Research Article |
Corresponding author: Claudia Hemp ( claudia.hemp@uni-bayreuth.de ) Academic editor: Susanne Randolf
© 2021 Claudia Hemp, Bruno Massa.
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.
Citation:
Hemp C, Massa B (2021) Biogeographical and evolutionary aspects of a Guineo-Congolian bushcricket tribe: Revision of the genera Cestromoecha Karsch, 1893 and Poreuomena Brunner von Wattenwyl, 1878, with the description of new species (Orthoptera, Tettigoniidae, Phaneropterinae). Deutsche Entomologische Zeitschrift 68(1): 45-79. https://doi.org/10.3897/dez.68.60193
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The genera Cestromoecha and Poreuomena of the tribe Poreuomenini in Phaneropterinae are revised and new generic characters are given for both genera, and six new species are described in Poreuomena. The newly described species are P. biaculeata sp. nov., P. eala sp. nov., P. gracilicercata sp. nov., P. ivoriana sp. nov., P. matthaei sp. nov., and P. tshuapa sp. nov. Based on characters defining the two genera, three species so far listed under Cestromoecha are transferred to Poreuomena: P. crassipes Karsch, 1890, P. laeglae (Massa, 2015), and P. magnicerca (Massa, 2013). One species of Cestromoecha, C. mundamensis Karsch, 1896, is synonymised with C. tenuipes (Karsch, 1890) since no morphological differences were detected between the type specimens. Thus, two species remain with Cestromoecha, and Poreuomena now contains 16 species.
Morphological closely-related species of Poreuomena suggest rapid speciation in the Congo Basin due to several expansions and shrinkages of the Guineo-Congolian forest belt since the Oligocene. At least two different morphological lineages are discernible. On the other hand the genus Cestromoecha Karsch, 1893 is a species-poor taxon.
taxonomy, identification characters, synonymies, biogeography, speciation
Information on Phaneropterinae and also other Orthoptera of Central to West Africa is sparse. Most research activity, mainly the collecting of specimens, reaches back to colonial times. More recent studies in parts of Africa were conducted by, for example,
In this study, we focus on the tribe Poreuomenini which includes the three genera, Cestromoecha Karsch, 1893, Poreuomena Brunner von Wattenwyl, 1878 and the monotypic Paraporeuomena Massa, 2018, tentatively included in the tribe. Currently, the genera Cestromoecha and Poreuomena contain 13 species, all from central-western Africa. We review the genera Poreuomena and Cestromoecha providing new characters on generic level since both genera were described only on the few male specimens present at that time only by Brunner von Wattenwyl (1878) and Karsch (1893). A biogeographical analysis is conducted on the basis of the morphology of the outer male genitalic apparatus.
Most specimens studied in the present paper were studied in museum collections, others were collected at nighttime attracted to light (UV) both on the ground and in the canopy of central-western countries of tropical Africa during scientific expeditions. Before mounting specimens, the left wing of every male having different cerci was spread in order to allow the study of the stridulatory file on the underside of the left forewing; the number and arrangement of the teeth are diagnostic and are useful characters to identify whether a species is bioacoustically separated from one another (
ANHRT African Natural History Research Trust, Hereford, UK.
BMPC Bruno Massa Collection, Palermo, Italy.
CCH Claudia Hemp Collection, Bayreuth, Germany.
NHM Natural History Museum (formerly British Museum Natural History), London, UK.
PAPC Philippe Annoyer Collection, Sainte Croix Volvestre, France.
The name Poreuomena is probably derived from the Greek verb Πορεύω meaning “bringing something into a definite direction”. The name Cestromoecha is probably derived from Κέστρον, an instrument used in pyrography (drawing by means of heat), pointed on one side and rounded on the other. The same name was used by the Macedonians to indicate a particular type of arrow; the word indicates something that has the form of κέστρον (κέστρον μοι έχειν).
Brunner von Wattenwyl (1878) erected the group Poreuomenae and the genus on the single species Poreuomena africana from Gabon, the only species known in this group at that time. Brunner von Wattenwyl (1878) noted that the new genus is similar in habitus to the genus Phaneroptera. Regarding differences to Phaneroptera both sides of the tympanal organs are closed in Poreuomena (not true for all newly-described species though, see below) while they are open in Phaneroptera. Furthermore, the fore tibiae are smooth without any furrow at their dorsal sides and completely unarmed. These characters differentiate the Poreuomenini from Phaneropterini and place them near to the Holochlorini (formerly divided into Holochlorae and Psyrae). Since Poreuomena was erected on the species africana generic characters were the 10th abdominal tergite of the males being extended in two lobes and the Rs vein branching off near the base of the tegmen.
Venation of Poreuomenini. Arrows point at Rs branching of from the radius 1. Rs branching off after the middle in Cestromoecha tenuipes. 2. Rs branching off basally, for example, in Poreuomena africana. 3–5. Mirror area of Cestromoecha tenuipes lacking the flaps on both tegmina bases, but with a well-developed mirror on the right tegmen (arrows). 5. Mirror area of Cestromoecha tenuipes from the underside with a well-developed stridulatory file, typical for all Poreuomenini.
Morphological characters of the genera Poreuomena and Cestromoecha. Rs: Position of Rs branching off from radius. Flaps: flaps on both tegmina developed or not. Mirror: mirror on right tegmen developed or not. 10th: 10th abdominal tergite “bilobed” (derived from the basic structure of an elongated 10th tergite with lateral bulges or short or long, straight or downcurved lateral processes) or undifferentiated or differently shaped.
