length of ventral valve. Sexual dimorphism.—The average values of P for males are 87% of those for females. After normalization, the ratios of the vari- ous body dimensions are closely similar in both sexes; in the male, the interocular space (IOS) is relatively smaller than in the female, and the antennae are somewhat longer (Table 1). In this species, the average length of the hind foot is about equal in both sexes, whereas in some other species, the foot of the male is slightly longer. C POST D Fig. 3. Pterotiltus ngoylaensis sp. nov. A. Dorsal view of male termi- nalia; B. Thoracic sterna; C. Pronotal disc showing sulci; D. Prono- tal disc showing coloration and patterning in life. Scale bar: 1 mm. JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) C. OUMAROU NGOUTE AND C.H.E ROWELL 131 Table 1. Measurements of Pterotiltus ngoylaensis sp. nov. P: length of the pronotum in the midline; L: overall length from the tip of the fastigium to the most posterior part of the abdomen measured in lateral view; Ant: length of antenna from the scape to the tip of antenna; IOS: inter-ocular space measured dorsally; E-E: total ocular width measured dorsally; F: maximum length of the hind femur; FD: depth of hind femur measured as the distance between the two parallel lines drawn parallel to the long axis of the femur, running through the dorsal and ventral extremities of the femur; Tal, Ta2, Ta3: lengths of each of the three hind tarsal segments; Sp Ext: number of external spines on the hind tibia; Sp Int: number of internal spines on the hind tibia. Measurements and ratios are given as the range and average of individual measurements and are also presented after normalization [obtained by dividing the measured value by P (the length of the pronotum)]|. The foot formula shows relative lengths of the three tarsal segments within the total length of the foot and again allows for comparison between different species. N is the number of specimens analyzed/measured. Male Dimensions in mm 1? fh Ant IOS E-E F FD Tal Ta2 Ta3 Tal+2+3 External spines External spines Minimum 3:96. «18.30° 9.45, O47 4.53 12342 249» 0,95. 059° 242 3.87 7 9 Maximum 446 22.14 13.94 0.76 5.54 1349 3.10 1.29 0.85 2.87 4.78 7 9 Average 4.14 20.30 11.65 064 453 13.01 2.85 1.17 0.71 2.49 4.37 7 9 N 13 13 13 13 13 13 13 10 10 10 10 10 10 Foot formula 0.27 (0.16> 0:57 Ratios L/P Ant/P IOS/P E-E/P F/P FD/P Tal/P Ta2/P Ta3/P Tal+2+3/P F/FD Minimum W25 =2s32 BON «O94? P2189.9 059° 023° OMS. 0.50 0.92 4.32 Maximum oe" BS), Och 1.3820 “S32 Pe0s25- 70327 (02k 80272 ile | 5.16 Average 4.92 2.74 0.15 Li = 39S 0.69" “028° O.17 > 0.60 1.05 4.60 N 13 13 13 13 13 13 10 10 10 10 10 Female Dimensions in mm P 1B; Ant IOS E-E F FD Tal Ta2 Ta3 Tal+2+3 External spines External spines Minimum 4.61 2436 11.26 084 4.93 14.16 3.14 144 045 2.62 4.51 7 9 Maximum 4.86 2444 11.32 090 4.96 1453 3.43 158 0.76 2.86 5.20 7 9 Average 4.74 2440 11.29 0.87 4.95 14.35 3.29 1.51 O60 2.74 4.85 7 9 N 2 2. 2 2 2 2 2 2 2 2 2: 2 2 Foot formula 0:31 O13 0.56 Ratios L/P Ant/P IOS/P E-E/P F/P FD/P Tal/P Ta2/P Ta3/P Tal+2+3/P F/FD Minimum DOS: 233 Ose Ale -299° 0.68" .0:30" 0.09. 054 0.93 4.24 Maximum 5.28 244 0.20 1.08 3.07 O.71 O34 0.16 0.62 1a13 4.51 Average bilo. 2395 ls 11:05", =3,037 =0699, O32. 0018.5 MO,58 1503 4.37 N 2 2 2 2 2 2 2 2 2 2 2 Sexual dimorphism Average male/average 0.87 0.83 103s 6 Ou73 41-0792), 0 0291 W878 w OL7 7 Tals 0.91 0.90 1.05 female Normalized on 87% 1.00 0.96 1.19 0.84 1.05 1.04 100 089 1.35 1.04 1.03 Male/female ratio O25 AIS! ! 20:1 106 1.004 0.99 0.88 1.33 1.03 1.02 1.05 Coloration.—The description given here refers to the living ani- mal (Fig. 2A, B). When dried, specimens of P. ngoylaensis lose their natural colors and become almost uniformly dark olive brown. Body multicolored in both sexes, predominantly green. Scape and pedicel of antenna green-brown, flagellum green- brown, sometimes lighter brown apically; fastigium green; ver- tex blue-black with black longitudinal bands either side of the midline, which extend dorsally onto the pronotum; eyes con- spicuously violet in life, fading to brown when dried; upper half of frons blue-black, lower half whitish-yellow, extending poste- riorly across the genae as a white-yellow subocular stripe and forming two pale patches on the ventral margin of the pronotal lobes, extending rearwards to the mesothoracic and metathorac- ic pleura as well. Distal margin of clypeus black; distal region of labrum, palps, and lateral margins of mandibles green. Pronotal disc multicolored, predominantly black, with two longitudinal pale bands; these bands are prolonged dorsally onto the mes- othoracic and the metathoracic tergites; thoracic sterna green; hind femur green with the upper genicular lobe red; tibia and tarsi green. The first abdominal segment black, with two white- yellow longitudinal bands; the 2‘ and 3“ segment black, some- times slightly white dorsally; the 4° segment green, sometimes slightly white in dorsal view; all the remaining abdominal seg- ments green. Distribution.—Cameroon: _High-Nyong Division, _Ngoyla, Nki National Park: 2°27'26'N, 14°34'0O1"E, altitude 505 m; 2°35'37"N, 14°25'58'E, altitude 498 m; 2°29'58"N, 14°23'36"E, altitude 510 m. Remarks.—The eyes of Pterotiltus ngoylaensis are conspicuously vio- let in life, which is, to date, a unique character in the genus. The large forwardly curved blade-like outer lophi are typical of the ge- nus and are particularly close to the structure observed in Pterotil- tus coeruleocephalus Bolivar, 1905. However, P. ngoylaensis is unique within its genus in having oval sclerites with two small tubercles on their ventral surfaces. The type locality in the Nki national park (Fig. 1) bodes well for conserving its natural habitat against de- forestation. However, the species may well occur more widely in southern Cameroon. Its habitat is typical of the genus: rainforest, 400 to 600 m above sea level, with abundant Marantaceae and Commelinaceae in the understory. These conditions are found in many parts of the south Cameroon plateau and neighboring countries (Congo, Gabon, and Central African Republic). JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) 132 C. OUMAROU NGOUTE AND C.H.E. ROWELL lobes of valvular plate Ventrolateral sclerite valvular plate AJ lobes of valvular plate arch ; Fee arch lobes of pas valvular plate cingular apodeme Cc H Fig. 4. Diagrams of phallus of Pterotiltus ngoylaensis sp. nov. A. Lateral view of entire phallic complex, shown somewhat extended relative to the resting state. Ectophallic membrane is coarsely stippled and obscures most of the sclerites. The ventrolateral sclerite (shaded) lies in this membrane and is clearly visible in the extended preparation. The cingular apodeme and the cingular ramus are indistinctly vis- ible through the membrane (partially dashed line). Parts of the epiphallus and of the ventral aedeagal valve (shaded) are seen protrud- ing through the membrane. The cingular valves are replaced by a valvular plate (sensu Hollis, 1971), which is foliose in this genus, as shown in B. and E. below; B, C. Endo- and ectophallic structures dissected free of epiphallus and ectophallic membrane (scale bar does not apply) in (B) dorsal and (C) lateral views. In C, the approximate course of the endophallic flexure and aedeagal valve are shown with dashed lines; D. Lateral view of the arch sclerite and its associated valvular plate dissected free from the cingulum. The whole struc- ture is saddle-shaped, being concave ventrally and fitting over and around the ventral aedeagal valve (C); E. Dorsal view of the arch and its valvular plate showing its multiple lobes. The dashed vertical lines indicate denser areas seen in transmitted light, which suggest the presence of paired cingular valves but is illusory, as no such valves are present; F. Lateral view of endophallus dissected free of the arch and the cingulum. Heavily shaded areas are sclerotised. The lightly shaded area is the ventral aedeagal sclerite on which the endophallic process terminates and which forms the visible part of the ventral aedeagal valve; G, H. Epiphallus in (G) axial and (H) dorsal view. This species has only a single pair of external lophi, which are long, bladelike, and curved forward over the bridge. The “oval” sclerites are shaded in G and have two small tubercles on their ventral surface, indicated in H, a peculiarity of this species. Scale bar: 1 mm. JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) C. OUMAROU NGOUTE AND C.H.E ROWELL Pterotiltus campoensis sp. nov. https://zoobank.org/913ECCD9-0C05-424D-BOBO0-068E1C3CDC72 Figsall; 2€3 D607, Tabled Material examined.