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Revision of the Madagascan genera Oncodopus Brongniart and Colossopus Saussure (Orthoptera: Tettigoniidae: Conocephalinae; Euconchophorini), with description of Malagasopus gen. nov

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The endemic Madagascan genera Oncodopus Brongniart and Colossopus Saussure are revised using museum specimens, including the types, and recently collected material. A new genus, Malagasopus gen. nov., and seven new species, Malagasopus desutterae sp. nov., Malagasopus meridianus sp. nov., Oncodopus janetae sp. nov., Oncodopus brongniarti sp. nov., Oncodopus saussurei sp. nov., Oncodopus soalalaensis sp. nov. and Colossopus parvicavus sp. nov. are described. Lectotypes are designated for Oncodopus zonatus Brongniart, 1897 and Colossopus redtenbacheri (Brongniart, 1897). A new term, mesothoracic auricle, is proposed for a structure on the episternum of the mesothorax. A tabulated key to the genera and keys to the species are presented. All species are described and diagnosed, and their phylogenetic relationships, geographical distributions, habitat preferences and phenologies are documented. The life history of Colossopus grandidieri is described, and the unusual possible mate-guarding behaviour of several species is discussed. Maps showing the distribution of the species are presented, as too are 57 photographs of museum specimens, 51 drawings of morphological characters, 17 photographs of living specimens and one habitat photograph.
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Accepted by D. Rentz: 5 Sept. 2017; published: 31 Oct. 2017
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2017 Magnolia Press
Zootaxa 4341 (2): 193
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https://doi.org/10.11646/zootaxa.4341.2.2
http://zoobank.org/urn:lsid:zoobank.org:pub:05152B19-56AA-4CCD-A3C6-53EA3369A54C
Revision of the Madagascan genera Oncodopus Brongniart and Colossopus
Saussure (Orthoptera: Tettigoniidae: Conocephalinae; Euconchophorini),
with description of Malagasopus gen. nov.
MUSTAFA ÜNAL
1,3
& GEORGE W. BECCALONI
2
1
Abant İzzet Baysal Üniversitesi, Fen-Edebiyat Fakültesi, Biyoloji Bölüm, 14030 Bolu, Turkey. E-mail: unal@ibu.edu.tr
2
Department of Life Sciences, The Natural History Museum, Cromwell Road, South Kensington, London SW7 5BD, UK.
E-mail: g.beccaloni@gmail.com
3
Corresponding author
Abstract
The endemic Madagascan genera Oncodopus Brongniart and Colossopus Saussure are revised using museum specimens,
including the types, and recently collected material. A new genus, Malagasopus gen. nov., and seven new species, Mal-
agasopus desutterae sp. nov., Malagasopus meridianus sp. nov., Oncodopus janetae sp. nov., Oncodopus brongniarti sp.
nov., Oncodopus saussurei sp. nov., Oncodopus soalalaensis sp. nov. and Colossopus parvicavus sp. nov. are described.
Lectotypes are designated for Oncodopus zonatus Brongniart, 1897 and Colossopus redtenbacheri (Brongniart, 1897). A
new term, mesothoracic auricle, is proposed for a structure on the episternum of the mesothorax. A tabulated key to the
genera and keys to the species are presented. All species are described and diagnosed, and their phylogenetic relationships,
geographical distributions, habitat preferences and phenologies are documented. The life history of Colossopus grandi-
dieri is described, and the unusual possible mate-guarding behaviour of several species is discussed. Maps showing the
distribution of the species are presented, as too are 57 photographs of museum specimens, 51 drawings of morphological
characters, 17 photographs of living specimens and one habitat photograph.
Key words: Madagascar, Orthoptera, Conocephalinae, Oncodopus, Colossopus, Malagasopus, revision, new genus, new
species, lectotype, taxonomy, distribution, mate-guarding
Introduction
The genera Oncodopus and Colossopus with three included species were described at the end of the 19
th
century.
Oncodopus was erected by Brongniart (1897) to contain two new species O. zonatus and O. redtenbacheri.
Brongniart described O. zonatus from two syntypes, both of which are male nymphs. He described O.
redtenbacheri from two syntypes, one adult male and one adult female. Only O. zonatus was illustrated in this
paper. Saussure (1899: 628) erected Colossopus to contain a new species C. grandidieri, and included Oncodopus
redtenbacheri in this genus as C. (Oncodopus) redtenbacheri Brong. Since he recognized Oncodopus as a valid
genus on page 626 of his paper, the reason for stating the generic name Oncodopus in parentheses after Colossopus
was probably simply to indicate the previous generic placement of the species, not to propose Oncodopus as a
subgenus. Saussure (1899) also described the female and adult male of O. zonatus for the first time. Possibly
because of Saussure’s misleading combination Kirby (1906: 270) gave C. redtenbacheri under the genus
Oncodopus as O. Redtenbacheri, Brongn., but listed the others in their correct combinations.
Karny (1912) followed Saussure’s classification. He listed Oncodopus with a single species O. zonatus, and
Colossopus with two species C. grandidieri and C. redtenbacheri. Karny also included the generic characters of
both genera and seven figures of the three species. Chopard (1948), discussing the known Madagascan
Tettigoniidae, listed eleven endemic genera of Agraeciinae including Oncodopus and Colossopus. He included a
dorsal illustration of a female of O. zonatus. Gorochov (1988) erected the tribe Euconchophorini to contain the
genera Euconchophora Brongniart 1897, Odontolakis Redtenbacher 1891, Oncodopus Brongniart 1897,
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Colossopus Saussure 1899 and Amblylakis Redtenbacher 1891. He also gave one figure of Oncodopus. Zompro
(2008) mentioned a likely new species of Colossopus as Collossopus sp.. Finally, Ünal & Beccaloni (2008)
discussed the type specimens of Colossopus grandidieri and selected the male as lectotype.
The descriptions and figures given in the historical publications mentioned above are enough to identify both
described genera and their known species. However recently collected specimens and some unidentified material
preserved in museum collections necessitates a review of this group. From the rich material examined, a new
genus, Malagasopus, and 7 new species, Malagasopus desutterae sp. nov., Malagasopus meridianus sp. nov.,
Oncodopus janetae sp. nov., Oncodopus brongniarti sp. nov., Oncodopus saussurei sp. nov., Oncodopus
soalalaensis sp. nov. and Colossopus parvicavus sp. nov. are described. Lectotypes for O. zonatus and C.
redtenbacheri are designated. Generic characters for Oncodopus, Colossopus and Malagasopus are compared and
keys for identifying the species are provided. Phylogenetic relationships of the taxa are discussed based on a study
of their morphological characters.
We propose the name mesothoracic auricle for a structure on the episternum of the mesothorax which is
situated directly opposite the prothoracic auditory spiracle. Previous authors describing this structure, likened it to
a mussel or other seashell, cave, or scoop (Redtenbacher, 1891; Brongniart, 1897; Saussure, 1899; Karny, 1912).
Redtenbacher (1891) described its inner part as “foramen conchae mesosternalis and its outer lobe as “lobus
auricularis mesosterni. Ingrisch (1998) referred to it as a “conchate projection of mesopleura. According to
Ingrisch (1998) it is not known if the auditory spiracle has any special function. We suggest that it may gather
sound and reflect it into the prothoracic auditory spiracle. Although the meso- and metathorax are almost fused, the
prothorax can be moved from side to side and the insect can move the mesothoracic auricle towards or away from
the prothoracic auditory spiracle by turning to the right or left. Doing this may help the insect to determine the
direction of the sound.
Material and methods
The 146 specimens we studied were either already part of museum collections or were newly collected by one of
the authors (G.W. Beccaloni) or by a team from the California Academy of Sciences. Drawings were made using a
stereo microscope with an ocular tube for drawing. Photographs were taken using a SLR digital camera.
Measurements were made using a ruler and a micrometer attached to a stereo microscope. The taxa are arranged
taxonomically based on the relationships we inferred from the morphological data gathered during this work. The
locality data of specimens listed in the "Material examined" sections is as stated on the original data labels.
Locality names listed in the "Distribution" sections are given according to modern usage. The drawings and the
photographs of museum specimens were produced by M. Ünal.
The following abbreviations are used for the collections cited in the text:
CAS California Academy of Sciences, San Francisco
MHNG Museum d’Histoire Naturelle, Geneva
MNCN Museo Nacional de Ciencias Naturales, Madrid
MNHN Muséum National d‘Historie Naturelle, Paris
NHMUK Natural History Museum, London
NMW Naturhistorisches Museum Wien
SMFD Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt
ZMB Zoologischen Museum Berlin
Results
Family: Tettigoniidae Krauss, 1902
Subfamily: Conocephalinae Burmeister, 1838
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REVISION OF ONCODOPUS AND COLOSSOPUS
Tribe: Euconchophorini Gorochov, 1988
Based on the characters listed in Table 1 we propose that the three genera are related as follows: (Oncodopus +
Malagasopus) + Colossopus.
TABLE 1. Comparison of the genera Oncodopus, Colossopus and Malagasopus gen. nov.
Oncodopus Brongniart, 1897
Brongniart 1897: 125, 205. Type species: Oncodopus zonatus Brongniart, 1897.
Saussure 1899: 626; Kirby 1906: 270; Karny 1912: 8, 38; Chopard 1948: 125; Gorochov 1988: 176, 180; Otte 1997: 32.
Diagnosis. Body relatively depressed dorso-ventrally (Fig. 12). Fastigium of vertex elongated, simple, spiniform
(Figs. 1, 10). Face almost flattened, with distinct tubercles, four of which are larger and spiniform. Pronotum
almost covers tegmina in male (Figs. 10, 11). Mesothorax with a large seashell-shaped mesothoracic auricle.
Tympanal organ on fore tibia slit shaped. Forelegs strongly developed, large, with very strong and large spines
(Figs. 10, 16). Fore femur strongly widened, with a large inner spine on anterio-lateral margin; its inner and outer
sides without apical spine ventrally (Figs. 4, 10). Hind tibia without apical spur dorsally (Fig. 7). Male last tergite
very short, without or with a shallow small median notch on posterior margin (Figs. 13, 43). Ovipositor shorter
than abdomen (Fig. 18).
This genus is closely related to Malagasopus gen. nov. and Colossopus Saussure. See Table 1 for comparison
of these genera.
Key to species of Oncodopus
1 Male cercus (Fig. 34) with a small (sometimes indistinct) tooth on ventral margin of apex which is never dark in colour; its sin-
gle inner arm pointed with a distinct dark tooth. Posterior margin of male last tergite (Fig. 33) with a distinct median incision
compared to the other taxa in this genus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. brongniarti sp. nov.