Species | Rs | Flaps | Mirror | 10th abdominal tergite |
---|---|---|---|---|
Cestromoecha | ||||
C. longicerca Massa, 2013 | behind middle | no | large mirror | stout 10th tergite with almost straight posterior margin |
C. mundamensis Karsch, 1896 (synonym with the following species) | behind middle | no | small mirror | straight margin with median ridge |
C. tenuipes (Karsch, 1890) | behind middle | no | small mirror | undifferentiated, straight margin |
Poreuomena | ||||
P. africana B. v. Wattenwyyl, 1878 | basal | yes | no | bilobed: posteriorly produced with two thick downcurved processes |
P. biaculeata sp. nov. | basal | yes | no | bilobed: posteriorly produced with two laterally compressed downcurved lobes |
P. crassipes (Karsch, 1890) stat. nov. | basal | yes | no | bilobed; elongated downcurved flap forming two bulges |
P. duponti Griffini, 1908 | basal | yes | no | bilobed : posteriorly produced with two thick processes |
P. eala sp. nov. | basal | yes | no | bilobed: posteriorly produced with two bulgy lobes |
P. forcipata Sjöstedt, 1902 | basal | yes | no | bilobed: posteriorly produced with two thick somewhat laterally expanded and downcurved processes |
P. gracilicercata sp. nov. | basal | yes | no | bilobed: two stout bulges with long downcurved processes |
P. huxleyi Massa, 2013 | basal | yes | no | bilobed: posteriorly produced with short stout processes narrowing suddenly to a thin elongate, laterally compressed, flagellum-like shape |
P. laeglae (Massa, 2015) stat. nov. | basal | yes | no | two broad but narrow lobes with median gap |
P. lamottei Chopard, 1954 | basal | yes | no | bilobed: posteriorly produced with two stout and downcurved lateral processes |
P. magnicerca (Massa, 2013) stat. nov. | basal | yes | no | undifferentiated, straight margin |
P. matthaei sp. nov. | basal | bulge | no | broad, shield-like with median gap |
P. sanghensis Massa, 2013 | basal | yes | no | bilobed: broad with short lateral flanges forming a u-shaped gap between them |
P. tshuapa sp. nov. | basal | yes | no | broad with shallow median depression and two reduced knob-like processes |
P. wilverthi Griffini, 1908 | basal | yes | no | bilobed: broad with short lateral flanges |
Referring to the type species of Poreuomena, P. africana, and Cestromoecha, C. tenuipes, several species are currently misplaced and are here transferred to their respective genus on grounds of generic characters given above and below. A very obvious distinguishing character on generic level is the presence (Poreuomena africana) or absence (Cestromoecha tenuipes) of a mirror on the right side of the tegmen (Figs
Cestromoecha Karsch, 1893. Berlin Ent. Z. 38: 128; type species: Poreuomena tenuipes Karsch, 1890
Medium-sized, predominantly green, typical Phaneropterinae with narrow and elongate tegmina surpassed at their apices by the alae. Fastigium verticis smaller than the width of the scapus; triangular with rounded apex, shallowly sulcate, separated from conical fastigium frontis by a gap. Antennal sockets elevated beside fastigia. Fore coxa with a spine. Fore and mid femora dorsally rounded, ventrally with very tiny spinules. Hind femora slender, slightly thickened in basal part, with few tiny spinules along the ventral length. Fore tibiae very slender, thickened in the area of the tympana and slightly sulcate dorsally; tympana open on the inner, conchate on the outer side. Hind tibiae triangular or narrow rectangular in diameter, densely packed along each edge with slender spines; dorsally with a pair of tiny spurs on each side, ventrally with each one larger spur on each side. Left tegmen of typical Phaneropterine shape, with a slight bulge where the stridulatory file is located at the underside (Figs
Cestromoecha longicerca Massa, 2013. J. Orth. Res. 22: 142; type locality: Central African Republic, Dzanga-Ndoki National Park (BMPC).
Central African Republic, Dzanga-Ndoki National Park, Lake 1, UV trap, 11–12.II.2012 (holotype ♂); same data 20–23.II.2012 (paratype ♂) (BMCP); Central African Republic, Dzanga-Ndoki NP, Saline des Buffles, 25.I.2012 (1♂); Mboki, 25–26.I.2012 (light) (1♂); Lake 1, 14–15.II.2012, 19–20.II.2012 (light) (3♂) (PAPC). Gabon, Mikongo (Rougier), Mts de Cristal (secondary forest) (430 m alt.) 28.VII-12.VIII.2019 (MV Light Trap) (1♂) (ANHRT).
The stridulatory area of the left tegmen is short; it is slightly arched and has ca. 30 evenly-spaced teeth (Fig.
Green-brownish, tegmina green with a black marking on the stridulatory area (Figs
(mm). Males. Body length: 15.7–16.0; Pronotum length: 4.0–4.2; Pronotum height: 3.5–3.7; Length of hind femora: 17.4–17.5; Length of tegmina: 25.8–28.2.