—Holotype: CAMEROON e 3; Ocean Division, Campo, Campo National Park; altitude 5 m; 13 Nov. 2017; C. Ou- marou Ngoute leg.; specimen no. 2017242; MfN. Paratype: CAM- EROON e 19; same data as holotype; specimen no. 2017243; MfN. Other material examined: CAMEROON e 4 4, 1 9; same data as holotype; personal collection of C. Oumarou Ngoute. Type locality.—Cameroon, Ocean Division, Campo, Campo Na- tional Park. Etymology.—Toponymic, referring to the locality of Campo, Cam- eroon. Diagnosis.—Color of body as in Fig. 2C, D: multicolored, pre- dominantly shades of red and blue-black. Average size of body: 17.92 mm (male) and 20.07 mm (female) (Table 2). Metazona of pronotum very short, about one-fifth of the length of prozona (Fig. GE, F). Metasternal space very small, nearly closed (Fig. 6B). Apterous. Furcula of last abdominal tergite in male very small, lobiform, almost vestigial, as in Fig. 6A. Epiphallus as in Fig. 6C, D: a narrow bridge bearing a pair of large rectangular outer lophi and one or two pairs of smaller inner lophi; anterior processes of epiphallus weakly developed; small forward-pointing projections on anterior edge of bridge, which may represent the rudiments of ancorae; oval sclerites elongate and irregular in shape. Description.—(Figs 2, 6, 7, Table 2) Male. Body of small-medium size (L = 17.92 mm), somewhat more slender and relatively longer than the other species (L/P = 5.17, in most other species <5.0). Fastigium of vertex short, pen- tagonal in dorsal view, somewhat concave; frons slightly oblique; eyes large, oval, convex; inter-ocular space equal to or slightly wider than antennal scape. Pronotum (Fig. 6E, F) medial carinae absent, the space between sulci 2 and 3 longer and more convex laterally than elsewhere; metazona very short, less than one-fifth of the length of prozona (15.6% of pronotum); posterior mar- gin of metazona slightly embayed and notched in the midline, otherwise straight; anterior margin of pronotum slightly convex; prosternal process conical, sharply pointed; mesosternal space (Fig. 6B) as wide as long; metasternal space very small, nearly closed. Wings and tegmina absent. Hind tibia not significantly expanded apically, hind femur slender (F/FD ca. 4.5) with weakly impressed chevron markings. Tympanum oval or circular; last ab- dominal tergite incompletely divided, the hind margin with very small thickenings suggesting a minute furcula, almost indiscern- ible, separation of midpoints 0.39 mm; cercus straight, extending beyond the tip of the subgenital plate (Fig. 6A). Supra-anal plate very short, triangular in dorsal view, tip rounded; proximally, there is a short medial longitudinal groove, bounded posteriorly by a transverse ridge that extends across the width of the plate (Fig. 6A). Epiphallus with a rather narrow bridge bearing a pair of large rectangular outer lophi and one or two pairs of smaller inner lophi (Fig. 6C, D). Anterior processes of epiphallus weakly developed; what may be small rudiments of forward pointing an- corae present; oval sclerites elongate and irregular in shape. Ecto- and endophalli typical of genus. The valvular plate consists of a small saddle-shaped arch with two vertical lamellae, one either side of the ventral valve (Fig. 7). Fig. 5. Pterotiltus ngoylaensis sp. nov., elytral rudiments. A. Female; B. Male, lateral view of elytra (shaded) at posterior edge of prono- tum. PRNMZ, metazona of pronotum; T2 TG, mesothoracic ter- gite; T2 EPST, mesothoracic episternum, T2 EPMR, mesothoracic epimeron; T3 TG, metathoracic tergite; CXM, coxal membrane. Scale bar: 1 mm. POST E Fig. 6. Pterotiltus campoensis sp. nov. A. Dorsal view of male termi- nalia; B. Thoracic sterna; C, D. Epiphallus; C. Axial view; D. Dor- sal view; E. Pronotal disc showing sulci; FE. Pronotal disc showing coloration and patterning in life. Scale bar: 1 mm. Female. In general, similar to the male. Female particularities: small to medium size (L= 20.07 mm), last abdominal tergite in- completely divided but no furcula; cercus long and straight, slightly compressed laterally, tapering to an acute apex, slightly shorter than JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) 134 RHS cingular apodeme flexure C broken LHS apodeme C. OUMAROU NGOUTE AND C.H.F ROWELL valvular plate ventral aedeagal valve d / x” “ss RHS ramus endophallic process ventral aedeagal sclerite Fig. 7. Pterotiltus campoensis sp. nov. phallic structure. A. Endophallus and part of ectophallus after removal of the outer ectophal- lic sheath. The LHS cingular ramus and most of the LHS cingular apodeme has been removed to expose the endophallus. The basal endophallic valves give rise to paired narrow flexures that lead to the ventral aedeagal valves; B. Endophallus after removal of the ectophallic valvular plate and cingulum. The endophallic processes are short and terminate in spatulate endings that are appressed to ventral aedeagal sclerites, which form the greater part of the ventral aedeagal valve; C. More magnified view (scale does not apply) of the junction between the RHS endophallic process and the ventral aedeagal sclerites, viewed from the midline. The precise relations of the terminal sclerites and the membrane of the spermatophore sac are presently unclear. Scale bar: 1 mm. length of the supra-anal plate. Egg guide rod-like, pointed, produced rearwards for slightly more than half the length of the ventral valves. Sexual dimorphism.—The average values of P for the male are 85% of those of the female. After normalization, the ratios of the various body dimensions are similar in both sexes; only the in- terocular space (IOS) is relatively smaller in the male than in the female. Data on relative antennal length (which are significantly longer in the male in other species) are not available (Table 2). There is no difference between the sexes in the relative lengths of the hind foot. Coloration.—(Fig. 2C, D) Body multicolored in both sexes, pre- dominantly red in dorsal view; scape and pedicel of antenna green, flagellum entirely red, sometimes light brown apically; fastigium blue; vertex blue with two indistinct yellow or red lateral longitu- dinal bands diverging rearwards; eyes red in life, fading to yellow- ish-brown when dried; frons blue-black with lighter mottling on clypeus and labrum. Palps green. Genae blue-black, with a narrow yellow subocular stripe; post ocular stripe black, extending onto the pronotum. Pronotal disc multi-colored, predominantly black with two yellowish patches at the anterior margin of prozona and two more at the posterior margin of metazona, and red pigment in the midline between the yellow patches; mesothoracic tergite black with small red areas along its posterior margin either side of midline; metathoracic tergite mainly red; abdominal tergite 1 red medially with yellow pigment laterally, tergites 2 and 3 black, tergite 4 yellowish, remainder of abdomen in general blue. Tho- racic and abdominal sterna blue-black; prosternal spine brownish, probably green in life. Hind femur yellowish proximally and green JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) C. OUMAROU NGOUTE AND C.H.E ROWELL 135 Table 2. Measurements of Pterotiltus campoensis sp. nov. P: length of the pronotum in the midline; L: overall length from the tip of the fastigium to the most posterior part of the abdomen measured in lateral view; Ant: length of antenna from the scape to the tip of antenna; IOS: inter-ocular space measured dorsally; E-E: total ocular width measured dorsally; F: maximum length of the hind femur; FD: depth of hind femur measured as the distance between the two parallel lines drawn parallel to the long axis of the femur, running through the dorsal and ventral extremities of the femur; Tal, Ta2, Ta3: lengths of each of the three hind tarsal segments; Sp Ext: number of external spines on the hind tibia; Sp Int: number of internal spines on the hind tibia. Measurements and ratios are given as the range and average of individual measurements and are also presented after normalization [obtained by dividing the measured value by P (the length of the pronotum)]|. The foot formula shows relative lengths of the three tarsal segments within the total length of the foot and again allows for comparison between different species. N is the number of specimens analyzed/measured. Male ee - Dimensions in mm iE L Ant IOS E-E F FD Tal Ta2 Ta3 Tal+2+3 External spines External spines Minimum 3.31 1662 940 030 4.07 10.31 2.40 0.97 0.28 2.06 3.78 7 9 Maximum 3.86 18.91 1242 066 456 11.59 2.66 166 0.63 2.40 4.32 7 9 Average 3151 e792. IVA, 048, 423. J22) 2570 129. 