- Male cercus (Figs. 14, 24, 44, 54) pointed, with two or three distinct teeth (bidentate or tridentate); its apex and inner arm(s)
pointed with distinct teeth with dark tips. Posterior margin of male last tergite (Figs. 13, 23, 43, 53) without or with a very
small median notch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2 Male cercus (Figs. 14, 24) bidentate, with one inner arm. Posterior margin of male last tergite (Figs. 13, 23) with a small
Oncodopus Malagasopus gen. nov. Colossopus
Fastigium of
vertex
simple, large and spiniform (Fig. 1) with 3 short spiniform arms
(tridentate) (Fig. 2)
with 3 long spiniform arms
(tridentate) (Fig. 3)
Male tegmina almost fully covered by pronotum
(Fig. 10)
protrude from under pronotum (Fig.
60)
reaching to far beyond end of
pronotum (Fig. 76)
Fore femur without apical spine ventrally on
inner and outer sides (Fig. 4)
without apical spine ventrally on
inner and outer sides (Fig. 5)
with a large apical spine
ventrally on inner and outer sides
(Fig. 6)
Middle and
hind femora
with ventral spines on outer side
(except for a number of specimens
of Oncodopus saussurei)
without ventral spines with ventral spines on outer side
(except hind femur of
Colossopus parvicavus)
Hind tibia without apical spur dorsally (Fig. 7) without apical spur dorsally (Fig. 8) with a single apical spur dorsally
on inner side (Fig. 9)
Prosternum with two, U-shaped spines (except
Oncodopus brongniarti)
with two, U-shaped spines with two, V-shaped spines
Male last
tergite
very short; posterior margin without
or with a shallow or an indistinct
median notch (Fig. 13)
long; posterior margin elongated
backward, with a distinct small and
narrow median incision (Fig. 63)
long; posterior margin elongated
backward, with a distinct, deep
median incision (Fig. 80)
Ovipositor short, shorter than abdomen, slightly
upcurved (Fig. 18)
slightly shorter than abdomen,
slightly upcurved (Fig. 68)
long, longer than abdomen,
straight (Fig. 85)
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median notch. Ovipositor slightly upcurved (Figs. 18, 28). Body size strongly variable, but not less than 35 mm in female (Tab.
2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Male cercus (Figs. 44, 54) tridentate, with 2 inner arms. Posterior margin of male last tergite (Figs. 43, 53) with median notch
almost absent. Ovipositor almost straight (O. soalalaensis) (Fig. 58), if not, body size less than 32.5 mm in female (Tab. 2) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 Body with distinct transverse black bands (Figs. 12, 18). Metazona of pronotum black, prozona light in colour in typical form
(Figs. 10, 11, 16, 17). Inner arm of male cercus long (Fig. 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. zonatus
- Body with indistinct transverse bands (Figs. 22, 28). Metazona of pronotum concolorous with prozona in typical form (Figs.
20, 21, 26, 27). Inner arm of male cercus short (Fig. 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. janetae sp. nov.
4 Body light, unicolorous without dark band and spot (Figs. 52, 58). Pronotum distinctly saddle-shaped (Figs. 51, 57). Fore tibia
with 5 inner spines ventrally (Figs. 50, 56). Male cercus as in Fig. 54. Ovipositor long (19.7 mm), slightly shorter than abdo-
men, straight, quite dark in colour (Fig. 58) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .O. soalalaensis sp. nov.
- Body with distinct transverse dark bands and dark spots (Figs. 42, 48). Pronotum not markedly saddle-shaped, straighter (Figs.
41, 47). Fore tibia with 4 inner spines ventrally (Figs. 40, 46). Male cercus as in Fig. 44. Ovipositor short (at most 14.9 mm),
clearly shorter than abdomen, slightly upcurved, very dark in colour (Fig. 48). . . . . . . . . . . . . . . . . . . . . . O. saussurei sp. nov.
We hypothesise that these five species are related in the following way: (brongniarti + (zonatus + janetae)) +
(saussurei + soalalaensis).
Oncodopus zonatus Brongniart, 1897
(Figs. 1, 4, 7, 10–19, 110, 112)
http://lsid.speciesfile.org/urn:lsid:Orthoptera.speciesfile.org:TaxonName:15096
Oncodopus zonatus: Brongniart 1897: 207; Saussure 1899: 626; Kirby 1906: 270; Karny 1912: 39; Chopard 1948: 125; Otte
1997: 32; Ünal & Beccaloni 2008: 35.
Type locality. Madagascar, Mozondeva (Morondava). Lectotype male, here designated, in alcohol (MNHN).
Redescription. Male: Body size quite variable (Tab. 2). Fastigium of vertex (Figs. 1, 10, 11) forming a simple
short horn, spiniform; almost conical, upper surface flattened with a wide furrow, 2.2 times longer than diameter of
eye, its basal part broader than antennal scape. Fastigium of frons (Fig. 11) with a short spiniform slightly
downcurved tooth. Pronotum (Figs. 10, 11) large, anterior margin narrow and convex, widened in middle, posterior
margin with a distinct median incision; 3.7 times longer than its height, and only 1.1 times longer than its width.
Prothoracic auditory spiracle large, partially concealed under pronotum. Opening of mesothoracic auricle large and
elliptical; ventral lobe distinct. Fore coxa with an elongate spine, projected forward dorsally. Fore femur (Fig. 10)
very muscular, short and broad; with a large inner spine on anterio-lateral margin; inner margin with 5–7 strong
spines ventrally. Fore tibia (Fig. 10) with very strong, curved 4 inner spines ventrally; its outer side with smaller 4–
5 spines. Prosternum with 2 long, U-shaped spines. Mid femur with 2–5 outer spines ventrally. Hind femur with 4–
8 ventral spines on outer side. Mid and hind tibia bear ventral spines. Hind tibia without apical spur dorsally (Fig.
7). Tegmina (Figs. 10–12) fully concealed under pronotum, their apex visible in some specimens, reaching to half
of metanotum. Abdomen (Fig. 12) large, mostly oval, anterior and posterior parts narrow, mid part broad. Last
tergite (Fig. 13) short and broad, posterior margin convex with a small median notch. Cercus (Fig. 14) with a short,
incurved apical tooth, inner arm very long with a distinct tooth at apex. Subgenital plate (Fig. 15) simple, with a
very short styli.
Female: As male. Hind margin of pronotum (Fig. 16) with a strong and broadly rounded incision. Tegmina
(Fig. 17) strongly reduced, small scale like laterally, not reaching halfway down of mesonotum. Cercus simple,
basal half cylindrical, apical half conical. Subgenital plate (Fig. 19) almost triangular with a distinct median carina,
its posterior margin with a small median notch. Ovipositor (Fig. 18) slender, slightly upcurved, almost 1.2–1.3
times longer than hind femur.
Colour (dry specimens). Body milky brown with distinct transverse dark bands in typical form. Head dark
except fastigium of vertex, eyes and posterior margin of occiput. Anterior margin of pronotum with a narrow dark
band; posterior part, (mainly of the metazona) dark , remaining part milky brown. Fore tibia light brown with
indistinct longitudinal dark stripes. Apical half of femora dark. Abdominal tergites with distinct dark band on
posterior margin. Male subgenital plate and cercus milky brown. Ovipositor reddish brown in apical part.
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REVISION OF ONCODOPUS AND COLOSSOPUS
FIGURES 1–9. Fastigium of vertex from above. 1 Oncodopus zonatus, 2 Malagasopus desutterae, 3 Colossopus grandidieri.
Figs. 4–6. apical half of left fore femur, inner view. 4 Oncodopus zonatus, 5 Malagasopus desutterae, 6 Colossopus
redtenbacheri. Figs. 7–9. apical part of left hind tibia from above. 7 Oncodopus zonatus, 8 Malagasopus desutterae, 9
Colossopus grandidieri. Scale bars 1 mm.
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FIGURES 10–19. Oncodopus zonatus. Male: 10 forelegs, head and pronotum, dorsal view, 11 head and pronotum, lateral view,
12 habitus, dorso-lateral view, 13 last abdominal tergite, 14 cercus, 15 subgenital plate. Female: 16 forelegs, head and
pronotum, dorsal view, 17 head and pronotum, lateral view, 18 habitus, dorso-lateral view, 19 subgenital plate. Scale bars 1
mm.
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REVISION OF ONCODOPUS AND COLOSSOPUS
Diagnosis. This species appears to be most closely related to O. janetae sp. nov. based on the structure of head,
last tergite of male, female subgenital plate, the variable body size. But this species differs from it by the structure
of pronotum, male cercus (with long inner arm) and colouration (body and pronotum with transverse dark bands) in
typical form. It is also near to O. brongniarti sp. nov. based on its colouration (with distinct transverse bands) and
male cercus with long inner arm. However, the male cercus is different to that of brongniarti (apical arm distinctly
pointed with a tooth), as is the male last tergite with a wide median incision on posterior margin.
Material examined. Madagascar, Mozondeva (Morondava), 1891, 2 male nymphs (Lecto- and paralectotype),
(leg. M. Grandidier et M. Doulliot), in alcohol; Madagascar, 1867, 2 male nymphs (leg. M. Grandidier);
Madagascar (S.O.), Forets du Fiher[en]ana, 1905, 1 male (leg. F. Geay); Prov. de Tulear, Bas Fiher[en]ana, 4 males,
8 females (leg. F. Geay); Madagask., 1 female (leg. M. Grandidier); Madagascar, Ikongo, 1902, 1 male (leg. M.
Grandidier); Madagascar Sud, Akiliazivo, 25.11.1963, 2 males, 1 female (leg. D. Wintrebert) (all in MNHN).
Madagascar, Morondava, February 2004, 1 male, in dead tree; Madagascar, Toliara Province, Zombitse-Vohibasia
National Park, (-22.60, 44.81) [22
°
36'0'' S, 44
°
48'36'' E], 17.11.2004, Coll. G.W. Beccaloni, 1 male, 1 female, in
forest near park headquarters, under bark of dead tree; Madagascar, Toliara Prov., c. 20 km North of Belo Sur
Tsiribihina (on main road north), 19
°
30'08.72" S, 44
°
30'25.17" E, 15.11.2007, Coll. G.W. Beccaloni, 3 males, 2
females, under bark of dead tree (all in NHMUK). SW Madagascar, Tulear, March 1904, 1 male (leg. S. Voltzkow)
(ZMB). Madagascar, Toliara Prov., Lake Ranobe, elev. 30 m, 23
°
00'629" S, 43
°
336' E, 2–10 May 2003, 1 male
nymph (leg. Frontier, Wilderness), grassland, transitional spiny-tamarind forest; Toliara Prov., Fiheranana
[Fiherenana], elev. 65 m, 23
°
13'351" S, 43
°
52'853" E, 5–7 August 2003, 1 male nymph (leg. Frontier &
Wilderness), leaf litter extraction degraded riparian forest close to water (all in CAS).
Measurements (mm). Lectotype (male) is a nymph. See Table 2 for the range of measurement of the adults.
Distribution. S.W. of Madagascar (Fig. 110). Toliara Province: Morondava, Toliara, Ikongo, Akiliazivo,
Zombitse-Vohibasia N.P., Belo Sur Tsiribihina, Fiherenana, Lake Ranobe.