The cerci of the male have a very stout base, with the inner part concave bending sharply upwards and becoming very narrow; the cerci surpass clearly the 10th tergite and the apices are very pointed (Figs
Dzanga-Ndoki National Park (Central African Republic) (
Cestromoecha tenuipes
Karsch, 1890. Entom. Nachricht. 16: 363; type locality: Cameroon, Barombi Station (
Syn. Cestromoecha mundamensis Karsch, 1896 syn. nov.
Cameroon, Barombi Station (2♀ syntypes) (
Equatorial Guinea, Fernando Poo, Musolo I. 1902, L. Fea (1♂, 1♀); Fernando Poo, Basile (1♀) (
Sjöstedt (1912) stated that Griffini (1905/6) found no differences between C. tenuipes and C. mundamensis, both described by Karsch. Sjöstedt mentions that a generic character on which Cestromoecha was erected, is a missing mirror on the right tegmen, while C. mundamensis has a well-developed mirror. However, both C. tenuipes (Fig.
Poreuomena Brunner von Wattenwyl, 1878. Monographie der Phaneropteriden 187; type species: Poreuomena africana Brunner von Wattenwyl, 1878
Medium-sized, predominantly green typical Phaneropterinae with narrow and elongate tegmina surpassed at their apices by the alae. Fastigium verticis smaller than the width of the scapus; triangular and sulcate above, separated from the conical fastigium frontis by a gap. Antennal sockets elevated beside fastigia. Fore coxa with a spine. Fore and mid femora dorsally rounded, ventrally with very tiny spinules. Hind femora slender, slightly thickened in basal part, with few tiny spinules along the ventral length. Fore tibiae very slender, thickened in the area of the tympana and sulcate dorsally; tympana open on inner, conchate on outer side. Hind tibiae triangular or narrow rectangular in diameter, densely packed along each edge with slender spines; dorsally with a pair of tiny spurs on each side, ventrally with one larger spur on each side. Bases of the tegmina differentiated into flaps; on each of these flaps, a similar shaped stridulatory file on the underside is present. No mirror developed on the right tegmen. Rs vein branching off from the radius basally. Male 10th abdominal tergite usually bilobate, either with evenly rounded lobes, lobes reduced to short bulges or strongly elongated. Male cerci simple, expanded at their tips or differentiated in differently-shaped branches. Subgenital plate elongated (but not markedly surpassing abdominal apex) with mostly a pair of short stout to more slender lobes or unlobed with two rounded apices at the posterior margin.
All checked specimens also have an area of dark cells surrounded by the green more elevated veins in the cubital area of the tegmina.
Poreuomena africana
Brunner von Wattenwyl, 1878. Monographie der Phaneropteriden, 187; type locality: Gabon (
Cameroon (1♂, 1♀) (MCNM); Cameroon, Campo Ma’an National Park (lowland rainforest) (950 m alt.) 10–22. III. 2018 (UV Cold Cathode Light Trap), Fotsing, Ishmael, Miles, Safian (2♂ in ANHRT; 1♂ in BMPC); Gabon, Mikongo (Rougier), Mts de Cristal (secondary forest) (430 m alt.) 0°29'47"N, 11°10'42"E, 28. VII-12. VIII.2019 (LepiLED Light Trap), Albert, Aristophanous, Bie Mba, Dérozier, Moretto (1♂) (ANHRT); Gabon, Mts de Cristal National Park, Kinguelé 3. XII. 2015 (UV) (BMPC); Gabon, M’Bigon 26. I. 1986, A Pauly (1♂) (
The 10th abdominal tergite (Fig.
Tegmina about 5.8–5.9 longer than broad. Stridulatory file 0.3 mm long; teeth more widely spaced at inner side, becoming more densely set distally (Fig.
Small species, uniformly green-yellowish, a little brownish around the stridulatory area (Fig.
(mm). Males. Body length: 18.5–19.9. Pronotum length: 3.7–3.8. Pronotum height: 2.8–2.9. Length of hind femora: 19.6–19.8. Length of tegmina: 26.5–26.8. Width of tegmina: 4.5–4.6.
Morphological details of male Poreuomena africana. Drawing Fig.
Democratic Republic of the Congo. Léopoldville (today Brazzaville) 1937, A. Tinant (holotype ♂) (
Very closely related to P. wilverthi, a species widespread in the Congo basin and the Albertine Rift. P. biaculeata sp. nov. has the 10th abdominal tergite deeply split into two lobes (Figs
Male. Typical Poreuomena species with wings protruding over the body by only a few mm (Fig.
Female. Unknown.
(mm). Male (n = 1). Body length: 22; pronotum length: 3.7; length hind femora: 21.2; length of tegmina: 28.3; width of tegmina: 4.3.
Congo Basin.
Named after the shape of cerci that are similar to two stings, from Latin biaculeatus (= with two stings).
Morphological details of male Poreuomena biaculeata sp. nov. Habitus (29), left tegminal flap with stridulatory file on the underside (30), stridulatory file on the underside of the left tegminal flap (31), abdominal apex in lateral (32) and rear view (33), cerci (34), subgenital plate (35).