20.49% 22:23 4.02 7 3) N 5 5 3 5 5 4 4 4 4 4 4 4 4 Foot formula 032 (0135 0:55 Ratios L/P Ant/P IOS/P E-E/P F/P FD/P Tal/P Ta2/P Ta3/P Tal+2+3/P F/FD Tal+2+3/F Minimum 5.02. 284° \5009 (1,22. 3.1 “0.735 -0:37 “006 0,66 0.76 4.30 0.33 Maximum 5.40 3.75 0.14 1.30 3.48 0.79 O53 #O.19 0.72 1Wg2. 4.43 0.39 Average SslZ. B22. 2Ocld 1252) BBS O77 0.46" “OZ 0.69 1.46 4.37 0.36 N 5 3 5 5 4 4 4 4 4 4 4 4 Female Dimensions in mm R 1B; Ant IOS E-E F FD Tal Ta2 Ta3 Tal+2+3 External spines External spines Minimum 4.07 19.60 0:65 4:86 133999 3:02 1.20 0.58 3.00 4.91 7 9 Maximum 4.19 20.54 0.69 4.90 14.03 3.16 146 0.71 3.16 5.20 7 9 Average 4.13 20.07 0.67 4.88 14.01 3.09 1.33 0.64 3.08 5.05 7 9 N 2 2. 2 2 2 2 2 2: 2 2: 2 2 Foot formula 0.26 0.13 0.61 Ratios L/P Ant/P IOS/P E-E/P F/P FD/P Tal/P Ta2/P Ta3/P Tal+2+3/P F/FD Tal+2+3/F Minimum 4.82 0.16 1.16 3.35 0.72 O35 O14 0.74 1.24 4.65 0.35 Maximum 4.90 0.16 1.16 344 0.72 O35 O14 0.75 1.24 4.65 0.37 Average 4.86 O06" 1,16 23:39 “O72 “O35 -Ojl4= “0.75 1.24 4.65 0.36 N 2 2 2 2 2 2. 2 2 2. 2 2 Sexual dimorphism Average male/average 0.85 0.89 O72 AQ 7S * X80" 90:83" O97 ~OAT* » 0372 0.80 female Normalized on 85% 1.00 1.05 0.85 1.002 094 0.98 1.15 0.91 0.85 0.94 Male/female ratio 1.06 O82 “108 <0:99) “L06" 1:33" 124 0:92 1.18 0.94 1.01 distally; knee blue-black with upper lobe tinged brownish, lower lobe blue; tibia and tarsi blue-black. Distribution. —CAMEROON, Ocean Division, Campo: 2°20'43"N, 9°59'28"E, altitude 5 m. Remarks.—Pterotiltus campoensis is most noticeable for its predomi- nantly red and blue coloration, a unique character in the genus. The epiphallus with a pair of large rectangular outer lophi is close in this genus to Pterotiltus apicalis Bolivar, 1905 (See fig. 63 in Hollis 1975). However, P. campoensis is distinguishable by for- ward-pointing projections on anterior edge of bridge, which may represent the rudiments of ancorae. The new taxon is found in hu- mid coastal forests at very low altitude (less than 15 m above the sea level). It was found both in wet areas and in the drier part of the forest, where it could be caught on young tree branches and on dead branches. We recorded the species in the Campo Ma’an Na- tional Park (Fig. 1); its natural habitat is therefore relatively well protected, despite the accelerated rate of deforestation in southern Cameroon. As with P. ngoylaensis, this species might well be found in other similar localities in south Cameroon and possibly in the neighbouring countries of Equatorial Guinea and Gabon. Genus Parapterotiltus gen. nov. https://zoobank.org/F5809A37-C3F8-43B3-9C9A-EIB5COF5C64E Table 3 Type species. —Pterotiltus minimus Ramme, 1929. Etymology.—Greek Para, near, close to; Pterotiltus, a genus of oxyine grasshopper, erected by Karsch, 1893. Diagnosis.—Differs from Pterotiltus Karsch, 1893 only in the fol- lowing particulars (Table 3): Size generally smaller; males aver- age 12-14 mm in length, females 16-22 mm. Cuticle of fasti- gium and vertex in head and of thoracic and proximal abdominal tergites, coarsely punctate. Tegmina present but micropterous, barely exceeding the mesothoracic segment, rounded and some- times brightly colored (Fig. 8). Female subgenital plate with two robust posteriorly directed projections, arising slightly anterior to the posterior edge, that project horizontally below the ventral ovipositor valves. Valvular plate of phallus simple, not foliose. Epiphallus with the bridge reduced, outer lophi long, upwardly and inwardly curved, either pointed and “horn-like” or bifid at their tips. Inner lophi absent or very small. Ancorae absent. The JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) 136 anterior processes of the epiphallic bridge, if present at all, tend to be elongated and inwardly pointed. Distribution.—The 3 known species are all from the Cameroonian highlands. Parapterotiltus minimus (Ramme, 1929) Figs 1, 8-10, Tables 4, 7 Material examined.—Holotype: CAMEROON e ©; West-Central Tropical Africa, Bamenda; 5°95'12"N, 10°16'69"E; W. Ramme leg.; DORSA no. 000031801; MfN. Topotype: CAMEROON « 10; Mezam division, Bamenda; 5°57'35"N, 10°08'50"E; altitude 1598 m; C. Oumarou Ngoute leg.; specimen no. 2017248; MfN. e 19; Mezam division, Bamenda; 5°57'35"N, 10°08'50"E; alti- tude 1598 m; C. Oumarou Ngoute leg.; specimen no. 2017247; MfN. Other material examined: CAMEROON e 1, 19; same data as topotype; personal collection of C. Oumarou Ngoute. Type locality.—Cameroon, Mezam division, Bamenda. Diagnosis.—Color of body as in Fig. 8A-D: multicolored, predom- inantly brown. Thoracic and abdominal terga brown, each seg- ment usually with a single, midline yellow spot in male (Fig. 8A). Average size of body, 13.23 mm (male) and 21.59 mm (female) (Table 4). Fastigium of vertex slightly concave, sloping down- wards. Pronotal medial carinae absent; metazona much shorter than prozona, making up only 17% of pronotal length; metaster- nal interspace very small, narrowly open; tegmina very small with rounded, almost spherical tip. Cerci compressed laterally, with acute apex not exceeding the tip of subgenital plate. Outer lophi of epiphallus long, pointed, horn-like; inner lophi absent (Fig. 9A, B); absence of anterior processes (Fig. 9A, B). Redescription.—(Figs 8-10, Tables 4, 7). Male. Size of body, 13-14 mm (male) and 21-22 mm (fe- male) (Table 4). Fastigium broadly triangular, sloping down- wards, its surface slightly concave with no medial carina, its tip merging smoothly with the frontal ridge. Pronotum devoid of me- dial carinae. Anterior margin slightly excurved, posterior margin transverse, straight, minutely notched in midline. Metazona much shorter than prozona, making up only 17% of pronotal length. Ventral margin of lateral lobes strongly sinuate, posterior angle rounded but produced slightly rearwards. Prosternal process verti- cal, acutely pointed. Mesosternal interspace about as wide as long, Table 3. Tabular key to genera. C. OUMAROU NGOUTE AND C.H.F ROWELL lateral lobes rounded. Metasternal interspace very small, narrowly open. Hind femur slender, outer face with weak chevron-shaped impressions. Hind knee smooth dorsally, with no medial terminal point. Upper genicular lobes rounded, hind tibia slender. Inter- nal hind tibial spurs somewhat larger than external spurs. Ventral face of hind tibia rather densely hirsute, dorsal face only sparsely. Arolium large. Tegmen very small (total length 0.94 mm), with rounded, almost spherical tip. Wing absent. Tympanum circular. Tenth abdominal tergite divided, furcula minute, points sepa- rated by 0.39 mm. Supra anal plate short, triangular, pointed, wider than long, not completely covering pallium. Cerci laterally flattened, wide at base, tapering to acute melanised points not exceeding subgenital plate. Subgenital plate short, rounded. Phal- lus with a voluminous ectophallic membrane containing a vent- rolateral sclerite. Aedeagus consists only of the ventral valves and the valvular plate. This plate is, however, simple, resembling that of Oxya (Hollis 1971). The endophallic processes do not reach the exterior but end shortly after their flexure on ventral aedeagal scle- rites that constitute the actual tips of the ventral valves. Epiphallus distinctive (Fig. 9), with divided bridge, long pointed “horn-like” lophi, ancorae and anterior processes absent. “Oval” sclerites pre- sent, elongate, roughly triangular. Female. Similar to male, but much larger and more robust (L = 21.6 mm), and some differences in coloration (see below). Egg guide produced into a pointed rod lying between the bases of the ventral ovipositor valves (Fig. 10B, C). Supra-anal plate lingulate, with a rounded tip and a prominent midline depression proximal- ly; cerci straight, slightly shorter than supra-anal plate (Fig. 10A). Sexual dimorphism.