Habitat. This species is known to occur in spiny forest and dry deciduous forest habitats as well as in riparian
forest. Individuals have been found resting under the bark of dead trees and one nymph was extracted from leaf litter.
Phenology. Adults of this species have been found between November and March, and nymphs in May and
August.
Remarks. The body size is highly variable in this species (see Table 2). Piotr Naskrecki (pers. commun., 2005)
believes that the largest males may be 'satellite males'. This variation in size amongst the adults of a species is also
seen in some of the other species of Oncodopus (see Table 2).
Oncodopus janetae sp. nov.
(Figs. 20–29, 110, 113)
http://lsid.speciesfile.org/urn:lsid:Orthoptera.speciesfile.org:TaxonName:499685
Type locality. Madagascar, Toliara Province, Andohahela National Park. Holotype male (NHMUK).
Description. Male (Holotype): Body size average for genus strongly variable in other males (Tab. 2).
Fastigium of vertex (Figs. 20, 21) simple, conical, elongated into a short horn; upper surface flattened with a wide
furrow; 2.6 times longer than diameter of eye; its basal part broader than antennal scape; its apex slightly
downcurved, straight in some males. Fastigium of frons (Fig. 21) with a large spiniform tooth. Face flattened,
granulated, with 4 very distinct spiniform teeth. Pronotum (20, 21) large, anterior margin convex, posterior margin
with a large incision; pronotum clearly concave in lateral view, three times longer than its height, very slightly
longer than its width, in Bekily population pronotum clearly longer than wide; prozona 1.5 times longer than
metazona. Prothoracic auditory spiracle large, triangular but with corners rounded, a large portion concealed under
lateral lobes of pronotum. Mesothoracic auricle and its opening large, ventral lobe distinct. Prosternum with 2 long,
U-shaped spines, reaching to level of ventral margin of fore coxa. Fore coxa with a long dorsal spine projected
forward. Fore femur (Fig. 20) with a large inner spine on anterio-lateral margin; with 5 inner spines ventrally, in
other males with 5–7 inner spines ventrally. Fore tibia (Fig. 20) with 4 very large inner and 4 smaller outer ventral
spines. Mid femur with 3 ventral spines on outer side (other males have 2–4 spines). Hind femur with 2 ventral
spines on outer margin of right leg, left leg with 4 spines (other males have 2–5 spines). Fore and mid tibia without
dorsal spines. Hind tibia with 5–6 dorsal spines on inner margin (other males have 3–6 spines); with 4 apical spurs
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ventrally, inner 2 slightly smaller than outer ones. Tegmina fully concealed under pronotum (Figs. 20, 21), reaching
to end of mesonotum (in some males they protrude a little beyond the hind margin of pronotum). Last tergite (Fig.
23) narrow and very wide, with a small median notch. Male cercus (Fig. 24) stout, short and thick; apical arm
rounded, with a short spine, slightly curved inwards; inner arm short, pointed with a spine. Subgenital plate (Fig.
25) with long styli, longer than depth of posterior incision of plate.
Female: Head as in male. Pronotum (Figs. 26, 27) broader than long and 3 times longer than its height; straight
in lateral view; its hind margin distinctly concave; metazona very short, almost 1/6 of total length of pronotum;
prozona 4.7 times longer than metazona in typical form. Legs as in male. Tegmina (Fig. 27) strongly reduced, scale
like laterally, reaching just beyond the middle of mesonotum, in some females fully concealed under pronotum.
Subgenital plate (Fig. 29) triangular, broader than long, hind margin with a small median notch. Last tergite very
short, with an indistinct median notch. Cercus simple, triangular, pointed at apex. Ovipositor (Fig. 28) slender,
gradually upcurved, in some females straighter; slightly longer than hind femur.
Colour (dry specimens). Body dark with dark reddish brown and black in typical form. Head reddish brown,
labrum, eye, scapus, lighter. Pronotum dark yellowish brown in male, blackish brown in female. Apical part of fore
femur blackish brown, basal and outer parts yellowish with dark spots in male, much darker in female with black.
All tibiae yellowish in male, brownish in female. Apical part of mid and hind femora dark with black, basal part
yellowish. Tarsi darker than tibiae with blackish brown. Abdomen pale reddish and yellowish brown with
transverse black stripes and spots, darker in female. Pronotal and abdominal sternites yellowish milky brown.
Subgenital plate and cercus yellowish brown. Ovipositor dark, with black, apical part reddish black.
Diagnosis. This new species is most closely related to O. zonatus judging by the structure of the head, the last
tergite of the male, the female subgenital plate, and possibly the variable body size. But the new species differs
from it in having a wider pronotum, a shorter inner arm of the male cercus, and its colouration (without distinct
transverse band, especially on the pronotum) in the typical form. In some populations the shape of pronotum is also
near to O. zonatus, but, it is easily recognizable by the male cercus and colouration. Without the male, it is difficult
to separate the females of O. zonatus and this new species except by the colour.
Material examined. Madagascar, Toliara Province, Andohahela National Park: Mangatsiaka parcel, (-24.83,
46.57) [24
°
49'48'' S, 46
°
34'12'' E], 30.11.2004, Coll. G.W. Beccaloni, 1 male (Holotype), 1 female; Madagascar,
Toliara Province, Andohahela National Park: Tsimelahy parcel, (-24.93, 46.65) [24
°
55'48'' S, 46
°
38'60'' E],
1.12.2004, Coll. G.W. Beccaloni, 1 male, 1 female, at base of [a living leaf of] Dypsis decaryi; Madagascar, Toliara
Province, littoral forest fragment, nr. Saint Luce, (-24.78, 47.18) [24
°
46'48'' S, 47
°
10'48'' E], 2.12.2004, Coll. G.W.
Beccaloni, 1 male; Fort Douphin [Dauphin], 1 male (leg. Scott Elliot) (all in NHMUK). Bekily, Red sud de L’ile,
January 1932, 3 males, 4 females, 4 male nymphs, January 1933, 1 female (leg. A. Seyrig); Col de Sakaralama,
Alluaud, 1901, 1 female; Sud de Morondava, Torev de Be Fasy, 1 female; Madagascar, Rég d’Ankazoabo, 1920, 4
male nymphs, 2 female nymphs (leg. C. Le Barbier); Andohahela [National Park?], 1800 m, 1 female, 1500 m, 1
female; Madagascar, Fénerive, 1902, 1 male (leg. F. Genot); Ambatoloma [Ambatalaona?], 1907, 1 female (leg.
Bernard); Madagascar, Forét d’Andampy, 60 km S.O. de Voheinar [Vohémar], Co’te N.E., 1889, 3 male nymphs
(leg. M. Grandidier) (all in MNHN). Madagascar, 1 male nymph (ZMB). Ampandrandave, Sud de Madagascar,
12.1931, 3 females (leg. A Seyrig) (MNCN). Madagascar, Toliara Prov., Réserve Spéciale de cap Sainte Marie,
14.9 km 261°W. Marovato, elev. 160 m., 13–19 Feb. 2002, 25
°
35'40" S, 45
°
8'49" E, 1 female, 12.3 km 262°W.
Marovato, elev. 200 m., 11–15 Feb. 2002, 25
°
34'54" S, 45
°
10'6" E, 1 female (leg. Fisher, Griswold et al.), in spiny
forest thicket; Toliara Prov., Parc National d’Andohahela, Forét de Manantalingo, 33.6 km 63 ENE Amboasary, 7.6
km. 99 E Hazofotsy, 12–16 January 2002, 24
°
49'1" S, 45
°
36'36" E, elev. 150 m., 1 male (leg. Fisher, Griswold et
al.), in spiny forest thicket; Toliara prov., Parc National de Zombitse, 19.8 km 8 E Sakaraha, elev. 770 m., 5–9 Feb.
2003, 22
°
50'36" S, 44
°
42'36", 1 female (leg. Fisher, Griswold et al.), in tropical dry forest (all in CAS).
Measurements (mm). Holotype (male): Body length: 44; pronotum length: 16.4; width of pronotum: 14.6;
fore femur length: 16.1; width of fore femur: 6.8; fore tibia length: 15.9; hind femur length: 18.7. See Table 2 for
the range of measurement of the other specimens including females.
Etymology. This species name is dedicated to Janet Beccaloni who provided considerable help to the second
author with collecting work in Madagascar.
Distribution. Southern and eastern Madagascar (Fig. 110). Toliara Province: Morondava, Toliara, Bekily, Fort
Dauphin, Ankazoabo, Andohahela N.P., Marovato, Zombitse-Vohibasia N.P. Toamasina Province (see remarks
below): Fénerive, Ambatoloma [Ambatalaona?], Forét d’Andampy.
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FIGURES 20–29. Oncodopus janetae sp. nov. Male: 20 forelegs, head and pronotum, dorsal view, 21 head and pronotum,
lateral view, 22 habitus, dorso-lateral view, 23 last abdominal tergite, 24 cercus; 25 subgenital plate. Female: 26 forelegs, head
and pronotum, dorsal view, 27 head and pronotum, lateral view, 28 habitus, dorso-lateral view, 29 subgenital plate. Scale bars 1
mm.
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Habitat. This species is known from littoral forest, spiny forest and tropical dry forest. Records for rainforest
are dubious (see remarks below). It has been found under dead bark and in the natural cavity found at the base of
living leaves of the palm Dypsis decaryi.
Phenology. Adult individuals have been found between November and February and nymphs in January.
Remarks. We regard the three old (1889, 1902, 1907) locality records in the Toamasina Province as dubious
since these sites are located in the humid climatic zone (Cornet 1974) which has rainforest vegetation, whereas all
other records are from dry forests in arid regions with a long dry season. Very extensive collecting of arthropods
since 1992 at more than 300 sites all over Madagascar (Fisher 2017), including many in the humid climatic zone,
by a team led by Brian Fisher of the California Academy of Sciences has failed to collect any specimens of O.
janetae, or indeed any of the other taxa discussed in our study, in rainforest habitats. Also note that old collecting
localities are often difficult to identify in Madagascar due to the large number of synonymous locality names and
because geographic coordinates are absent from the data labels.
The body size is highly variable in this species as it is in O. zonatus (see the remarks under O. zonatus and
Table 2).
Oncodopus brongniarti sp. nov.
(Figs. 30–39, 110, 114–116)
http://lsid.speciesfile.org/urn:lsid:Orthoptera.speciesfile.org:TaxonName:499686
Type locality. Madagascar, Toliara Province, S. Morondava, Belo Sur Mer. Holotype male (NHMUK).
Description. Male (Holotype): Body size average for genus (Tab. 2) (in one male body size quite large).
Fastigium of vertex (Figs. 30, 31) simple, conical, forming a short horn, upper surface flattened with a narrow
longitudinal median furrow, 2.3 times longer than diameter of eye, basal part 1.4 times wider than antennal scape.