Poreuomena crassipes
Karsch, 1890. Entom. Nachricht. 16: 364; Type locality: Cameroon (
Central African Republic, Dzanga-Ndoki National Park, Ndoki, Lake 1, UV trap 1, 31.I.-2.II.2012 (♂); 11–12.II.2012 (♂); 20–23.II.2012 (♂); Central African Republic, Dzanga-Ndoki NP, Lake 1, camp 1, 15–16.II.2012 (♂); Central African Republic, Dzanga-Ndoki NP, Lake 1, UV trap 2, 15–16.II.2012 (♂); Central African Republic, Dzanga-Ndoki NP, Lake 1, 11–12.II.2012 (light) (3♂); Lake 3, 25–26.II.2012 (light) (1♂) (BMPC); Central African Republic, Dzanga-Ndoki NP, Lake 1, 29–30.XI.2010 (1♂, 1♀); 30.XI-1.XII. 2010 (light) (1♂) (PAPC).
Morphologically closely related to P. biaculeata sp. nov. and P. wilverthi. All three species share a similar 10th abdominal tergite that is flap-like, deeply divided into two lobes in P. biaculeata sp. nov. (Figs
(mm). Males. Body length: 17.3–19.6; pronotum length: 3.8–4.1; pronotum height: 3.3–3.6; length of hind femora: 19.3–20.2; length of tegmina: 27.3–30.6.
P. crassipes is known from Cameroon (
Poreuomena duponti
Griffini, 1908. Mem. Soc. entom. Belgique, Bruxelles 15: 84; type locality: Cameroon, Mukonje Farm (
Cameroon, Mukonje Farm (♂ holotype, 4♀) (
The 10th abdominal tergite is hood-shaped with a small median indentation (Fig.
Democratic Republic of the Congo, Eala, IV 1935. J. Ghesquiére (Holotype ♂) (
Rather stout species with a 10th abdominal tergite differentiated into two bulges (Figs
Male. Stout species with wings projecting over the body by about half of their length. Where tegmina meet, when folded, interior part of cells of dark colour while surrounding and elevated veins green. Rs branching off in the first half of the basal part of the tegmen. Tegmina with two flaps at their base, the flap of the left wing smaller and more pointed than the broader flap with a rounded tip on the right tegmen. The stridulatory rib marked dark brown on the flap of the left tegmen, uniformly brown on the right tegmen. Beneath flaps, tegmina with narrow longish brown markings. Stridulatory file on the underside of the left flap about 1.3 mm long; teeth at apical part of the left flap very densely set and gradually getting larger, about from half of the file teeth large and more widely to very widely spaced; inner part then strongly curved and teeth becoming smaller and then obsolete; with about 45–50 teeth (Fig.
Female. As male comparatively stout, of uniform green colour without brown markings (Fig.
(mm). Males (n = 2). Body length: 22.7–23.1; pronotum length: 4.4–4.5; length hind femora: 22–22.5; length of tegmina: 30–31.5; width of tegmina: 4.4–5.2.
Females (n = 2). Body length: 19.5–22; pronotum length: 4.2–4.4; length of hind femora: 21–22.1; length of tegmina: 32–32.7; width of tegmina: 5.4–5.5; length of ovipositor: 6.4–6.5.
DRC, Équateur Province.
Named after the area Eala where specimens of this species were collected.
Morphological details of Poreuomena eala sp. nov. Habitus, female (45), subgenital plate (46) and lateral view on ovipositor (47), semi-lateral (48) and lateral (49) view on the male abdominal apex, stridulatory file on the underside of the male left tegminal flap (50), titillators (51).
Poreuomena forcipata
Sjöstedt, 1902. Bihang Kungl. Svenska Vet. Akad. Handl. 27 (3): 12; type locality: Cameroon (
Central African Republic, Dzanga-Ndoki NP, Lake 1, 14–15.II.2012, 19–20.II.2012 (light) (2♂) (BMPC).
Poreuomena gladiator
Bolívar, 1906. Mem. Soc. espan. Hist. nat. 1: 337; type locality: Cameroon (
Cameroon, holotype ♂ of P. gladiator (
Rather stout species with comparatively broad tegmina (Fig.
Cameroon and the Central African Republic.
Zaire (Democratic Republic of the Congo), 180 km W from Bukavu, rainforest, 14.V.1988, leg. A. Vojnits et al., Arthropoda collected at 160 W MV lamp, No. 320 (Teleki expedition) (Holotype ♂) (
The elongated and slender shape of the male cerci – thickened in the middle with an inner dent – and the 10th abdominal tergite forming two long downcurved processes are unique.
Male. Typical Poreuomena species with wings protruding over the body by only a few mm. Where tegmina meet when folded, interior part of cells of dark colour while surrounding and elevated veins green (or tawny in preserved insect). Rs branching off in the first half of the basal part of the tegmen. Tegmina with two flaps at their bases, the flap of the left wing smaller and more pointed than the broader flap with a rounded tip on the right tegmen. Left flap completely marked brown, right one partly brown, especially along the bulge of the stridulatory file on the underside (Fig.
Female. Unknown.
(mm). Males (n = 2). Body length: 18.2–18.6; pronotum length: 3.9–4.05; length hind femora: 19.3–19.6; length of tegmina: 28.6–29.15; tegmina width: 4.98–5.0.
Democratic Republic of the Congo.
Poreuomena huxleyi
Massa, 2013. J. Orth. Res. 22: 140; type locality: Equatorial Guinea, Fernando Poo, Santa Isabel (
Equatorial Guinea, Bioko (Fernando Poo), Santa Isabel (♂ holotype) (MCNM); Cameroon, Gulf of Guinea Is, Bioko (Fernando Poo), Santa Isabel (Malabo) (1♂) (
This species is easy to recognise by the short apical lobes on the 10th tergite of the male and cerci clearly upcurved and with a lateral spine. The most closely-related species is P. africana, which, however, has much longer lobes of the 10th abdominal tergite and more robust cerci.