—(Table 4) P male/P female is only 0.71, indicat- ing a considerable size difference between the sexes. Once again, the male antenna is proportionately longer and the inter-ocular space smaller than in the female. The eyes are slightly more protuberant (E-E) in the male than the female. The foot formula is practically identical in the two sexes; the second tarsal segment is very short (14% of the foot) as is typical for non-arboreal grasshoppers. The entire hind foot is relatively somewhat longer in the male than the female. Note however that N, the sample size, is low in this species. Coloration.—Male. Vertex, occiput, and posterior half of the genae matte black. Eyes shiny black in life, drying brown. Fas- tigium, frons, and anterior half of genae dull green. Basal seg- ments of antenna and proximal flagellar segments green, distal segments orange with paler tips. Clypeus and labrum proximally green, distally black; the upper outer corners of the labrum bear Characters Pterotiltus Parapterotiltus Size of body Small to medium: males average 17-21 mm in length, Generally smaller: males average 12-13 mm in length, females females 20-25 mm (Tables 1, 2). 16 - 22 mm (Tables 4-6). Tegmina Absent or extremely reduced, usually represented only — Present but micropterous, barely exceeding the mesothoracic by small ridges of integument not extending beyond segment, rounded and sometimes brightly colored (Fig. 8). the hind margin of the mesothoracic segment. Female subgenital plate directed processes. Ventral surface flat or smoothly rounded, its posterior margin straight or concave, never with posteriorly Ventral face with two robust posteriorly directed projections, arising slightly anterior to the posterior edge, that project horizontally below the ventral ovipositor valves (Figs 10, 12). Valvular plate of phallus Foliose, extensively ornamented with cuticular folds Simple, not foliose. and lamellae, often of species-specific form. Epiphallus Outer lophi long, slender, blade-like, and curved forwards; ancorae absent or rudimentary, forward- pointing projections on anterior edge of bridge (Fig. 4). Outer lophi long, upwardly and inwardly curved, either pointed and “horn-like” or bifid at their tips; ancorae absent; anterior processes of the bridge, if present, elongated and inwardly pointed (Fig. 12D). JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) C. OUMAROU NGOUTE AND C.H.E ROWELL enn y 5 bg Sie aes fr cs ‘ . beste. = Fig. 8. Parapterotiltus minimus (Ramme, 1929). A. Typical form; B. Another individual lacking yellow markings; C. Female, typical form; D. Female lacking yellow color on the elytron. two orange or bright yellow patches. Palps and outer surfaces of mandibles green. Pronotum, thorax, and first four abdominal tergites dull brown. Lateral lobes of pronotum with ventral margins irregu- larly marked with pinkish orange. In some specimens, there is a small orange patch in the midline of the anterior pronotal mar- gin. Tegmina bright yellow, contrasting strongly with the dark brown thoracic pleura. Posterior margins of first three abdominal tergites marked with orange or yellow patches in the midline. Ab- dominal segment 5 and all more distal segments light leaf green. Tips of cerci blackish. Fore and middle legs green but with orange femora. Hind femur green, knee black, but lower genicular lobes dark blue. Outer and inner faces of hind femur marked with two dark green longitudinal bands, converging distally. Condyle of hind tibia black, remainder of tibia green, tibial spines and spurs blue black. Female. General impression, dull brown with green legs and tip of abdomen. Differs from male in detailed marking and in the absence of orange femora on fore and middle legs. Posterior half of gena white (tinged with pink), continuing onto ventral margin of the pronotal lobes, thus replacing the orange color of the male in this area. Frons brown, not green. The paired yellow marks on the upper margin of the male labrum are replaced in the female by a yellow central spot on the lower labrum, and the orange medial markings of the pronotum and proximal abdominal tergites are absent. Hind femora green, with darker chevron markings. Inner face of the femur with a single wide dark green longitudinal band. In some specimens, the distal femur and the hind knee are suf- fused with reddish brown instead of the black seen in males. It should be noted that in both the Bamenda and the Ban- ganté populations of P. minimus, there is marked variability in coloration between individuals, especially in the extent of orange markings on the tergites, and in some females the tegmina are not contrastingly colored (Fig. 8). Distribution.—Cameroon: Bamenda (type locality): 5°57'35"N, 10°08'50"E, altitude 1598 m; Banganté: 5°09'30"N, 10°31'58"E, altitude 1325 m. Remarks.—Ramme’s description referred only to the male; the fe- male has been unknown until now. Its divergent characters, espe- cially the female subgenital plate, have prompted a re-examina- tion and, together with the new species described below, have led to our erection of this new genus. The above description differs slightly from that of Ramme (1929). He wrote that the antennae are shorter than described here. His holotype (examined) had broken antennae when exam- ined in 2016. He also described the P. minimus male as having a truncated, squared-off tip to its supra-anal plate and even illus- trated this (his fig. 36) as a specific character. However, the holo- JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) 138 Fig. 9. Parapterotiltus minimus (Ramme, 1929). A. Drawing of epiphallus of holotype from Hollis 1975 (scale bar does not ap- ply); B. Photograph from a recent dissection of a topotypic indi- vidual. Note absence of ancorae and anterior processes and the large divergent “horn-shaped” lophi that are hooked but not bifid at their tips. Scale bar: 1 mm. type presently has a normal triangular supra-anal plate but shows an old fold line. Apparently, Ramme did not realise that the tip of the plate was folded under and probably it did not re-emerge until the specimen was relaxed by Hollis (1975), who dissected out and figured the epiphallus. Parapterotiltus minimoides sp. nov. https://zoobank.org/6E34A7 D8-A20D-4312-BAF5-90B82C01AE09 Figs 1, 11A—D, 12, Tables 5, 7 Material examined.—Holotype: CAMEROON e <; Ndé Division, Banganté; altitude 1335 m; 13 Nov. 2017; C. Oumarou Ngoute leg.; specimen no. 2017241; MfN. Paratype: CAMEROON e 19; same data as holotype; specimen no. 2017240; MfN. Other mate- tial examined: CAMEROON e 104, 59; same data as holotype; personal collection of C. Oumarou Ngoute. Type locality.—Cameroon, Ndé Division, Banganté. C. OUMAROU NGOUTE AND C.H.FE ROWELL Etymology.—From Latin, minimoides, “similar to minimus”. Refers to the external habitus of the species, almost indistinguishable from P. minimus. Diagnosis.