Fastigium of frons (Fig. 31) with a spiniform slightly downcurved tooth. Face flattened, strongly granulated; each
tubercle like a small tooth, of which the lower two are larger. Pronotum (30, 31) long, 3.2 times longer than its
height and 1.2 times longer than its width; anterior margin convex; posterior margin with an angular, large median
incision; upper surface slightly concave in lateral view; prozona 1.9 times longer than metazona. Prothoracic
auditory spiracle large, oval; a small portion of its opening concealed under pronotum. Mesothoracic auricle small,
short; its opening almost triangular; ventral lobe short. Prosternum with two long, V-shaped spines, reaching to
level of ventral margin of fore coxa; widely separated towards to apex. Fore coxa with a short spine projecting
forward. Femora without dorsal spines. Fore femur (Fig. 30) with a large inner spine on anterio-lateral margin; with
6 inner spines ventrally. Fore tibia (Fig. 30) with 5 inner and 5 outer spines ventrally. Mid femur with 2 ventral
spines on outer margin. Hind femur with 2 outer spines ventrally (1–4 spines in other males). Fore and mid tibia
without dorsal spines. Hind tibia with 7 dorsal spines on inner margin (other males with 5–6 spines); with 4 apical
spurs ventrally, inner 2 slightly smaller than outer ones; without apical spurs dorsally. All tibia with ventral spines
on both margins. Tegmina (Figs. 30, 31) concealed under pronotum, its apex visible from the posterior incision of
pronotum in dorsal view, reaching to half of metanotum. Last tergite (Fig. 33) wider than long, posterior margin
with a shallow, wide, rounded median incision. Cercus (Fig. 34) stout, apical part with a very small tooth (which is
not spiniform) on its ventral margin; inner arm very long, its apex slightly bent backward, pointed with a distinct
tooth. Subgenital plate (Fig. 35) almost as long as wide, its styli distinctly longer than depth of median incision.
Female: Face as in male. Fastigium of vertex (Fig. 36) short, 1.8 times longer than diameter of eye, its basal
part 1.5 times wider than antennal scape. Pronotum (Figs. 36, 37) as wide as long; posterior margin with a large
rounded incision; pronotum 2.9 times longer than its height; prozona 3 times longer than metazona. Tegmina (Fig.
37) strongly reduced, scale like laterally, reaching to half of mesonotum. Prosternal spines V-shaped, slightly
shorter than that of male. Fore femur (Fig. 36) with 6–7 ventral spines on inner margin and a large inner spine on
anterio-lateral margin. Tibial spines as in male. Mid femur with 2 outer spines ventrally, in other females with 2–4
spines. Hind femur with 2 ventral spines on outer side. Last tergite very short and wide, its posterior incision
distinct. Cercus stout, basal part broad, apical third strongly and sharply narrowed, its apex slightly incurved and
pointed. Subgenital plate (Fig. 39) almost triangular with a median carina, its apex with a small rounded incision.
Ovipositor (Fig. 38) clearly longer than hind femur, slightly upcurved.
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FIGURES 30–39. Oncodopus brongniarti sp. nov. Male: 30 forelegs, head and pronotum, dorsal view, 31 head and pronotum,
lateral view, 32 habitus, dorso-lateral view, 33 last abdominal tergite, 34 cercus, 35 subgenital plate. Female: 36 forelegs, head
and pronotum, dorsal view, 37 head and pronotum, lateral view, 38 habitus, dorso-lateral view, 39 subgenital plate. Scale bars 1
mm.
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Colour (dry specimens). Body milky brown with distinct transverse dark reddish-brown bands. Head reddish
brown. Eye, fastigium of vertex, labrum, maxillary and mandibulary palps slightly lighter. Anterior part of
pronotum milky brown, posterior part (metazona) very dark with reddish-brown. In female metazona dark brown.
All tibiae milky brown. Apical part of each femur dark, with a large black or reddish-black band. Posterior half of
first abdominal tergite in male and the whole of it in the female dark, black or reddish brown; posterior part of each
abdominal tergite with transverse dark band, black or reddish brown. All sternites, cerci and subgenital plate
yellowish milky brown. Ovipositor milky brown in basal 1/3, reddish brown in remaining part.
Diagnosis. Based on the colour pattern (with distinct transverse dark bands), the male cercus with a long inner
arm, and the shape of the ovipositor this new species appears to be most closely related to O. zonatus. It differs
from it by the absence of a distinct apical spine on the male cercus, the larger median incision of the last tergite in
both sexes, the more pronounced tubercles of the face and the female cercus. It is recognized by the shape of the
male cercus and the last tergite.
Material examined. Madagascar, Toliara Province, Belo Sur Mer (South of Morondava), 20
°
44'53.63" S,
43
°
59'36.97" E, 22.11.2007, Coll. G.W. Beccaloni, 2 males (including Holotype), 2 females, under bark of dead
tree; Madagascar, S. Centr. Madagascar, 1 male [no more data]; Toliara province, Forest near village of Mangily, c.
27 km N. of Toliara, (-23.15, 43.60) [23
°
8'60'' S, 43
°
36'0'' E], 11.11.2004, Coll. G.W. Beccaloni, 1 female, under
bark of dead tree; Madagascar, Foret de Zombitsy [Zombitse-Vohibasia N.P.], near Sakaraha, 650 m, 16.12.1959, 1
female (leg. E. S. Ross) (all in NHMUK).
Measurements (mm). Holotype (male): Body length: 40.3; pronotum length: 14.2; width of pronotum: 12.1;
fore femur length: 14.5; width of fore femur: 5.7; fore tibia length: 13.3; hind femur length: 15. See Table 2 for the
range of measurement of the other specimens including females.
Etymology. This species name is dedicated to Charles Brongniart who was the first to discover this group of
insects in 1897. He is the author of the genus Oncodopus and the species Oncodopus zonatus and Colossopus
redtenbacheri.
Distribution. South-west and south-central Madagascar (Fig. 110). Toliara Province: Toliara, Mangily, Foret
de Zombitsy [Zombitse-Vohibasia N.P.], Morondava.
Habitat. This species is known from spiny forest, dry costal scrub and dry deciduous forest. It has been found
under the bark of dead trees.
Phenology. Adults have been found in November and December.
Remarks. The body size is variable as in above 2 species (see under the Remarks section for O. zonatus and
Table 2).
Oncodopus saussurei sp. nov.
(Figs. 40–49, 110, 117)
http://lsid.speciesfile.org/urn:lsid:Orthoptera.speciesfile.org:TaxonName:499687
Type locality. Madagascar, Fianarantsoa Prov., South of Ambohimahasoa. Holotype male (NHMUK).
Description. Male (Holotype): Body size smallest in the genus (Tab. 2). Fastigium of vertex (Figs. 40, 41)
simple, almost conical, forming a short horn, upper surface flattened without furrow, twice as long as eye diameter,
its basal part 1.5 times wider than antennal scape, its apex slightly downcurved. Fastigium of frons (Fig. 41) with a
spiniform tooth. Face flattened, with few indistinct tubercles, with 2 distinct teeth. Pronotum (Figs. 40, 41) slightly
longer than wide, 3.8 times longer than its height and 1.1 times longer than its width; anterior margin convex,
posterior margin slightly concave; metazona long, only 1.4 times shorter than prozona in middle; pronotum
strongly concave in lateral view. Prothoracic auditory spiracle small almost triangular, only a small part visible in
lateral view. Mesothoracic auricle small, its inner gap oval, lacking ventral lobe. Prosternum with 2 short, U-shaped
spines, almost parallel, not reaching to level of ventral margin of fore coxa. Fore coxa with a short spine projecting
forward. Fore femur (Fig. 40) with a large inner spine on anterio-lateral margin; with 5 ventral spines on inner
margin (other males have 3–5 spines). Fore tibia (Fig. 40) with 4 inner and 4 outer spines ventrally. Mid femur with
1 outer spine ventrally (some males lack this spine). Hind femur without spines. Hind tibia with 2–4 dorsal spines
on inner margin (other males have 3–5 spines); 4 apical spurs ventrally, inner two slightly smaller than outer ones.
Tegmina (Figs. 40, 41) fully concealed under pronotum, reaching to mid point of metanotum. Last abdominal
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REVISION OF ONCODOPUS AND COLOSSOPUS
tergite (Fig. 43) very short and wide, its posterior margin slightly concave, without notch or incision. Cercus (Fig.
44) short, apical part rounded, with an incurved apical tooth; with 2 inner arms, upper one very short, pointed with
a tooth; lower one longer and pointed with a tooth. Subgenital plate (Fig. 45) slightly longer than wide, its styli
broad and longer than depth of posterior incision.
Female: Head as in male. Pronotum (Figs. 46, 47) 1.2 times wider than long and 2.7 times longer than its
width; posterior margin strongly concave; prozona 3.5 times longer than metazona. Ventral half of prothoracic
auditory spiracle visible in lateral view. Fore femur (Fig. 46) with 3–6 ventral spines on inner margin. Mid and hind
femora without or with 1–2 ventral spines on outer margin. Hind tibia with 3–5 dorsal spines on inner margin.
Spines of prosternum reaching to level of ventral margin of fore coxa, in some specimens a little shorter. Tegmina
(Fig. 47) strongly reduced, scale like laterally, not reaching to half of mesonotum, largely concealed under
pronotum. Last tergite short and wide, its posterior margin convex, without median incision. Subgenital plate (Fig.
49) triangular, with a median carina, posterior margin with a small incision. Cercus triangular, slightly incurved,
pointed at apex with an indistinct spine. Ovipositor (Fig. 48) longer than hind femur, slightly upcurved.
Colour (dry specimens). Body various shades of greenish brown, black and yellowish brown. Face brown in
male, blackish brown in female. Dorsal surface of head greenish brown with irregular black pattern. Pronotum
greenish brown, with short longitudinal black stripes. Fore femur and tibia milky yellowish brown; dorsal surface
of fore femur with distinct black spots, ventral surface completely black including spines; fore tibial spines black.
Mid femur as fore femur. Hind femur dark apically, basal part with dark spots. Anterior part of each abdominal
tergite except last 2–3 ones with black transverse band on both sides, not touching in middle, behind it with a faint
band; hind margin of abdominal tergites completely black. All sternites, male cercus and subgenital plate
unicolorous milky brown. Female subgenital plate with a black macula in middle. Ovipositor milky brown at base,
remaining part dark with black or brownish black.
Diagnosis. This species is the smallest member of the genus. It is unique in the shape of the male pronotum
and cercus, the last tergite in both sexes, the ovipositor and colouration. It appears to be most closely related to O.
soalalaensis sp. nov. judging by the shape of posterior margin of pronotum and the last tergite of the male, the
tridentate male cercus and its small body size. It differs from it by the general structure of pronotum in both sexes,
the male cercus, the fore tibia with 4 ventral spines on both sides (O. soalalaensis has 5 spines), the shape of its
shorter ovipositor, its colouration and its distribution.