Typical Poreuomena species with narrow tegmina surpassed by the alae by a few mm (Fig.
P. huxleyi is brownish coloured, with green tegmina and hind tibiae; a black marking is typical at the base of the tegmina and some small black spots on the posterior margin of the tegmina.
Côte d’Ivoire, Taï National Park, Research Station 22.III-4.IV.2017 (light) (♂ holotype) (BMPC).
P. ivoriana sp. nov. is a comparatively small species of Poreuomena, characterised by its green colour, the blackish triangular areas on the left and right tegmina (Fig.
Male. Green coloured with a faint blackish stripe on the stridulatory area; the triangular area close to the stridulatory file above the left tegmen and the corresponding area on the right tegmen are black. Antennae thin, fastigium of the vertex separated from the fastigium of the frons, face smooth, eyes round, prominent. Pronotum with a flat and smooth disc, the anterior margin straight, the posterior margin broadly rounded. Tegmina narrow, ca. 5.8 times broader than long, stridulatory area of the left tegmen short. The flap of the left tegmen comparatively broad (damaged in the holotype) (Fig.
(mm). Male. Body length: 19.0; pronotum length: 4.1; pronotum height: 2.3; length hind femora: 18.2; length of tegmina: 23.3; width of tegmina: 4.0.
P. ivoriana sp. nov. is named after the Côte d’Ivoire.
Presently known only from the Taï National Park (Côte d’Ivoire).
Poreuomena laeglae
Massa, 2015. ZooKeys 524: 38; type locality: Côte d’Ivoire, Tuba, Biémasso (
Côte d’Ivoire, Tuba, Biémasso (441 m) 9.VII.2014 (UV trap) (♂ holotype) (
P. laeglae was described in the genus Cestromoecha, but it actually belongs to the genus Poreuomena because it has two well-developed tegminal flaps and no mirror. The Rs vein branches off near the base and not past the middle as typical for Cestromoecha. It is morphologically most closely related to P. forcipata (both species share an asymmetrical supra-anal plate with a spine on the left side) and also to P. magnicerca and P. tshuapa sp. nov. The cerci of the males are differentiated into two branches in the first two species, with a finger-like apical or outer branch and a blade-like expanded inner branch, however, differently shaped between P. laeglae and P. forcipata. In P. magnicerca and P. tshuapa sp. nov., the third blade-like branch or expansion is present giving the expression of the cerci having three branches. The flaps on the left tegmina bearing the stridulatory files on the underside are differently shaped between P. tshuapa sp. nov. (oval and pointed) and P. magnicerca (laterally slightly expanded and not pointed) as is the shape of the stridulatory file itself supporting species status for P. tshuapa sp. nov. The 10th abdominal tergites, however, are very similar amongst these four species since they are rather undifferentiated with a more or less straight posterior margin and thus very likely derived from the bilobed condition as the generic character of the genus Poreuomena.
The stridulatory file of P. laeglae is similar to that of P. magnicerca with a distal part with less and more widely-spaced teeth than in the proximal part (
Typical Poreuomena species with elongated habitus. Fore and mid femora with 4–5 very small spines, fore tibiae with 3 ventral spines + 1 spur on each side, mid tibiae with 6–7 ventral spines + 1 spur on each side, hind tibiae with 3 spurs on each side. Ventral margins of hind femora with 2 small basal spines. Tegmina narrow, stridulatory area of left tegmen black and straight; stridulatory file downcurved with ca. 50 teeth, distal part with asymmetrical and widely-spaced teeth (Fig.
Probably predominantly green when alive, brown to tawny when preserved. Stridulatory area of left tegmen and area below black. Small black spots are present on the posterior margins of the tegmina. Two longitudinal parallel dark lines are present on the outer surface of the hind femora.
(mm). Males. Body length: 18.5–19.4; pronotum length: 4.0–4.2; pronotum height: 3.4–3.6; hind femur: 18.2–20.7; tegmina: 26.4–27.5.
Female. Body length: 21.7; pronotum length: 4.0; pronotum height: 3.4; hind femur: 20.8; tegmina: 29.4; ovipositor: 6.1.
Presently only known from the Ivory Coast and Liberia in West Africa.
Morphological details of male Poreuomena laeglae stat. nov. Head, pronotum and part of abdomen in lateral view (70), stridulatory file on the underside of the left tegminal flap (71), lateral view on cerci and 10th abdominal tergite (72), cerci lateral view (73), subgenital plate and cerci (74). Arrow in Fig.
Poreuomena lamottei
Chopard, 1954. Mem. Inst. franc. Afr. Noire 40(2): 40; type locality: Guinea, Nimba, N’zo (
Côte d’Ivoire, Azagny National Park (light trap) (3♂); Côte d’Ivoire, Taï National Park, Res. Station 5–10.VII.2015 (light trap) (1♂) (NHM).
P. lamottei has a similar 10th abdominal tergite as P. eala sp. nov., but the lobes are much narrower and have a large gap between them in P. lamottei. In addition, the male cerci are similar between the two species, stout and upcurved at their tips, however, with bifid tips in P. eala sp. nov.