—Color of body as in Fig. 11E-H: multicolored, pre- dominantly brown. Thoracic and abdominal terga brown: each segment with two, midline yellow spots in male (Fig. 11E, F). Aver- age size of body, 13.95 mm (male) and 17.32 mm (female) (Table 5). Fastigium of vertex flat, punctate; interocular space with a small medial depression. Medial carina of pronotum very weak or ab- sent; metazona short, about one-third the length of prozona, and 22% of the pronotum (Fig. 12E, F); metasternal space open, about half the width of the mesosternal space (Fig. 12B). Micropterous: elytra very small, roughly circular, not extending beyond the meso- thorax. Cerci strongly compressed laterally, slightly incurved, acute apex, exceeding the tip of subgenital plate. Outer lophi of epiphal- lus large, tilted rearwards, bifid at their tip; inner lophi absent (Fig. 12C, D), anterior processes well-developed (Fig. 12C, D). Description. —(Figs 11, 12, Tables 5, 7). Male. Length of body 13-14 mm; fastigium of vertex short, about twice as wide as long, roughly pentagonal in dorsal view, flat, punctate; frons slightly oblique; eyes large, oval, and convex; in- terocular space equal to or slightly wider than antenna scape with a small medial depression. Pronotum (Fig. 12E, F) medial carina very weak or absent; space between sulci 2 and 3 almost equal to the length of the metazona; metazona short, about one-third the length of prozona and 22% of the pronotum; posterior margin of metazona slightly embayed in the midline. Anterior margin of pro- notum slightly convex; micropterous: elytra very small, roughly cir- cular, not extending beyond the mesothorax laterally, not touching at the dorsal midline; prosternal process conical and pointed; mes- osternal space (Fig. 12B) as wide as long; metasternal space open, about half the width of the mesosternal space; hind femur slender (F/FD = 4.5). Internal tibial spurs twice as long as external spurs. Tympanum oval; last abdominal tergite incompletely divided (Fig. 12A), the hind margin with very small lobiform processes forming a minute furcula near the midline; cercus long, slightly incurved, strongly compressed laterally with acute apex, extending beyond the subgenital plate; supra-anal plate triangular in dorsal view with acute apex; there is a short medial groove proximally bounded pos- teriorly by a traverse ridge that extends across the width of the plate. Subgenital plate short, rounded. The phallus valvular plate is rela- tively simple, with a short quadratic “anvil-like” process dorsally and paired processes distally. Epiphallus (Fig. 12C, D) remarkable, with large backwardly tilted bifid outer lophi and well-developed anterior processes; “oval” sclerites of irregular strap-like shape. Female. In general, similar to the male. Female particularities: Length of body 15-18 mm; the last abdominal tergite divided, but the hind margin lacks a furcula; cercus is long and straight, compressed laterally with acute apex, equalling or slightly ex- ceeding the supra-anal plate; supra-anal plate triangular in dorsal view, elongated, the apex rounded. Ovipositor valves slender and straight, upper surface of dorsal valves slightly grooved. Sexual dimorphism.—P male/P female = 0.79. After normalisation using this factor, it is seen (Table 5) that the male has proportionate- ly much longer antennae and very slightly more protuberant eyes. Coloration.—(Fig. 11E-H). Body multicolored in both sexes, predominantly brown in dorsal view. Antennal scape and pedicel green, flagellum mostly brown JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) C. OUMAROU NGOUTE AND C.H.F ROWELL 139 Fig. 10. Parapterotiltus minimus (Ramme, 1929). Female terminalia. A. Dorsal view; B. Ventral view, subgenital plate and egg guide shaded; C. Lateral view, subgenital plate and egg-guide shaded; D, E. Pterotiltus impennis showing differing structure of subgenital plate in that genus, ovipositor valves slightly separated for clarity (not to same scale as A-C); D. Lateral view; E. Ventral view. Scale bar: 1 mm. but green basally and sometimes yellow apically; fastigium of vertex green; vertex green-blue or black; eyes black in life, turning brown or brown-yellow when dried; frons green; genae green-blue or black. Pronotal disc brown, with two yellow patches in the anterior part of prozona and two more (sometimes confluent) in the posterior part of metazona; pronotal lateral lobes mainly brown, but in life with two prominent yellow patches at the anterior and posterior ventral angles, sometimes confluent to form yellow ventral stripe that can extend on to the mesothoracic and metathoracic pleura; these yellow markings tend to fade to brown in dried specimens. Elytra yellow; mesothoracic and metathoracic tergites brown; fore and middle femora brown; hind femur green, hind knee blue with a black upper lobe, tibia and tarsus green. Abdominal tergites 1 and 2 brown with two yellow patches at the posterior margin; tergite 3 entirely or only laterally blackish; in the latter case, the medial part is brown. All other abdominal segments green. Distribution. —Cameroon: Makénéné: 4°53'58"N, 10°47'59"E, altitude 790 m; Bangangté: 5°09'030"N, 10°31'58"E, altitude 1325 m; Bamenda: 5°57'35"N, 10°08'50"E, altitude 1598 m. Remarks.—Parapterotiltus minimoides is almost indistinguishable from Parapterotiltus minimus in coloration, hence its specific name. Typically, they differ only in the number and placement of yel- low spots on the thoracic and abdominal terga in males: in P. minimus, each segment normally has only a single, midline yel- low spot, whereas the spots are paired in P. minimoides. However, there is considerable individual variation. Morphologically, the two species are more distinct: the metazona is shorter in P. mini- mus (17% of pronotal length) than in P. minimoides (22% of the pronotum length). In minimus, the metasternal interspace is very small and narrowly open, while this space is clearly open in mini- moides, at about half the width of the mesosternal space. The cerci of minimus are shorter, not exceeding the tip of subgenital plate, while the cerci extend beyond the tip of subgenital plate in mini- moides. The epiphalli, however, are dramatically different. P. mini- moides is unique within its genus in having large backwardly tilted bifid lophi, carried on a fairly wide bridge with well-developed anterior processes. In P. minimus, the bridge is reduced in size, anterior processes are absent, and the lophi are long but simple, pointed, and horn-like. Unexpectedly, these two outwardly very JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) 140 C. OUMAROU NGOUTE AND C.H.F ROWELL Table 4. Measurements of Parapterotiltus minimus. P: length of the pronotum in the midline; L: overall length from the tip of the fasti- gium to the most posterior part of the abdomen measured in lateral view; Ant: length of antenna from the scape to the tip of antenna; IOS: inter-ocular space measured dorsally; E-E: total ocular width measured dorsally; F: maximum length of the hind femur; FD: depth of hind femur measured as the distance between the two parallel lines drawn parallel to the long axis of the femur, running through the dorsal and ventral extremities of the femur; Tal, Ta2, Ta3: lengths of each of the three hind tarsal segments; Sp Ext: number of external spines on the hind tibia; Sp Int: number of internal spines on the hind tibia. Measurements and ratios are given as the range and aver- age of individual measurements and are also presented after normalization [obtained by dividing the measured value by P (the length of the pronotum)]. The foot formula shows relative lengths of the three tarsal segments within the total length of the foot and again allows for comparison between different species. N is the number of specimens analyzed/measured. Male Dimensionsinmm P L Ant IOS E-E F ED Tal Ta2 Ta3 Tal+2+3 External spines External spines Minimum Poy 215s O00" SUBS" 3253," 49:05% 87" 0-86 0.43 1.50 29) 4 9 Maximum 2:63, 14,317 “9353-" T0AL “SAS 9.51" —=2500-~ *1:00 0.43 1.50 253 rs 9 Average QAD 13.23, 9727 038 BtL6- 9,28 1.94. —-6:93 0.43 1.50 2.86 7 5 N 2. 2 2 yi 2. 2 2 2. 2 2 2 2 2. Foot formula 0.33 0.15 0.52 Ratios L/P Ant/P IOS/P E-E/P Tal/P Ta2/P Ta3/P Tal+2+3/P F/FD Tal+2+3/F Minimum S017 3 49ee Oa Qe O37 8 “O.l6n “O57 1.11 4.76 0.53 Maximum 5.44 406 0.16 1.33 038 0.18 0.64 1.19 4.84 0.55 Average 533i 3174. “OS 8h27 WBZ O87 20:60 TS 4.80 0.54 N 2 2 2 2 2 2 2! 2 2 a Female Dimensionsinmm P L Ant IOS E-E F FD Tal Ta2 Ta3 Tal+2+3 External spines External spines Minimum 3.50 «21.56 947 ~0.73 3.78 RIO 25ST LEO 0.48 1.94 3.61 az 9 Maximum 3.50. 21.61 10,22 .0.88° 3:89 —(12:10+ 2:51 1.19 0.48 1.94 3.61 7 9 Average 3150: «2159. “9:85r OG . B84 gl2t10" 2:51 119 0.48 1.94 3.61 7 i) N 2 2 2 2 2 2 2 2 2 2 2 2 2 Foot formula 0.33 0.14 0.53 Ratios L/P Ant/P IOS/P E-E/P Tal/P Ta2/P Ta3/P Tal+2+3/P F/FD Tal+2+3/F Minimum 6.16. 271 “0x21 1.08 0.34 0.14 0.55 1.03 4.63 0.40 Maximum Gil7 3292" 025 It “O03 "0.1'5- “058 1.09 4.82 0.41 Average G7 e231. 0:23. _ Iel0. 2035" LOn4 0057 1.06 4.72 0.41 N 2 2 2 2 2 2 2 2 2 2 Sexual dimorphism Average male/ 0.71 0.61 O94 047 O82 O.75 £40.74 0.75 0.85 0.76 0.77 average female Normalized on 71% 1.00 0.86 1.33 0.66 1.16 1.06 1.05 1.06 1.20 1.07 1.08 Male/female ratio 0°86; 133° 0.66, TAG" 31.06; 20: 41.07 1.08 1.02 1.33 similar species occur together at some localities (Banganté and Bamenda) between 1000 and 1600 m of altitude (Fig. 1). Parapterotiltus bamboutos sp. nov. https://zoobank.org/BCF83209-4B55-41C4-9E52-82EE26754CAD Figs 1, 11E-H, 13-15, Tables 6, 7 Material examined.