Material examined. Madagascar, Fianarantsoa Prov., beside R.N. 7 south of Ambohimahasoa, (c. -21.17,
47.25) [21
°
17' S, 47
°
25' E], 13.11.2004, Coll. G.W. Beccaloni, 1 male (Holotype), 1 female, under a stone;
Antananarivo, (-18.92, 47.52) [18
°
55'12'' S, 47
°
31'12'' E], on R.N. 7 to the south of the city, 12.11.2004, Coll. G.W.
Beccaloni, 2 females, under a stone, (all in NHMUK). Madagascar, Pic de Ivohibe, 2200 m, 8.11.1950, 3 males, 1
female, 1 female nymph (leg. A.R.); Madagascar-centre, Plateau Soaindrana, 2090 m, Andringitra-Ambalavao,
15.1.1958, 1 female (leg. R. Paulian); Andringitra centre, plat. Andohariana, 2000–2100 m, 9.11–10.12.1970, 1
female (all in MNHN).
Measurements (mm). Holotype (male): Body length: 30.9; pronotum length: 9.5; width of pronotum: 8.2; fore
femur length: 7.9; width of fore femur: 3.6; fore tibia length: 7.5; hind femur length: 9.9. See Table 2 for the range
of measurement of the other specimens including females.
Etymology. This species name is dedicated to Henri Louis Frédéric de Saussure, who was the second person to
study these insects (in 1899). Saussure is the author of the genus Colossopus and the species C. grandidieri.
Distribution. Central and south-central Madagascar (Fig. 110). Antananarivo Province: Antananarivo,
Soavinandriana. Fianarantsoa Province: Ambohimahasoa, Andringitra, Ambalavao, Ivohibe.
Habitat. This species occurs in seasonally dry habitats on the Central Plateau of Madagascar which are largely
grassland today but which were probably sclerophyllous woodland in the past. It is known to occur up to 2200
metres—the highest altitude of any species in this group. It has been found under stones.
Phenology. Adults have been found between November and January and one nymph (instar unknown) in
November.
Oncodopus soalalaensis sp. nov.
(Figs. 50–59, 110)
http://lsid.speciesfile.org/urn:lsid:Orthoptera.speciesfile.org:TaxonName:499688
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Type locality. Madagascar, Mahajanga, Soalala Reserve. Holotype male (MNHN).
Description. Male (Holotype): Body size small for genus (Tab. 2). Fastigium of vertex (Figs. 50, 51) simple,
elongated into a short horn, upper surface flattened, 1.7 times longer than eye diameter, its basal part 1.5 times
wider than antennal scape. Fastigium of frons (Fig. 51) with a small spiniform tooth. Face convex, not distinctly
flattened, with tubercles. Pronotum (Figs. 50, 51) almost saddle shaped, 1.1 times longer than wide and 3.1 times
longer than its height; depression of typical sulcus deep and distinct; anterior margin slightly convex, posterior
margin concave in middle; prozona 1.5 times longer than metazona. Tegmina (Fig. 50, 51) almost concealed under
pronotum, only the apex visible from above; reaching to a little beyond the middle of the metanotum. Prothoracic
auditory spiracle large, oval; its dorsal half concealed under lateral lobe of pronotum. Mesothoracic auricle small,
without ventral lobe; its dorsal part concealed under lateral lobe of pronotum. Prosternum with two, U-shaped
spines, almost parallel, not reaching to level of ventral margin of fore coxa. Fore coxa with a spine projecting
forward. Fore femur (Fig. 50) with a large inner spine on anterio-lateral margin and 6 ventral spines on inner
margin. Fore tibia (Fig. 50) with 5 inner and 5 outer spines ventrally. Mid and hind femora with 2 ventral spines on
outer margin apically. Hind tibia with 6 dorsal spines on inner side (5–6 spines in other males); with 4 apical spurs
ventrally, all of which almost equal in length. Last abdominal tergite (Fig. 53) short and broad, posterior margin
convex without median notch (other males have a very indistinct median notch). Cercus (Fig. 54) stout, with 3
arms; apical arm short, roundly curved inwards, pointed with a spine; basal inner arm short and broad, pointed with
a tooth; median inner arm larger, near to ventral surface of cercus, pointed at apex with a large tooth. Subgenital
plate (Fig. 55) longer than wide, its styli longer than depth of posterior median incision.
Female: Head as in male. Fastigium of vertex (Figs. 56, 57) 1.3 times wider than antennal scape. Pronotum
(Fig. 56, 57) short and wide, saddle shaped, 1.2 times wider than long; anterior margin convex, posterior margin
broadly concave; prozona 3 times longer than metazona. Tegmina (Fig. 56) reduced, like a very small scale placed
on anterio-lateral margin of mesonotum. Only a small portion of prothoracic auditory spiracle covered by lateral
lobe of pronotum. Mesothoracic auricle without ventral lobe, not covered by pronotum. Legs and their spines as in
male. Last abdominal tergite very short and broad, posterior margin without median notch. Cercus short conical, its
apex slightly incurved. Subgenital plate (Fig. 59) triangular, with distinct median carina; its apex with a small
notch, hind margin curved upwards in posterior view. Ovipositor (Fig. 58) distinctly longer than hind femur,
slightly shorter than abdomen, almost straight, apical part very slightly upcurved.
Colour (dry specimens). Body various shades of light brown. Upper portion of face yellowish brown, lower
part darker with reddish brown; labrum black; upper surface of head greenish brown; eye darker. Pronotum slightly
reddish brown with some short longitudinal stripes; posterior half of metazona greenish brown in male. Legs
yellowish brown; mid and hind tibia of male reddish brown. Abdomen slightly darker reddish brown, with
indistinct transverse dark bands in male, greenish brown in female. All sternites reddish brown in male and
concolorous in female. Basal part of ovipositor greenish brown, turning to reddish brown in apical part.
Diagnosis. This species has the following unique characters: the shape of its pronotum in both sexes, the
morphology of the male cercus, the shape of the ovipositor and its colour. Features indicating it is closely related to
O. saussurei sp. nov. are the tridentate male cercus, the small body size, and the last tergite without median
incision. However, this new species differs from O. saussurei by the fore tibia with 5 inner and 5 outer spines
ventrally (O. saussurei has 4 spine), the shape of the pronotum, features of the male cercus, the shape of its longer
ovipositor and its colour. The fore tibia has a similar structure to that of O. brongniarti sp. nov., having 5 ventral
spines on both sides. But, all other characters are very different from it.
Material examined. Madagascar, Mahajanga, Soalala Reserve nat. VIII, 3 males (including Holotype), 1
female nymph; Namoroka, 9.1952, 1 male nymph; Madagascar centre, Rég. de Bétafo d’Ambositra, et de
Miandrivazo, 1905, 1 female (leg. Bauet) (all in MNHN).
Measurements (mm). Holotype (male): Body length: 33.6; pronotum length: 10.6; width of pronotum: 9.4;
fore femur length: 8.8; width of fore femur: 3.7; fore tibia length: 9.4; hind femur length: 13. See Table 2 for the
range of measurement of the other specimens including female.
Etymology. The species name is derived from “Soalala” is the type locality of this new species.
Distribution. Central and western Madagascar (Fig. 110). Mahajanga Province: Soalala, Namoroka.
Fianarantsoa-Toliara Province: Betafo, between Ambositra and Miandrivazo.
Habitat. The region in which this species occurs contains dry deciduous forest. Living specimens have not
been studied.
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REVISION OF ONCODOPUS AND COLOSSOPUS
FIGURES 40–49. Oncodopus saussurei sp. nov. Male: 40 forelegs, head and pronotum, dorsal view, 41 head and pronotum,
lateral view, 42 habitus, dorso-lateral view, 43 last abdominal tergite, 44 cercus, 45 subgenital plate. Female: 46 forelegs, head
and pronotum, dorsal view, 47 head and pronotum, lateral view, 48 habitus, dorso-lateral view, 49 subgenital plate. Scale bars 1
mm.
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FIGURES 50–59. Oncodopus soalalaensis sp. nov. Male: 50 forelegs, head and pronotum, dorsal view, 51 head and
pronotum, lateral view, 52 habitus, dorso-lateral view, 53 last abdominal tergite, 54 cercus, 55 subgenital plate. Female: 56
forelegs, head and pronotum, dorsal view, 57 head and pronotum, lateral view, 58 habitus, dorso-lateral view, 59 subgenital
plate. Scale bars 1 mm.
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REVISION OF ONCODOPUS AND COLOSSOPUS
Phenology. A nymph of this species was found in September. The exact collecting dates of the adults were not
recorded on the labels.
TABLE 2. Measurements (mm) of the body parts of Oncodopus spp. M male, F female
Malagasopus gen. nov.
Type species. Malagasopus desutterae sp. nov., here designated.
Description and Diagnosis. Body slightly depressed dorso-ventrally (Fig. 62). Head wide. Fastigium of vertex
(Figs. 2, 60) relatively elongated, with 3 short spiniform arms all of which are slightly directed upwards. Face
almost flattened, with distinct tubercles, four of which larger, spiniform. In M. desutterae a large portion of the
male tegmina is covered by the pronotum and only the apical parts are visible from above (Fig. 60), while in the
other species, M. meridianus, a large apical portion of the male tegmina is visible from above (Fig. 70).
Mesothorax with a large seashell-shaped mesothoracic auricle. Tympanal organ on fore tibia slit shaped. Forelegs
(Figs. 60, 66) strongly developed, large, with very strong and large spines. Fore femur (Figs. 5, 60) strongly
widened, with a large inner spine on anterio-lateral margin; its outer and inner sides without apical spine ventrally.
Hind tibia without apical spur dorsally (Fig. 8). Last male tergite (Fig. 63) long, with a narrow median incision on
posterior margin. Ovipositor (Fig. 68) slightly shorter than abdomen. For more details, see the descriptions of M.
desutterae sp. nov. and M. meridianus sp. nov. below.
This new genus appears to be closely related to Oncodopus judging by the structure of pronotum, fore femur,
hind tibia and by the slightly protruding male tegmina in M. desutterae. It differs from Oncodopus in the tridentate
fastigium of its vertex (one simple, elongated spine in Oncodopus), protruding male tegmina (almost covered by
pronotum in Oncodopus), the elongated hind margin of the last male tergite (very short in Oncodopus) and the
wider head especially in the female. It shares with Colossopus the tridentate fastigium of the vertex, the elongated
hind margin of the last male tergite and the strongly protruding tegmina of the male in one species, M. meridianus.