Typical Poreuomena species with stridulatory area marked dark brown (Fig.
Guinea, Ghana (
Poreuomena magnicerca
Massa, 2013. J. Orth. Res. 22: 142; type locality: Central African Republic, Dzanga-Ndoki National Park (
Central African Republic, Dzanga-Ndoki National Park, Lake 1, UV trap, 6–8.II.2012 (Holotype ♂) (
Compare diagnoses at P. laeglae, P. tshuapa sp. nov., and P. forcipata.
P. magnicerca was described in the genus Cestromoecha, but actually belongs to the genus Poreuomena since sharing all generic characters with this genus and lacking a mirror on the right tegmen, typical for Cestromoecha. The stridulatory area of the left tegmen is black (Figs
Probably predominantly green when alive, brown to tawny when preserved. The stridulatory area of the left tegmen and the area below black. Small black spots are present on the posterior margins of the tegmina.
(mm). Males. Body length: 19.5–21.0; pronotum length: 3.9–4.0; pronotum height: 3.4–3.6; length of hind femora: 19.4–21.7; length of tegmina: 30.3–33.0.
Females. Body length: 17.7–21.0; pronotum length: 4.1–4.9; pronotum height: 3.1–4.0; length of hind femora: 18.8–21.7; length of tegmina: 27.7–29.2; width of tegmina: 4.6–5.0; length of ovipositor: 5.1–5.9.
Known only from the Central African Republic.
Gabon, Lope National Park, Ogooue-Ivindo 4.IV.2014, Ecotrop Team (♂ holotype) (BMPC).
P. matthaei sp. nov. is a comparatively large Poreuomena species, characterised by its yellowish colour, whitish triangular areas on the left and the right tegmina and incurved cerci with a flattened blackish tip. The 10th abdominal tergite is similar to that of P. gracilicercata sp. nov. Both species, however, have differently-shaped male cerci. While all other Poreuomena species have flaps at the bases of the tegmina, in P. matthaei sp. nov. only the area at the base of the right tegmen is developed as a flap, while on the right side, a bulge is present (Fig.
Male. Yellowish-cream coloured with a faint blackish stripe on the stridulatory area (Fig.
Female. Unknown.
(mm). Males. Body length: 23.6; pronotum length: 4.8; pronotum height: 2.7; length hind femora: 22.8; length of tegmina: 32.0; width of tegmina: 3.0.
P. matthaei sp. nov. is dedicated to the late Matteo Griggio, who at the age of 43 left us on 14 May 2020 due to an aneurysm. Matteo was a lively behavioural ecologist who was also very committed to nature conservation, curious about every particular aspect of nature, including Orthoptera as a food source for the Rock Sparrow in Sardinia.
Presently known only from the Lope National Park (Gabon).
Poreuomena sanghensis
Massa, 2013. J. Orth. Res. 22: 140; type locality: Central African Republic, Dzanga-Ndoki National Park (
Central African Republic, Dzanga-Ndoki National Park, Lake 1, UV trap, 10–12.II.2012 (holotype ♂) (
Small and fragile species, green (alive) or brown (preserved) coloured, with a brown-reddish upper area of the abdominal tergites. The 10th abdominal tergite of P. sanghensis is similar to P. wilverthi, however, not downcurved, but produced posteriorly with a deep median fold. The male cerci are also similar between these two species, however, much more fragile-built in P. sanghensis.
Typical Poreuomena species with an elongate habitus (Fig.
Predominantly green when alive, brown to tawny when preserved; dorsal area of abdominal tergites brown-reddish. A black marking is present at the base of the tegmina of the male, absent in the female.
(mm). Males. Body length: 15.2–17.5; pronotum length: 3.0–3.2; pronotum height: 2.5–2.7; length hind femora: 16.0–18.8; length of tegmina: 23.6–24.8.
Female. Body length: 19.4; pronotum length: 3.3; pronotum height: 2.8; length of hind femora: 19.2; length of tegmina: 30.0; length of ovipositor: 3.9.
Central African Republic (Dzanga-Ndoki National Park and Dzanga-Sangha Special Reserve), Cameroon (
Democratic Republic of the Congo, Tshuapa, Bokungu 1949, M. Dupuis (Holotype ♂); same data as holotype (Paratype ♂); Democratic Republic of the Congo, Haut-Lopori V-VI 1927, J. Ghesquiére (Paratype ♂) (
Similar in the outer male genitalic apparatus to P. magnicerca from further west in the Congo Basin. Differentiated from P. forcipata and P. laeglae by a different 10th abdominal tergite which is clearly divided into two lobes in P. forcipata, while P. tshuapa sp. nov. only has small humps and the posterior margin of the 10th abdominal tergite of P. laeglae is more-or-less straight. P. forcipata has male cerci differentiated into two branches, an outer rather blunt, finger-like part and an inner blade-like expanded branch with an acute tip, similar to the inner branch of P. tshuapa sp. nov. and P. magnicerca. However, P. tshuapa sp. nov. has an additional branch just below the finger-like part of the cerci, absent in P. forcipata, but also present in P. magnicerca (also see diagnosis at P. laeglae). In P. magnicerca, the blade-like subapical section or the “third” branch is more roundish and larger than in P. tshuapa sp. nov. and the subgenital plate differs between these two species. In P. tshuapa sp. nov., the subgenital plate is bilobate with the processes clearly separated, while in P. magnicerca, the lobes of the subgenital plate are shorter and situated more closely to each other. Further, in P. forcipata and P. laeglae, asymmetrical spines are present on the left side of the supra-anal plates.