—Holotype: CAMEROON e <4; Bamboutos Di- vision, Mount Bamboutos; altitude 2400 m; 15 Nov. 2017; C. Ou- marou Ngoute leg.; specimen no. 2017246; MfN. Paratype: CAM- EROON e 19; same data as holotype; specimen no. 2017245; Mf£N. Other material examined: CAMEROON e 2, 49; same data as holotype; personal collection of C. Oumarou Ngoute. Type locality. Cameroon, Bamboutos Division, Mount Bamboutos. Etymology.—Toponymic, referring to the Mount Bamboutos, Cameroon. Diagnosis.—Color of body as in Fig. 11A-D: multicolored, pre- dominantly black in male and olive-brown or green in female. Thoracic and anterior abdominal terga black: each segment with two midline yellow spots in male (Fig. 11A, B). Average size of body 12.74 mm (male) and 16.83 mm (female) (Table 6). Fastigi- um of vertex slightly concave. Pronotal medial carina very weak or absent; metazona short, accounting for only 19.5% of the length of pronotum (Fig. 13E, F); metasternal space open, about half the width of mesosternal space (Fig. 13B). Micropterous: elytra minute, rounded, barely exceeding the mesothoracic segment. Cerci strongly compressed laterally, slightly incurved, acute apex, extending beyond the tip of the supra-anal plate and sometimes exceeding the tip of the subgenital plate as well. Outer lophi of epiphallus large, horn-like, and pointed; a single weak pair of in- ner lophi (Fig. 13C, D); anterior processes absent (Fig. 13C, D). Description.—(Figs 11, 13-15; Tables 6, 7) Male. Average length of body 12.7 mm; fastigium of vertex short, roughly pentagonal in dorsal view, slightly concave in the middle; frons slightly oblique, frontal ridge shallowly sulcate; eyes large, convex, interocular space equal to or slightly wider than an- tenna scape. Pronotum (Fig. 13E, F) medial carina very weak or absent; the space between sulci 2 and 3 almost equal to the meta- zona in length; metazona short, accounting for only 20% of the length of pronotum; posterior margin of metazona slightly convex JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) C. OUMAROU NGOUTE AND C.H.E ROWELL ee - 7 Fig. 11. Parapterotiltus bamboutos sp. nov. and Parapterotiltus minimoides sp. nov. habitus. A, B. P. bamboutos male; C, D. P. bamboutos female, showing both brown and green color forms; E, FE. P. minimoides male; G, H. P. minimoides female. JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) 142 C. OUMAROU NGOUTE AND C.H.FE ROWELL E POST F Fig. 12. Parapterotiltus minimoides sp. nov. A. Dorsal view of male terminalia; B. Thoracic sterna. C, D. Epiphallus: C. axial view; D. dor- sal view; E. Pronotal disc showing sulci; F. Pronotal disc showing coloration and patterning in life. Scale bar: 1 mm. in female and slightly notched in midline in the male. Anterior margin of pronotum slightly convex; prosternal process conical, vertical, and pointed; mesosternal space (Fig. 13B) as wide as long; metasternal space open, about half the width of mesosternal space; micropterous: elytra minute, rounded, barely exceeding the mesothoracic segment. Hind femur slender, 4.7 times longer than wide (Table 6). Paired internal tibial spurs nearly twice as long as the external spurs. Tympanum oval or circular; last abdomi- nal tergite divided (Fig. 13A), the hind margin with very small lobiform processes forming a minute furcula, almost invisible, tips separated by 0.45 mm; cercus strongly compressed laterally, long, slightly incurved, with acute apex extending beyond the tip of the supra-anal plate and sometimes exceeding the tip of the subgenital plate as well; supra-anal plate triangular in dorsal view, very short, the apex angular. Proximally in the supra-anal plate, there is a short medial longitudinal groove bounded posteriorly by a curved transverse ridge that extends across the width of the plate. Epiphallus (Fig. 13C, D) with a reduced bridge and very large, horn-like pointed outer lophi and a single weak pair of inner lophi, ancorae absent. Female. In general, similar to the male. Female particularities: Length of body 16-18 mm, posterior margin of metazona slightly convex; last abdominal tergite short, divided, but the hind margin without a furcula. Cercus slightly compressed laterally, long, straight with acute apex, equalling or slightly exceeding the supra-anal plate; supra-anal plate triangular in dorsal view, elongated, the apex lin- gulate and rounded. Dorsal ovipositor valves laterally compressed with rounded tips, upper surface of dorsal valves slightly grooved; ventral valves slender and straight, rodlike. Egg guide rod-like, pro- jecting below ventral valves, acutely pointed (Fig. 14). JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) C. OUMAROU NGOUTE AND C.H.E ROWELL 143 Table 5. Measurements of Parapterotiltus minimoides sp. nov. P: length of the pronotum in the midline; L: overall length from the tip of the fastigium to the most posterior part of the abdomen measured in lateral view; Ant: length of antenna from the scape to the tip of antenna; IOS: inter-ocular space measured dorsally; E-E: total ocular width measured dorsally; F: maximum length of the hind femur; FD: depth of hind femur measured as the distance between the two parallel lines drawn parallel to the long axis of the femur, running through the dorsal and ventral extremities of the femur; Tal, Ta2, Ta3: lengths of each of the three hind tarsal segments; Sp Ext: number of external spines on the hind tibia; Sp Int: number of internal spines on the hind tibia. Measurements and ratios are given as the range and average of individual measurements and are also presented after normalization [obtained by dividing the measured value by P (the length of the pronotum)]|. The foot formula shows relative lengths of the three tarsal segments within the total length of the foot and again allows for comparison between different species. N is the number of specimens analyzed/measured. Male a - Dimensions in mm P. L Ant IOS E-E F FD Tal Ta2 Ta3 Tal+2+3 External spines External spines Minimum 2.44 13.36 9.87 0.34 3.34 9.09 1.97 0.84 0.34 1.51 2.89 7 9 Maximum 3.07 14.64 1062 0.53 3.54 10.38 2.26 1.07 0.61 2.08 3.50 7 9 Average 2.84 13.95 987 044 344 9.72 2.14 0.96 048 1.74 3°19 7 9 N 11 11 6 11 11 11 11 10 10 10 10 10 10 Foot formula 0.30 0.15 0.55 Ratios L/P Ant/P IOS/P E-E/P F/P FD/P Tal/P Ta2/P Ta3/P Tal+2+3/P F/FD Tal+2+3/F Minimum 4.43 3.31 Oe “O57 2307" £064 O30" “Osh. 8:0:50 0.98 4.04 O29 Maximum 5.81 3.64 0.19 1.22 4.05 0.86 0.39 0.20 0.73 122 5.06 0.37 Average 494 346 0.16 1.09 3.43 0.76 O34 O17 0.62 Lag 4.55 0.33 N 11 6 11 11 11 11 10 10 10 10 11 10 Female Dimensions in mm P bs Ant IOS E-E F FD Tal Ta2 Ta3 Tal+2+3 External spines External spines Minimum 3.40 15.62 7.78 049 3.44 10.71 2.25 0.87 40.53 1.79 3.23 7 9 Maximum 3.76 18217 28.82 —0:68" 896 1165 2:69) “1:387 ~0:60 -2,06 3.87 7 9 Average 3:59: “hyas2 ESI” S056", 364°. d1O3 2.50" WeOSt, 06> =1.95 3°55 7 9 N 6 6 3 6 6 6 6 6 6 6 6 6 Foot formula O30) [0°15 90:55 Ratios L/P Ant/P IOS/P E-E/P F/P FD/P Tal/P Ta2/P Ta3/P Tal+2+3/P F/FD Tal+2+3/F Minimum 459 2.24 014 1.004 2.99 063 0.24 0.16 0.55 0.95 4.18 029 Maximum A.90: 9248 ~“O0Z- «109° =3.23) ©0:72) 030 0.16 “0,58 1.03 4.83 0.34 Average 4.75 2.34 0.16 1.08 3.12 069 0.29 0.16 0.54 1.00 4.51 082 N 6 5 6 6 6 6 6 6 6 6 6 Sexual dimorphism Average male/ 0.79 0.81 120 _ 0.78 ~0:90- 0:87-0:86 “0:92 -0:86. .0.90 0.90 average female Normalized on 79% 0.99 1.01 LO” 408. wi S08 EOF) “if s08° > 12 Deh? Male/female ratio 104 148 0.99 1.01 lO: TROD IES OOS) Ths 1.13 1.01 103 Sexual dimorphism.—P male/P female = 0.76. Antennae of male are much longer than those of the female, and the male also has slightly longer feet, more protuberant eyes, and a smaller interocular space. Coloration.