It differs from Colossopus in the morphology of the fastigium of the vertex (with very short arms all of which are
directed slightly upwards whilst in Colossopus the longer 2 lateral arms are parallel and directed slightly
downwards and the longer middle arm is upcurved), the shape of pronotum, the structure of the fore femur (the new
genus lacks an apical spine on both sides ventrally whilst Colossopus possesses such spines), the hind tibia without
Body length Length of
pronotum
Width of
pronotum
Length of
fore femur
Width of
fore femur
Fore tibia Hind femur Ovipositor
O. zonatus M 41.75±11.25
N=14
14.2±3 13.25±3.2
5
14.4±3.6 5.85±1.65 14.2±4.3 16.3±3.4
F 45.5±10.5
N=15
11.9±1.8 11.95±1.65 13.35±1.95 4.95±0.95 13.5±1.5 17.95±1.85 20.3±1.6
O. janetae M 44.5±9.5
N=10
14.95±3.8
5
13.85±4.0
5
15.5±5 6.3±2.1 15.5±5.3 16.75±4.25
F 43.05±8.05
N=18
11.1±2.2 11.85±2.85 12.2±3.2 4.3±1.1 11.9±3.1 17.8±4.2 21.55±6.45
O. brongniarti M 49.1±9.1
N=3
15.45±1.9
5
13±1.8 15.1±2.1 6.3±0.6 14.3±2.3 17.45±2.65 ―
F 51.95±10.25
N=3
13±2 12.85±2.4
5
14.4±3 5.4±0.9 13.8±3.2 19.3±4.7 23.6±1.7
O. saussurei M 25.45±5.45
N=4
8.65±0.85 7.35±0.85 7.1±0.8 3±0.6 6.85±0.6
5
9.45±0.45
F 27.65±4.75
N=6
7.2±1.1 7.9±0.9 6.95±0.85 2.95±0.45 6.95±1.0
5
10.65±1.15 13.5±1.4
O. soalalaensis M 32.45±1.65
N=3
10.6±0.2 9.45±0.15 8.65±0.15 3.7±0.1 9.05±0.3
5
12.4±0.6
F 32.2
N=1
9 9.6 9.1 3.4 9.9 15.4 19.7
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apical spur dorsally (Colossopus, has an apical spur dorsally on inner side), the shorter ovipositor and the wider
head especially in female.
Etymology. The generic name is derived from “Malagasy” meaning Madagascan.
Key to species of Malagasopus
1 Depression of typical sulcus of pronotum narrow and deep (Figs. 61, 67); hind margin of pronotum with a shallow median
incision (Fig. 60); metazona long, 4.5 mm; prozona 1.5 times longer than metazona in male. Male tegmina covered by prono-
tum except for their apical margin (Figs. 60, 61). Fore tibia with 4 ventral spines on its inner side (Fig. 60). Male cercus as in
Fig. 64. Subgenital plate as in Fig. 65. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. desutterae sp. nov.
- Depression of typical sulcus of pronotum broad and relatively shallow (Fig. 71); hind margin of pronotum with a deep median
incision (Fig. 70); metazona short, 1.9 mm; prozona 3.4 times longer than metazona in male. Only basal part of male tegmina
covered by pronotum, a large part visible from above (Figs. 70, 71). Fore tibia with 5 ventral spines on inner side (Fig. 70).
Male cercus as in Fig. 74. Subgenital plate as in Fig. 75 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. meridianus sp. nov.
Malagasopus desutterae sp. nov.
(Figs. 2, 5, 8, 60–69,111, 118)
http://lsid.speciesfile.org/urn:lsid:Orthoptera.speciesfile.org:TaxonName:499690
Type locality. S. Madagascar, Beheloka. Holotype male (MNHN).
Description. Male (Holotype): Head wide, 1.2 times longer than its width. Fastigium of vertex (Figs. 2, 60, 61)
short and broad, 1.2 times narrower than eye, 1.5 times wider than antennal scape; triarmed, its arms short. Face
slightly convex, almost flattened, with 2 rows of vertical tubercles, and 2 spiniform tubercles placed both sides of
frons. Fastigium of frons (Fig. 61) with a short, distinct spiniform tooth. Pronotum (Figs. 60, 61) slightly longer
than wide; depression of typical sulcus very deep and distinct; anterior margin convex, posterior margin with a
rounded median incision; prozona 1.5 times longer than metazona in middle, 2.8 times longer than its height.
Tegmina (Figs. 60, 61) slightly protrude from under pronotum, only their apex visible from above; reaching
slightly beyond of posterior margin of metanotum. Prothoracic auditory spiracle fully concealed under lateral lobe
of pronotum. Mesothoracic auricle small, dorsal half covered by lateral lobe of pronotum, its inner gap elliptical;
without ventral lobe. Fore femur (Figs. 5, 60) with a small inner spine on anterio-lateral margin, 4 ventral spines on
inner side, no apical spine ventrally, outer side without ventral spines. Fore tibia (Fig. 60) with 4 strong inner and 4
smaller outer spines ventrally. Mid and hind femora without spines. Hind tibia with 6 dorsal spines on outer side;
without apical spur dorsally (Fig. 8); with 4 apical spurs ventrally, inner 2 slightly shorter than outer ones.
Prosternum with 2, U-shaped spines, not reaching to level of ventral margin of fore coxa. Fore coxa with a forward
projecting dorsal spine. Last abdominal tergite (Fig. 63) long and broad, 2.6 times wider than long, posterior
margin elongated with a very narrow small median incision, like a split. Cercus (Fig. 64) short and broad, apical
part strongly incurved; with 2 arms, dorsal arm short and narrow, pointed with a sharp tooth, ventral arm long,
broad with a larger apical tooth. Subgenital plate (Fig. 65) almost twice as long as its width, with a distinct median
carina; posterior margin with a wide triangular median incision; its styli small, almost rectangular, longer than
posterior incision of plate.
Female: Head very wide, almost rounded in frontal view, only 1.1 times longer than its width. Fastigium of
vertex (Figs. 66, 67) short and broad; triarmed, all arms directed slightly upwards; as wide as diameter of eye and
almost twice as wide as antennal scape. Fastigium of frons (Fig. 67) with a short spiniform tooth. Pronotum (Figs.
66, 67) wider than long, typical sulcus very distinct like a deep depression; anterior margin convex, posterior
margin concave; almost twice as long as its height; prozona 2.5 times longer than metazona. Tegmina strongly
reduced, like small scales, covered by pronotum. Prothoracic auditory spiracle almost rounded, only a small part
visible in lateral view. Mesothoracic auricle like an oval bowl, without ventral lobe. Forelegs as in male but larger.
Mid and hind legs as in male but hind tibia with very small 5 dorsal spines on inner side. Prosternum with 2, U-
shaped spines. Last abdominal tergite very narrow and broad, with a distinct narrow median incision which
separates the hind margin into 2 rounded lobes. Cercus conical, short and broad, its apex slightly incurved.
Subgenital plate (Fig. 69) short and broad, median part swollen, 2.6 times wider than long, triangular but hind
margin with a large deep incision. Ovipositor (Fig. 68) sword shaped, slightly upcurved, 1.5 times longer than hind
femur and slightly shorter than abdomen.
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FIGURES 60–69. Malagasopus desutterae gen. nov. et sp. nov. Male: 60 forelegs, head and pronotum, dorsal view, 61 head
and pronotum, lateral view, 62 habitus, dorso-lateral view, 63 last abdominal tergite, 64 cercus, 65 subgenital plate. Female: 66
forelegs, head and pronotum, dorsal view, 67 head and pronotum, lateral view, 68 habitus, dorso-lateral view, 69 subgenital
plate. Scale bars 1 mm.
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Colour (dry specimens). Body various shades of brown in male and milky brown in female. Lower margin of
frons and mandibles dark, reddish brown. Labrum light, yellowish brown. Spines of fore femur and basal parts of
fore tibial spines black. Dorsal surface of fore femur with black stripes and black spots; outer sides of mid and hind
femora with irregular, large black spots and stripes. Pronotum with 2 median and 2 lateral short black longitudinal
stripes in prozona; metazona with several dark brown spots. Visible part of male tegmina with several small black
spots. Abdominal tergites reddish brown in male and milky brown in female, with indistinct irregular transverse
dark stripes. Basal third of ovipositor milky brown, apical two thirds dark.
Diagnosis. This new species shows its close affinity to M. meridianus sp. nov. by the structure of the legs, the
last male tergite, the subgenital plate and the cercus. It differs from it in the shape of male pronotum with longer
metazona (4.5 mm in the middle), the length of the male tegmina (only hind margin visible from above, almost
fully covered by pronotum), the structure of the male cercus, the number of spines on the fore tibia (4 inner and 4
outer spines ventrally in this species, and 5 inner and 5 outer spines in M. meridianus sp. nov.). Although it has a
somewhat similar pronotum and tegmina to Oncodopus spp. the generic characters given in the table 1 and under
the genus Malagasopus show that these taxa are very different.
Material examined. Madagascar sud, Beheloka, April 1963, 1 male (Holotype), 2 males nymph, 2 females
nymph (MNHN). Madagascar, Toliara Prov., Mahafaly Plateau, 6.2 km 74 ENE Itampolo, elev. 80 m, 21–25 Feb.
2002, 24
°
39'13" S, 43
°
59'48" E, 1 female (leg. Fisher, Griswold et al.), in spiny forest thicket (CAS).
Measurements (mm). Holotype (male): Body length: 31.2; pronotum length: 11.9; width of pronotum: 10.3;
fore femur length: 8.2; width of fore femur: 3.4; fore tibia length: 8.4; hind femur length: 12.4. See Table 3 for the
measurement of the female.
Etymology. This new species name is dedicated to our colleague Dr. Laure Desutter-Grandcolas from the
MNHN Paris in recognition of her very important contributions to Orthoptera research. She very kindly assisted M.
Ünal during his research visits to the MNHN and loaned the specimens of this new species as well as many others.
Distribution. South-west of Madagascar (Fig. 111). Toliara Province: Beheloka, NE Itampolo.
Habitat. This species is known from spiny forest. Living specimens have not been studied.
Phenology. Adults have been found in February and nymphs in April.
TABLE 3. Measurements (mm) of the body parts of Malagasopus spp. M male, F female.
Malagasopus meridianus sp. nov.
(Figs. 70–75 ,111)
http://lsid.speciesfile.org/urn:lsid:Orthoptera.speciesfile.org:TaxonName:499691
Type locality. Madagascar, Toliara, Raharizonina. Holotype male (MNHN).
Description. Male (Holotype): Head wide, 1.3 times longer than its width. Fastigium of vertex (Figs. 70, 71)
with tridentate, lateral 2 denticles very small, but probably not developed sufficiently (its normal form is probably
as in M. desutterae, but it may has been damaged during immature stages); its basal part 1.5 times narrower than
eye, 1.3 times wider than antennal scape. Face slightly convex, with distinct tubercles. Fastigium of frons with a
small spiniform tooth. Pronotum (Figs. 70, 71) slightly longer than wide, 3.1 times longer than its height; anterior
margin convex, posterior margin with a strong median incision; depression of typical sulcus wide and shallow;
prozona 3.4 times longer than metazona in middle. Tegmina (Figs. 70, 71) strongly protrude from under pronotum,
a large portion visible from above, reaching to slightly beyond posterior margin of metanotum. Prothoracic
auditory spiracle concealed under lateral lobe of pronotum. Mesothoracic auricle small, inner gap elliptical, ventral
lobe distinct. Fore femur (Fig. 70) with a small inner spine on anterio-lateral margin; with 5 ventral spines on inner
side, no apical spine ventrally; outer side of fore femur without ventral spines. Fore tibia (Fig. 70) with 5 large
Body
length
Length of
pronotum
Width of
pronotum
Length of
fore femur
Width of
fore femur
Fore
tibia
Hind
femur
Ovipositor
M. desutterae M 31.2 11.9 10.3 8.2 3.4 8.4 12.4
F 41 11.8 12.5 10.4 3.9 10.6 14.8 22.2
M. meridianus M 33.1 10.6 9.9 8.9 3.3 9.4 13.6
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inner and 5 smaller outer spines ventrally. Mid and hind femora without spines. Hind tibia with 5 dorsal spines on
outer side, without dorsal apical spur; with 4 apical spurs ventrally, inner 2 slightly shorter than outer ones.