Male. Probably predominantly green when alive, preserved specimens of tawny colour (Fig.
Female. Unknown.
(mm). Males (n = 3). Body length: 17.7–18.2; pronotum length: 3.8–4.1; length hind femora: 20.3–21.2; length of tegmina 29.6–30.4; width of tegmina: 4.6–5.0.
Poreuomena wilverthi
Griffini, 1908. Mem. Soc. entom. Belgique, Bruxelles 15: 85; type locality: Democratic Republic of Congo, Umangi (
Democratic Republic of Congo, Umangi IX-XI.1896 (♂ holotype) (
Male. Typical Poreuomena species with wings protruding over the body by only a few mm (Figs
Female. Unknown.
(mm). Males (n = 7). Body length: 17.2–19.5; pronotum length: 3.7–4.0; pronotum height: 2.6–3.0; length hind femora: 18.5–23.2; length of tegmina: 27.9–30.0. Width of tegmina: 3.8–5.1.
Widespread west of the Albertine rift into the area of the Central African Republic.
At present only the holotype was known. The new material studied coming from the Natural History Museum of Budapest and the Africamuseum Tervuren considerably enlarges the known area of distribution of this species (Fig.
1 | Tegmina broad, ratio length/width 4.6 | Paraporeuomena signata Massa, 2018 |
1´ | Tegmina narrow, ratio length/width 6 to 8.75 (Cestromoecha 7.25–7.4; Poreuomena 6.1 (P. wiverthi – 8.75 P. biaculeata sp. nov.)) | 2 |
2 | Mirror present, Rs branching off at the middle or past the middle of the tegmen... Cestromoecha | 3 |
2´ | Mirror absent, Rs branching off near the base of the tegmen... Poreuomena | 4 |
3 | Cerci strongly curved downward at base and then sharply bent upwards, at tips very acute (Fig. |
C. longicerca |
3´ | Cerci rather short and of normal shape, slightly in- and upbent at tips (Figs |
C. tenuipes |
4 | 10th abdominal tergite bilobed; cerci not differentiated into well-developed branches (sometimes with flattened tips) | 5 |
4´ | 10th abdominal tergite broad and with straight or slightly incurved posterior margin, but not bilobate; cerci differentiated into two or three apical branches | 16 |
5 | 10th abdominal tergite deeply divided into two lobes or into processes | 6 |
5´ | 10th abdominal tergite flap-like and not deeply divided into two lobes or processes | 9 |
6 | Processes of 10th abdominal tergite long and downcurved; cerci very elongated, but stout with an inner dent about middle (Figs |
P. gracilicercata sp. nov. |
6´ | Processes of 10th abdominal tergite much shorter and stouter; cerci not very elongated but of different shapes | 7 |
7 | 10th abdominal tergite with two short and stout processes from which each a lobe is arising (Fig. |
P. huxleyi |
7´ | 10th abdominal tergite with two processes without additional lobes | 8 |
8 | Processes of 10th abdominal tergite short and stout, only slightly downcurved (Fig. |
P. africana |
8´ | Processes of 10th abdominal tergite more slender, evenly downcurved (Fig. |
P. lamottei |
9 | Cerci with two stout tips | 10 |
9´ | Cerci pointed, flattened, or with bent apices but not differentiated into two tips | 11 |
10 | 10th abdominal tergite differentiated into two thickened bulges that are slightly downcurved (Fig. |
P. eala sp. nov. |
10´ | 10th abdominal tergite flap-like with median groove along length (Fig. |
P. biaculeata sp. nov. |
11 | Cerci slender, evenly tapering and with acute tips | 12 |
11´ | Cerci flattened, blunt or with acute angle at tips | 14 |
12 | Cerci very slender evenly tapering to tips; 10th abdominal tergite with elevated margins and deep median groove; medially at posterior margin with u-shaped incision with two acute tips at each side (Fig. |
P. sanghensis |
12´ | Cerci more stout along whole length with strongly narrowed acute tips (Figs |
P. wilverthi |
13 | Cerci laterally flattened | 14 |
13´ | Cerci with blunt or acute tips | 15 |
14 | Cerci flattened laterally in apical third, with an inner apical flange and a blunt outer tip (Fig. |
P. ivoriana sp. nov. |
14´ | Cerci flattened from base onwards, tips narrowed and curved, sclerotised and with acute tips (Fig. |
P. crassipes stat. nov. |
15 | Cerci with a strong angle at tips (Figs |
P. duponti |
15´ | Cerci with a blunt sclerotised tip, evenly incurved (Fig. |
P. matthaei sp. nov. |
16 | Cerci differentiated into two branches | 17 |
16´ | Cerci differentiated into three branches (Fig. |
P. tshuapa sp. nov. |
17 | Cerci with two branches and an inner flange (Figs |
P. magnicerca stat. nov. |
17´ | Cerci with two branches, without an inner flange | 18 |
18 | Spine on left side of supra-anal plate long and conspicuous (Fig. |
P. forcipata |
18´ | Spine on left side of supra-anal plate small (Fig. |
P. laeglae stat. nov. |
Distribution patterns of Central and West African Orthoptera taxa are poorly understood. As forest-bound taxa the comparatively species-rich genus Poreuomena very likely diversified in the course of several expansions and shrinkages of the Guineo-Congolian forest belt caused by large-scale climatic changes. Humid tropical Africa has been thought to have been covered more or less continuously by forest. About 33 m years ago (Ma) during the Oligocene a period of drastic global cooling fragmented the Eocene pan-African forest (
In the light of these dramatic changes of the climate in the past and thus leading to fragmentation and recurrent expansion of forest cover, the high diversity found in Poreuomena could be explained. However, Cestromoecha, on the other hand is a species-poor genus with only two known species at present. Either the latter genus did not cope well with recurrent fragmentation events of its habitat and species became extinct or the picture we see is due to poor collecting activities and gaps in our knowledge of the diversity of many Central and West African taxa including Cestromoecha. In Poreuomena however, several morphological closely related species suggest a diversification during the past couple of million years, similar to time-scales of speciation found in East African Orthoptera (e.g.