—(Fig. 11A-D). Male. Body multicolored, predominantly black; scape and pedicel green, flagellum green proximally with more apical seg- ments orange or brown. Fastigium green to blackish green; vertex black, tinged with green in life; eyes black in life, fading to brown or brown-yellow when dried. Frons in male green above medial ocellus and white below; genae black in upper half and white in inferior half. Pronotal disc black with two yellowish patches at the anterior margin of prozona and two more at posterior mar- gin of metazona. Elytra yellow or whitish; mesothoracic and metathoracic tergites black. Pronotal lateral lobes black, with a ventral whitish stripe prolonging the white genal stripe and con- tinuing on to the mesothoracic pleura. Fore and middle femora orange-brown, tibiae and tarsi green in life, fading to brown or green-brown when dried; hind femur dark green; hind knee black, with the upper lobe black, lower lobe very dark green; hind tibia and tarsus greenish, condyle of tibia black. Tergites of abdominal segments 1, 2, and 3 black, with paired yellow patches at their posterior margins; 4" abdominal tergite black in the anterior part, otherwise green; all other abdominal segments green. Female. Green or brown (either a polymorphism or a poly- phenism). Body multicolored, predominantly olive-brown or green. Scape and pedicel green, flagellum green with apical seg- ments orange or brown; fastigium brown-green or green. Vertex black or brownish green with two ill-defined paler brown longi- tudinal bands running behind the eyes; eyes black in life, fading to brown or brown-yellow when dried. Frons entirely brown or green; genae black or green in upper half and light brown in in- ferior half. Pronotal disc brown-green or green. Elytra brown or orange; mesothoracic and metathoracic tergites brown-green or green. Fore and middle femora brown or green, tibia and tar- sus green in life fading brown or green-brown when dried; hind femur entirely olive-brown in brown females; in green female, fe- mur brown apically while basal part is green; hind knee brown, with the upper lobe black; hind tibia and tarsus blue in both color forms. Abdomen entirely brown-green or green. Distribution.—Mount Bamboutos: 5°43'60"N, 10° 4’ 0” E, altitude 2400 m. P. bamboutos is known, at the moment, only from Mount Bamboutos (Fig. 1), hence its specific name. The population there is JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) 144 C. OUMAROU NGOUTE AND C.H.E ROWELL Table 6. Measurements of Parapterotiltus bamboutos sp. nov. P: length of the pronotum in the midline; L: overall length from the tip of the fastigium to the most posterior part of the abdomen measured in lateral view; Ant: length of antenna from the scape to the tip of antenna; IOS: inter-ocular space measured dorsally; E-E: total ocular width measured dorsally; F: maximum length of the hind femur; FD: depth of hind femur measured as the distance between the two parallel lines drawn parallel to the long axis of the femur, running through the dorsal and ventral extremities of the femur; Tal, Ta2, Ta3: lengths of each of the three hind tarsal segments; Sp Ext: number of external spines on the hind tibia; Sp Int: number of internal spines on the hind tibia. Measurements and ratios are given as the range and average of individual measurements and are also presented after normalization [obtained by dividing the measured value by P (the length of the pronotum)]. The foot formula shows relative lengths of the three tarsal segments within the total length of the foot and again allows for comparison between different species. N is the number of specimens analyzed/measured. Male a Dimensions in mm IP L Ant IOS E-E F FD Tal Ta2 Ta3 Tal+2+3 External spines External spines Minimum Dol T2298") 6:57 (02809 3322, 6909" SLI TOi83? 045" “47 3.02 7 | Maximum 274 13337 +8045 «Oly -3:33, 9:56" 2505 | 15055 “O17 - “2212 3157 7 9 Average 2364~ “1274 T52 O35 “3.27. “9:35 °2.00,. 0,94 051 Lvs 3.24 7 9 N 3 3 3 3 3 3 3 3 3 3 3 3 Foot formula 0:29 0:16. “0:55 Ratios L/P Ant/P IOS/P E-E/P F/P FD/P Tal/P Ta2/P Ta3/P Tal+2+3/P F/FD Tal+2+3/F Minimum 459 e240 BOLO: “11225 §3.32"0:735 "0319 O16. "0:64 1.14 4.57 0.34 Maximum SOL e297" BOIS 12229" eSS e077 8 O36- Oat et 1.34 4.77 0:37 Average A483 ‘2:86 °O0:13 1.22 .3.55 0:76 0:34 0:19. .0:68 1,23 4.67 0.35 N 3 a 3 3 3 3 3 3 3 3 3 Female Dimensions in mm i? iB Ant IOS E-E F FD Tal Ta2 Ta3 Tal+2+3 External spines External spines Minimum Sob7~ "15.83" S90. 0354 “3.46. 11005 2227 -OF73 0:53 = 1.88 3.50 7; 9 Maximum Bio 82S § 5905 S005. A062! L230" 2-70» 9 1932> O72" $2356 3.96 7 9 Average 3.48. <1633° 35.90 0:60 23.68. "11,53" 2427 1709. “O:G0, 22.05 But @ 3) N 5 5 1 5 5 5 5 5 5 5 5 5 Foot formula 0:29, “O16 v0.55 Ratios L/P Ant/P IOS/P E-E/P F/P FD/P Tal/P Ta2/P Ta3/P Tal+2+3/P F/FD Tal+2+3/F Minimum 4.59 1.71 O15 1.002 3.20 066 0.21 O14 40.54 1.01 4.54 0.31 Maximum 5.26 “1,71 “O19 1,09» 347 0.72 ~OAI 0.21 0.63 1.17 4.98 0.34 Average 4.85 ele FOE AKOGS §3232" 0270) "4032 O17 20°59 1.08 4.78 0.33 N 5 1 5 5 5 5 5 5 5 5 5 Sexual dimorphism Average male/ O76'5 O46. 1227 (0.58. °0;89" 0:31 . 083° 036% 0:84h -0.87 0.86 average female Normalized on 76% 1.00 1.00 1.68 0.76 1.17 4.1.07 1.09 1.13 Lait 1.15 1.14 Male/female ratio OO, Mu6e “OMe TZ* MOF = SIO mcd. i! | ae: 1.14 0.98 1.06 very small, and it is difficult to find a male in the field. Mount Bam- boutos is one of the most degraded mountains in Cameroon; its human populations have destroyed almost all the pockets of forests present on the mountain and replaced them with crops. Conserva- tion measures are urgent for the habitat of this species, which may be endemic to this mountain. However, Mount Bamboutos belongs to the volcanic line of Cameroon. It is possible that this species also occurs at altitude (more than 2000 m above sea level) elsewhere in that line, for example on Mounts Kupe and Manengouba, which are not far from the type locality. A photograph from Mt. Kupe taken by Dr. Ulf Bjelke supports this suggestion (Fig. 15). Remarks.—Parapterotiltus bamboutos and Parapterotiltus minimus are superficially very different. Body is predominantly brown in both sexes of P. minimus, while it is blackish in males and olive-brown or green in females of P. bamboutos (Table 7). However, the two species are similar in other respects, and here, we summarize the morphological differences: The metazona is shorter in P. minimus (17% of pronotal length) than in P. bamboutos (20% of pronotum length). In minimus, metasternal interspace is very small and nar- rowly open, while this space in bamboutos is clearly open, about half the width of the mesosternal space. The cerci of minimus are shorter, not exceeding the tip of the subgenital plate, while the cer- ci are longer in bamboutos, extending beyond the tip of the supra- anal plate and sometimes exceeding the tip of the subgenital plate (Table 7). The epiphallus of P. bamboutos is similar to that of P. minimus (Ramme, 1929). Both species have long horn-like pointed outer lophi, but P. bamboutos is unique within the genus for having an additional pair of weak inner lophi that are absent in minimus. Discussion The known species of Pterotiltus are morphologically very homogenous, mostly differing only in size and coloration. Our examination shows that while their phalli are also superficially similar, the species differ markedly in the structure of the epiphal- lus and the valvular plate [see also Hollis (1975) and Rowell (2005)|. The present work also shows that some populations in the Western Cameroonian highlands have differentiated enough from their lowland congeners that they can be considered a sepa- rate, though clearly closely related, genus, Parapterotiltus gen. nov. The two genera are similar, and it is plausible that one is evolu- tionarily derived from the other or that they share an immediate common ancestor. While their differences in size, degree of wing JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) C. OUMAROU NGOUTE AND C.H.E ROWELL reduction, and coloration would not alone justify generic status, the differences in the male epiphalli and the female subgenital plates are considerable. In Pterotiltus spp., the female subgenital plate is simple and the epiphallus has long, slender, blade-like, and forwardly curved outer lophi, while in Parapterotiltus spp., the female subgenital plate has two robust posteriorly directed pro- jections and the epiphallic outer lophi are long, upwardly and inwardly curved, and either pointed and “horn-like” or bifid at their tips. The highland region in which Parapterotiltus spp. occur