Prosternum with 2 U-shaped spines, not reaching to level of ventral margin of fore coxa. Fore coxa with a dorsal
spine projecting forward. Last abdominal tergite long and broad, 2.2 times longer than wide, posterior margin
elongated with a small, very narrow median incision, like a split. Cercus short and broad, strongly incurved
apically; with 2 incurved arms, dorsal arm short and broad, pointed with a distinct tooth, ventral arm longer,
pointed with a larger tooth. Subgenital plate (Fig. 75) 1.3 times longer than wide, with a distinct median carina;
posterior margin with a narrow triangular median incision; its styli small and rounded, almost as long as depth of
posterior incision of plate.
FIGURES 70–75. Malagasopus meridianus gen. nov. et sp. nov. Male: 70 forelegs, head and pronotum, dorsal view, 71 head
and pronotum, lateral view, 72 habitus, dorso-lateral view, 73 last abdominal tergite, 74 cercus, 75 subgenital plate. Scale bars 1
mm.
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Female: Unknown.
Colour (dry specimens). Body light brown with some darker shades. Lower margin of frons and mandibles
dark reddish brown. Spines of forelegs dark. Wing membrane between veins of apical part of tegmina dark. Hind
margins of abdominal tergites with transverse light milky brown bars.
Diagnosis. This new species is similar to M. desutterae sp. nov. in the structure of its legs, last male tergite,
subgenital plate and cercus. It different from M. desutterae in the shape of male pronotum with shorter metazona
(1.9 mm in the middle), the length of the male tegmina (a large part visible from above, not fully covered by
pronotum), structure of the male cercus, the spine number of the fore tibia (it has 5 inner and 5 outer spines
ventrally whilst M. desutterae has 4 inner and 4 outer spines). Whilst the pronotum and tegmina of M. meridianus
are similar to Colossopus spp. there are many other differences (see the generic characters given in the Table 1 and
under the genus Malagasopus).
Material examined. Madagascar, Sud Quest, Tulear, P.K. 860, Raharizonina, 7.1959, 1 male (Holotype)
(MNHN).
Measurements (mm). Holotype (male): Body length: 33.1; pronotum length: 10.6; width of pronotum: 9.9;
fore femur length: 8.9; width of fore femur: 3.3; fore tibia length: 9.4; hind femur length: 13.6.
Etymology. The new species name “meridianus” means southern in Latin. This species in common with many
of the other species in this paper, is found in South Madagascar.
Distribution. South-west of Madagascar (Fig. 111). Toliara Province: S.E. Toliara, Raharizonina.
Habitat. The region in which this species occurs has spiny forest. Living specimens have not been studied.
Phenology. This species has been found in July as an adult.
Remarks. Lateral denticles of fastigium of vertex are very small and not in natural forms. Possibly they may
have been damaged during immature stages.
Colossopus Saussure, 1899
Saussure 1899: 627. Type species: Colossopus grandidieri Saussure, 1899.
Karny 1912: 8, 39; Kirby 1906: 270; Chopard 1948: 125; Gorochov 1988: 176; Otte 1997: 27; Ünal & Beccaloni 2008: 34.
Diagnosis. Body relatively depressed dorso-ventrally (Fig. 78). Fastigium of vertex distinctly elongated, with 3
long spiniform arms (Figs. 3, 76). Facet almost flattened, with distinct tubercles four of which larger, spiniform.
Male tegmina wavy, protruding from under pronotum and almost half of them visible from above (Figs. 76, 77).
Tympanal organ on fore tibia slit shaped. Mesothorax with a large seashell shaped mesothoracic auricle. Forelegs
(Fig. 76, 83) strongly developed, large, with very large and robust spines. Fore femur thickened, with a large inner
spine on anterio-lateral margin; its outer and inner sides armed with a large apical spine ventrally (Figs. 6, 76).
Hind tibia with an apical spur dorsally on inner side (Fig. 9). Male last tergite long, posterior margin elongated
backward, with a distinct narrow median incision (Fig. 80). Ovipositor longer than abdomen (Fig. 85).
This genus is most closely related to Malagasopus gen. nov. and Oncodopus Brongniart. See Table 1 for
comparison of these genera.
Key to species of Colossopus
1 Body small (Tab. 4). Opening of prothoracic auditory spiracle very small (Fig. 101). Hind margin of pronotum with a shallow
rounded incision (Fig. 98). Hind femur without ventral spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. parvicavus sp. nov.
- Body distinctly larger (Tab. 4). Opening of prothoracic auditory spiracle wide (Figs. 79, 90). Hind margin of pronotum with a
deep angular incision (Fig. 76, 87). Hind femur with ventral spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Body dark (Figs. 78, 85). Pronotum and forelegs black (Figs. 76, 83). Male cercus long and narrow, pointed with a strong api-
cal tooth (Fig. 81). Ovipositor black, very long (at least 35 mm); twice as long as hind femur and 1.5–1.6 times longer than
abdomen (Fig. 85) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. grandidieri
- Body very light in colour, almost without black markings (Figs. 89, 96). Pronotum and forelegs milky or light brown (Figs. 87,
94). Male cercus short and broad, its apex blunt, with a small apical tooth (Fig. 92). Ovipositor light brown, shorter (at most
29.8 mm), 1.5–1.6 times longer than hind femur and slightly longer than abdomen (Fig. 96) . . . . . . . . . . . . .C. redtenbacheri
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We hypothesise that these three species are related in the following way: (grandidieri + redtenbacheri) +
parvicavus.
Colossopus grandidieri Saussure, 1899
(Figs. 3, 9, 76–86, 111, 119–127)
http://lsid.speciesfile.org/urn:lsid:Orthoptera.speciesfile.org:TaxonName:15119
Colossopus grandidieri: Saussure 1899: 628; Kirby 1906: 270; Karny 1912: 39; Otte 1997: 27; Ünal & Beccaloni 2008: 35;
Hollier & Heads 2015: 310.
Type locality. Madagascar. Lectotype male (MHNG).
Redescription. Male: Body size large (Tab. 4). Fastigium of vertex (Figs. 3, 76) forming a long horn with 3
arms, upper 2 arms short and equal, lower one long, placed between the upper arms, slightly downcurved; its basal
part 1.7 times narrower than diameter of eye and 1.1 times wider than antennal scape. Fastigium of frons (Fig. 77)
with a simple, spiniform tooth. Face strongly flattened with distinct irregular tubercles. Pronotum (Figs. 76, 77)
large, almost as long as wide; 3.8 times longer than high; anterior margin convex, posterior margin with a deep
angular median incision; prozona 2.8 times longer than metazona. Tegmina (Figs. 76, 77) protruding from under
pronotum, reaching 2/3 of the way down metanotum; its upper surface wavy. Prothoracic auditory spiracle (Fig.
79) almost rounded, its dorsal half concealed under lateral lobe of pronotum in lateral view. Mesothoracic auricle
(Fig. 79) long, its inner gap oval without or with an indistinct ventral lobe. Prosternum with 2 long spines,
distinctly V-shaped. Fore femur (Fig. 76) with a large inner spine on anterio-lateral margin; with a large inner and a
large outer apical spine ventrally (in one male with an additional ventral spine on outer side apically); with 4–5
additional ventral spines on inner side. Fore tibia (Fig. 76) with 5 very large inner and 6–7 smaller outer spines
ventrally. Mid femur with 2–4 and hind femur with 2–4 ventral spines on outer side. Hind tibia with 5–7 dorsal
spines on inner side, with a single apical spur on inner side dorsally (Fig. 9); with 4 apical spurs ventrally, inner two
shorter than outer ones. Last abdominal tergite (Fig. 80) broad, posterior part elongated backward, with a deep
narrow median incision. Cercus (Fig. 81) long, narrow and slender, apical third sharply narrowed with a large,
slightly incurved apical tooth. Subgenital plate (Fig. 82) almost as long as wide; its styli long, much longer than
depth of posterior incision.
Female: Head as in male, relatively slightly larger. Fastigium of vertex (Figs. 83, 84) as wide as half eye
diameter, its basal part 1.1 times wider than antennal scape. Fastigium of frons (Fig. 84) with a simple, large,
slightly downcurved tooth. Pronotum (Figs. 83, 84) short and broad, anterior margin convex, posterior margin
broadly concave; prozona 1.4 times narrower and 3.9 times longer than metazona. Tegmina strongly reduced, like a
very small scale. Dorsal half of prothoracic auditory spiracle covered by pronotum. Mesothoracic auricle not
covered by pronotum. Fore femur (Fig. 83) with a large inner spine on anterio-lateral margin; with a large inner and
a large outer apical spine ventrally; with 5–6 additional inner spines ventrally. Fore tibia (Fig. 83) with 5 inner and
6–7 outer strong spines ventrally. Mid femur with 3 and hind femur with 1–3 outer spines ventrally. Hind tibia with
6–8 dorsal spines on inner side, with a single inner apical spur dorsally; with 4 apical spurs ventrally, inner 2
slightly shorter than outer ones. Last tergite very short and broad, posterior margin with a narrow, median incision.
Cercus conical, pointed at apex like a spine. Subgenital plate (Fig. 86) short and broad, with a wide median carina,
its lateral lobes with transverse ridges. Ovipositor (Fig. 85) almost straight, 2–2.3 times longer than hind femur and
1.5–1.6 times longer than abdomen; apical part of ventral valve with small teeth.
Colour (dry specimens). Body dark, black and dark brown. Face dark reddish brown; labrum light brown;
mandibles and upper surface of head black. Pronotum black with some irregular dark brown spots. Male tegmina,
mesonotum and metanotum in both sexes yellowish milky brown. Forelegs reddish brown and reddish black. Mid
and hind tibia yellowish milky brown. Abdominal tergites black or dark reddish brown in some specimens.
Sternites yellowish milky brown in male, darker in female. Ovipositor black.