In Poreuomena two morphological lineages are discernible: Lineage 1 consists of species with a (for Poreuomena) typical bilobed 10th abdominal tergite in males or a 10th tergite deviated from this condition (Fig.
The highest diversity of Poreuomena is found in forests between Gabon and Cameroon. The forests of this area persisted even during extremely arid times, for example during the Ice Ages as seen in various forest trees (
Ultimately, molecular analyses should show the relationships of the species and lineages of Poreuomena, very likely supporting that forest remains of the once strongly-fragmented Guineo-Congolian area served as refugia for flora and fauna and a motor for diversification, analogous to the Eastern Arc Mountains in East Africa.
We wish to thank Philippe Moretto and the late Philippe Annoyer, President of the Association Insectes du Monde (www.insectesdumonde.org) and organiser of the expedition Sangha 2012, who kindly let BM study the material collected during the 2005–2012 Sangha entomological missions (Central African Republic), Samuel Danflous, Matias Loubes for their collaboration and help during the collecting nights in the Taï National Park (Côte d’Ivoire) in March 2017, both on the ground and on the canopy. We are indebted with Richard Smith, Chairman of the African Natural History Research Trust (ANHRT) (Hereford, UK), who loaned to BM for study the specimens collected in Gabon and Cameroon, Hitoshi Takano, researcher and curator of ANHRT and the collectors and collaborators in the Cameroon and Gabon entomological expeditions carried out by ANHRT and the Association Catharsius (in Gabon Philippe Moretto, Marios Aristophanous, Violette Dérozier et Jean-Louis Albert). ANHRT and we thank very much the Ministre des Eaux et Forêts of Gabon, Prof. Lee White; Monsieur l’Administrateur Général of Centre National de la Recherche Scientifique for the research authorisations, the Rector of the Université des Sciences et Techniques of Masuku, Prof. Ella Missang Crépin, the Director of the Département de Biologie, Prof. Nicaise Lepengue and Dr Stefan Ntie. Finally we thank Nicolas Moulin for providing one specimen from Lope National Park, in Gabon.
We also would thank very much Haralabos Tsolakis for helping with the interpretation of the Greek origin of the scientific names of Poreuomena and Cestromoecha, and Rob Felix for his kind help in the map production.
This research received support from the Synthesys Project, which is financed by European Community Research Infrastructure Action under the FP7 “Capacities” Programme. For Bruno Massa at the Museo Nacional de Ciencias Naturales, Madrid (CSIC) (2013: ES-TAF-2438), the Museum für Naturkunde, Berlin (2014: DE-TAF-4109), the Naturhistorisches Museum, Vienna (2016: AT-TAF-5324), the National Museum, Prague (2016: CZ-TAF-5559) and the Royal Belgian Institute of Natural Sciences, Bruxelles (2017: BE-TAF-6319). For Claudia Hemp for the museums of Brussels and Tervuren, Belgium (2019: BE-TAF 2685) and Budapest, Hungary (2019: HU-TAF 2666). We are especially indebted to Mercedes Paris (Museo Nacional de Ciencias Naturales of Madrid), Michael Ohl (Museum für Naturkunde of Berlin), Susanne Randolf and Harald Bruckner (Naturhistorisches Museum, Vienna), Jérôme Constant (Royal Belgian Institute of Natural Sciences, Bruxelles), Martin Fikáček (National Museum Natural History, Prague), Laure Desutter (Muséum National d’Histoire Naturelle, Paris), Marc de Meyer and Stepháne Hanot of the African Museum in Tervuren, Belgium, facilitating the study of specimens preserved in their museums.
Collecting authorisations and research permits were obtained as follows: 019/UB/DSV2012 of 16.I.2012 from Bangui University, Central African Republic; 021/MESRS/DGRI of 15.II.2017 from the Ministère de l’Einsegnement Superieur et de la Recherche Scientifique of Côte d’Ivoire; AR0029/19/MESRSTT/CENAREST/CG/CST/CSAR of 11.VII.2019, issued from the Commission Scientifique d’Examen des Demandes d’Autorisations de Recherche of the Ministère de l’Enseignement Supérieur, de la Recherche Scientifique et du Transfert des technologies du Gabon.
Thanks again to Jérôme Constant of
Many thanks to the two reviewers Dr. K.-G. Heller and Dr. S. Ingrisch for very valuable comments on the manuscript.