has not been intensively collected, and more taxa may be awaiting discovery there. In Pterotiltus and Parapterotiltus, the external morphology is al- most homogenous, the species principally differing outwardly in size and coloration. In these genera, the species are separated on the basis of only a few characters and fundamentally by the struc- ture of the epiphallus. The most important references about the Oxyinae genera are the papers of Hollis (1971 and 1975), which revised the genus Oxya Serville, 1831 and reviewed the subfam- ily. Hollis (1971) separated approximately 20 Oxya species from various parts of the world; he based this partially on the structure of the valvular plate but mainly on the forms of outer and inner E POST Fig. 13. Parapterotiltus bamboutos sp. nov. A. Dorsal view of male terminalia; B. Thoracic sterna; C, D. Epiphallus: C. Axial view; D. Dorsal view; E. Pronotal disc showing sulci; EF. Pronotal disc showing coloration and patterning in life. Scale bar: 1 mm 145 lophi, posterior and anterior processes, and bridge, all of which are ephiphallic structures. Examination of the valvular plate often necessitates a destructive dissection, which is problematic if the available material is limited. Given the present work on Pterotiltus and Parapterotiltus, it is clear why Hollis (1971, 1975) concentrat- ed on the epiphallus, which is very diverse within the subfamily, cercus paraproct dorsal valve ventral valve subgenital plate egg-guide Fig. 14. Parapterotiltus bamboutos sp. nov. Female terminalia to show subgenital plate and egg-guide (shaded). The ovipositor valves have been slightly separated for clarity. Scale bar, 1 mm. Fig. 15. Parapterotiltus sp., probably P. bamboutos sp. nov., photo- graphed on Mt. Kupe by Dr. Ulf Bjelke in 2010. This picture is the best evidence to date that this species is not confined to Mt. Bamboutos. JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1) 146 C. OUMAROU NGOUTE AND C.H.F ROWELL Table 7. Tabular key to the species of the genus Parapterotiltus (males). Characters Parapterotiltus minimus Parapterotiltus minimoides Parapterotiltus bamboutos Color of body Thoracic and abdominal terga brown: Thoracic and abdominal terga brown: =‘ Thoracic and abdominal terga black: each each segment usually with a single, | each segment with two midline yellow segment with two midline yellow spots midline yellow spot (Fig. 8A). spots (Fig. 11E, F). (Fig. 11A, B). Vertex Fastigium of vertex slightly concave. Fastigium of vertex flat, punctate; Fastigium of vertex slightly concave. interocular with a small medial depression. Pronotum Medial carinae absent; metazona much _—— Medial carina very weak or absent; = Medial carina very weak or absent; metazona shorter than prozona, making up only metazona short, 22% of the pronotum short, accounting for only 19.5% of the 17% of pronotal length. (Fig. 12E, F). length of pronotum (Fig. 13E, F). Sternum Metasternal interspace very small, Metasternal space open, about half | Metasternal space open, about half the width narrowly open. the width of the mesosternal space of mesosternal space (Fig. 13B). (Fig. 12B). Tegmina Micropterous: very small with Micropterous: elytra very small, roughly Micropterous: elytra minute, rounded, barely rounded, almost spherical tip. circular, not extending beyond the exceeding the mesothoracic segment. mesothorax. Cerci Compressed laterally, acute apex, not Strongly compressed laterally, slightly Strongly compressed laterally, slightly incurved, exceeding the tip of subgenital plate. incurved, acute apex, extending beyond acute apex, extending beyond the tip of the the tip of subgenital plate. supra-anal plate and sometimes exceeding the tip of the subgenital plate as well. Lophi of epiphallus — Long pointed horn-like outer lophi. Large backwardly tilted bifid outer —_ Large, horn-like pointed outer lophi. A single Inner lophi absent (Fig. 9A, B). Anterior processes of Absent (Fig. 9A, B). epiphallus to define the species and even to establish the genera in Oxyinae while largely ignoring other characters. Acknowledgements We thank the Center for African Studies of the University of Basel in Switzerland and its director, Dr. Arlt Veit, for the admin- istrative and financial help that enabled us to carry out this work. We also thank Mr. G. R. Saroua and Dr. A. R. Nzoko-Fiemapong for help in sampling grasshoppers in Cameroon. Thanks are ex- tended to the Museum fiir Naturkunde, Berlin, for kindly lending us the holotype of Pterotiltus minimus, to Dr. Ulf Bjelke for permit- ting use of his photograph for Fig. 15, and to the Orthopterists’ Society for their support in the publication of this work. References Bolivar I (1905) Ortdpteros Acridioideos de la Guinea Espafiola. Memo- rias de la Real Sociedad Espanola de Historia Natural 1: 209-240. Brunner von Wattenwyl G (1893) Révision du Systeme des Orthopteres et description des espéces rapportées par M. Leonardo Fea de Birmanie. Annali del Museo civico di Storia naturale Giacomo Doria (2)13: 1-230. https://doi.org/10.5962/bh1.title.5121 Cigliano MM, Braun H, Eades DC, Otte D (2022) Orthoptera Species File. Version 5.0/5.0. http://Orthoptera.SpeciesFile.org [accessed October 04, 2022] Descamps M (1972) Geographical regions and taxonomic groups of Acri- domorpha in need of study. Proceedings of the International Study Conference. Current and Future Problems of Acridology, London 1970. Dirsh VM (1965) The African genera of Acridoidea. Anti-Locust Research Centre and Cambridge University Press, London, 579 pp. Dirsh VM (1970) Acridoidea of the Congo (Orthoptera). Annales de la Musée Royal de l'Afrique Centrale, Tervuren, Belgique, Serie IN-8°, Sciences Zoologiques 182, 605 pp. Hollis D (1971) A Preliminary Revision of the Genus Oxya Audinet-Serville (Orthoptera: Acridoidea). Bulletin of the British Museum of Natural History (Entomology) 26: 269-343. lophi. Inner lophi absent (Fig. 12C, D). Well-developed anterior processes (Fig. 12C, D). weak pair of inner lophi (Fig. 13C, D). Absent (Fig. 13C, D). Hollis D (1975) A review of the subfamily Oxyinae (Orthoptera: Acridoidea). Bulletin of the British Museum of Natural History (Entomology) 31: 189-234. _ https://doi.org/10.5962/bhl. part.29486 Jago ND (1964) Aspects of the ecology and distribution of grasshoppers in Ghana as a contribution to the zoogeography of West Africa. Journal of the West African Science Association 8: 190-204. Johnston HB (1956) Annotated Catalogue of African Grasshoppers. Cam- bridge University Press: Cambridge, 833 pp. Johnston HB (1968) Annotated Catalogue of African Grasshoppers. Sup- plement. Cambridge University Press: Cambridge, 448 pp. Karsch F (1891) Verzeichnis der von Herrn Dr. Paul Preuss in Kamerun erbeuteten Acridoideen. Berliner Entomologische Zeitschrift 36: 175-196. https://doi.org/10.1002/mmnd.18910360116 Karsch F (1893) Die Insekten der Berglandschaft Adeli im Hinterlande von Togo (Westafrika) nach dem von den Herren Hauptmann Eugen Kling (1888 und 1889) und Dr. Richard Bittner (1890 und 1891) gesammelten Materiale. I. Abtheilung: Apterygota, Odonata, Orthop- tera. Saltatoria, Lepidoptera Rhopalocera. Berliner Entomologische Zeitschrift 38: 1-266. Oumarou Ngoute C, Kekeunou S (2017) Redescription and diagnosis of the African genus Gemeneta Karsch, 1892 (Orthoptera: Acrididae: Cat- antopinae). Journal of Orthoptera Research 26: 161-170. https://doi. org/10.3897/jor.26.19995 Ramme W (1929) Afrikanische Acrididae. Revisionen und Beschreibungen wenig bekannter und neuer Gattungen und Arten. Mitteilungen des Zoologischen Museums Berlin 15: 247-492. Rehn JAG (1957) The grasshoppers and locusts (Acridoidea) of Australia. Vol. II. Family Acrididae. Melbourne: CSIRO. Rentz DCF (1996) Grasshopper country: The abundant orthopter- oid insects of Australia. Sydney: University of New South Wales Press. Rowell CHF (2005) A new Ugandan species of Pterotiltus (Orthop- tera, Acrididae, Oxyinae) with epiphyllic oviposition. Journal of Orthoptera Research 14: 33-43. https://doi.org/10.1665/1082- 6467(2005) 14[33:ANUSOP]2.0.CO;2 Stal C (1878) Systema Acridiodeorum. Essai d’une systematisation des Acridiodes I. Bihang till Kongl Svenska vetenskaps-akademiens han- dlingar 5, 100 pp. JOURNAL OF ORTHOPTERA RESEARCH 2024, 33(1)