Diagnosis. The shape of fastigium of the vertex and the forelegs indicates that this species is closely related to
C. redtenbacheri. It differs from it in the shapes of the pronotum, the male cercus and the ovipositor, in the ratios of
ovipositor/hind femur and ovipositor/abdomen, in its distinctly darker body colour and also its larger size. It differs
from C. parvicavus sp. nov. by its very larger body size, dark colouration, widely open prothoracic auditory
spiracle, the shape of male pronotum, and the presence of ventral spines on the hind femur.
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FIGURES 76–86. Colossopus grandidieri. Male: 76 forelegs, head and pronotum, dorsal view, 77 head and pronotum, lateral
view, 78 habitus, dorso-lateral view, 79 prothoracic auditory spiracle and mesothoracic auricle, 80 last abdominal tergite, 81
cercus, 82 subgenital plate. Female: 83 forelegs, head and pronotum, dorsal view, 84 head and pronotum, lateral view, 85
habitus, dorso-lateral view, 86 subgenital plate. Scale bars 1 mm.
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Material examined. Madagascar, 3 females (leg. M. Grandidier) (Paralectotypes); Prov. De Tulear, Bas
Fiherana [Fiherenana], 1906, 1 male, 2 females (leg. F. Geay) ; Madagascar (S.O.), Forest du Fiherana
[Fiherenana], 1905, 1 male, 1 male nymph (leg. F. Geay); Madagascar sud, 14.5.1963, 1 female nymph (leg. D.
Wintrebert); Madagascar, sud-wat, Efeda, 29.4.1963, 1 female nymph (leg. D. Wintrebert); Madagascar,
Fianarantsoa, Anpamaherana [Ampamaherana], station forestire, 1 female; Madagascar, Tulear, Manombo,
10.5.1956, 1 male (all in MNHN). Madagascar, S. Centr. Mdg., 2 males; Toliara Province, Forest near village of
Mangily, c. 27 km N of Toliara (-23.15, 43.60) [23
°
8'60'' S, 43
°
36'0'' E], 19.11.2004, Coll. G.W. Beccaloni, 2
males, 1 female (all in NHMUK). 1 female (det. Karny), without locality label (NMW). Madagascar, Toliara Prov.,
Forét de Tsinjoriaky 6.2 km 84 E Tsifota, elev. 70 m, 6–10 Mar. 2002, 22
°
48'8" S, 43
°
25'14" E, 1 female (leg.
Fisher, Griswold et al.), in spiny forest thicket; Madagascar, Toliara Prov., Lake Ranobe, elev. 20 m, 23
°
02'524" S,
43
°
37'214" E, 13–21 May 2003, 1 male, 1 female (leg. Frontier, Wilderness), spiny forest; Madagascar, Toliara
Prov., Mahafaly Plateau, 6.2 km 74 ENE Itampolo, elev. 80 m, 21–25 Feb. 2002, 24
°
39'13" S, 43
°
59'48" E, 1
female (leg. Fisher, Griswold et al.), in spiny forest thicket (CAS). In addition, 2 adult females were photographed
but not collected in spiny forest near Mangily, north of Toliara, Toliara Province by Martin Bader on 15.3.2015
(photographs and video footage were examined by the authors). They were found during the day hiding under bark
attached to dead trees.
Measurements (mm). Lectotype (male): Body length: 53; pronotum length: 14; width of pronotum: 12.5; fore
femur length: 16.9; width of fore femur: 6.2; fore tibia length: 14; hind femur length: 18. See Table 4 for the range
of measurement of the other specimens including female.
Distribution. South-west and south-central Madagascar (Fig. 111). Toliara Province: Toliara, Fiherenana,
Mangily, Efeda, Manombo, Tsifota, Lake Ranobe, NE Itampolo. Fianarantsoa Province: Anpamaherana
[Ampamaherana].
Habitat. This species was collected in spiny forest. The specimens collected by the second author were found
under loose bark still attached to dead trees.
Phenology. This species has been collected or observed between February and May: in November and March
as adults, and in April and May as nymphs.
Remarks. The record for Anpamaherana [Ampamaherana] is dubious since this locality is located in the
subhumid bioclimatic zone (Cornet 1974), rather than the subarid zone like all of the other collecting localities for
this species (see remarks for O. janetae).
Life history. C. grandidieri has been bred in captivity by several hobbyists in Europe and to our knowledge it
is the only one of the species discussed in this paper that has been studied in this respect. What follows is a
compilation of information from several people who have reared this species.
C. grandidieri is omnivorous, feeding on a wide range of leaves and fruit, as well as living and dead insects,
and manufactured food, e.g. lettuce, Rubus and Quercus leaves, rose flowers, zucchini, cucumber, banana (Röhrs
2017), living crickets, cockroaches, moths and Zophobas morio larvae (Fritzsche 2008), dried Tenebrio molitor
larvae, dog food and fish flakes (Guillermo González, pers. commun., 2017). All instars of both sexes have
raptorial forelegs, which are used to grasp living prey as well as being used for defence (see below).
The pale brown cigar-shaped eggs are deposited singly into soil (Röhrs 2017) and females can lay 150–200
during their lifetime (Tänzler 2012). Eggs measure c. 6mm in length when first laid (Fig. 119 right) and they
increase in volume over several weeks (Tänzler 2012), presumably by absorbing water from the surrounding soil
(Figs. 119 left, 120). The eggs hatch in ca. 3 months, although some may take more than a year to emerge (Röhrs
2017). Hatching appears to be triggered by alternating dry and wet conditions (Röhrs 2017), so it is possible that
the eggs which took a year to hatch did not receive the appropriate wet/dry regime.
First instar nymphs (Fig. 121) measure ca. 8mm in body length (Tänzler 2012), are dark brown/black, and lack
the turquoise markings on their hind- and midlegs which third instar and older nymphs and adults have . Third
instar and older nymphs have pale markings on their pronota, legs and tergites (Figs. 122–123), whereas, in adults
these markings are replaced by black (Figs. 124, 126). Nymphs take 4–5 months to reach adulthood and adults can
live for about six months (Tänzler 2012). It is unclear how many nymphal instars there are.
C. grandidieri is nocturnal and first and second instar nymphs hide singly in holes in dead bark during the day
(Guillermo González, pers. commun., 2017). Older nymphs and adults will rest together during the day in groups,
but smaller individuals may be eaten by larger ones (Tänzler 2012). In their natural habitat adults of several
species, including C. grandidieri, have been observed to rest by day as male-female pairs, never in groups of more
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than two individuals (see Discussion), so the aggregations seen in captivity may be the result of a lack of
harbourages.
Adult males and females have very similar colouration, except that the labrum of the male is yellow-orange
(Fig. 127), whilst that of the female is orange-red (Fig. 125). When disturbed adults and larger nymphs rear up,
open their mandibles and raise and spread their forelegs (Fig. 125). Adult males stridulate loudly when in this
defensive position (George Beccaloni, pers. obs.; Tänzler 2012; Guillermo González, pers. commun., 2017),
whereas the adult females do not make any audible sound (Martin Bader, pers. commun., 2017; Guillermo
González, pers. commun., 2017). If an aggressor approaches closely enough the insect will attempt to grab it with
its forelegs and bite it. The threat display of only one other species, O. brongniarti, has been observed (Fig. 115). It
was an adult female and the defensive display was the same as that of the female C. grandidieri (Martin Bader,
pers. commun., 2017).
Colossopus redtenbacheri (Brongniart, 1897)
(Figs. 6, 87–97, 111, 128)
http://lsid.speciesfile.org/urn:lsid:Orthoptera.speciesfile.org:TaxonName:15117
Oncodopus redtenbacheri: Brongniart 1897: 208; Kirby 1906: 270; Otte 1997: 32.
Colossopus (Oncodopus) redtenbacheri: Saussure 1899: 628.
Colossopus redtenbacheri: Karny 1912: 39; Otte 1997: 27.
Type locality. Madagascar. Lectotype male (in alcohol) (MNHN), here designated.
Redescription. Body size is intermediate between the other two members of the genus (Table 4). Fastigium of
vertex (Fig. 87, 88) with three-pronged horn; upper 2 arms short and equal, slightly upcurved; lower one longer and
situated between the upper arms; 1.4 times narrower than diameter of eye; its basal part 1.1 times wider than
antennal scape. Fastigium of frons (Fig. 88) with a large spiniform tooth. Face almost flattened with distinct
tubercles some of which are like short spines. Pronotum (Fig. 87, 88) slightly longer than wide, 3.4 times longer
than its height; anterior margin convex, posterior margin with a deep angular median incision; prozona 1.5 times
longer than metazona. Tegmina (Fig. 87, 88) wavy, clearly visible from above, reaching 4/5 of the way down
metanotum. Prothoracic auditory spiracle (Fig. 90) almost rounded, almost fully covered by lateral lobes of
pronotum. Mesothoracic auricle (Fig. 90) oval, visible in lateral view, longer than wide, without ventral lobe; its
inner gap long and elliptical. Prosternum with two, V-shaped spines, their apices not reaching to level of ventral
margin of fore coxa. Fore coxa with a short dorsal spine. Fore femur (Figs. 6, 87) with a large inner spine on
anterio-lateral margin; with a large inner and a large outer apical spine ventrally; with 5–6 additional ventral spines
on inner side. Fore tibia (Fig. 87) with 5 inner and 4–7 outer spines ventrally, inner ones larger. Mid femur with 2–
4 and hind femur with 1–3 outer spines ventrally. Hind tibia with 5–7 dorsal spines on inner side, with a single
inner apical spur dorsally; with 4 apical spurs ventrally, inner two slightly shorter than outer ones. Last abdominal
tergite (Fig. 91) twice as wide as its length, posterior margin with a deep narrow incision. Cercus (Fig. 92) short,
basal part broad, end of basal part with a distinct projection on inner margin; apical part narrower and slightly
incurved, apex blunt, slightly rounded, with a small apical tooth. Subgenital plate (Fig. 93) almost as long as its
width, its styli slightly longer than depth of posterior incision of plate.
Female: Head as in male. Fastigium of vertex (Figs. 94, 95) 1.4 times narrower than diameter of eye, 1.3 times
wider than antennal scape. Pronotum (Figs. 94, 95) with long, narrow prozona and wide, short metazona; prozona
1.4 times narrower and 4.3 times longer than metazona; anterior margin convex, posterior margin strongly concave.
Tegmina strongly reduced to very small scale-like projections. Prothoracic auditory spiracle almost concealed by
lateral lobe of pronotum. Mesothoracic auricle smaller than that of male, its inner gap narrow elliptical, not covered
by pronotum. Fore tibia (Fig. 94) with 5–6 ventral spines on outer side; mid femur with 1–4 and hind femur with 1–
4 outer spines ventrally; hind tibia with 6–7 dorsal spines on inner side, with an inner apical spur dorsally and 4
apical spurs ventrally. Last abdominal tergite broad and narrow, posterior margin with a small distinct median
incision. Cercus short, conical, its apical part slightly incurved, pointed at apex. Subgenital plate (Fig. 97) 3.5 times
broader than long, with a broad rectangular median carina; lateral lobes with many transverse ridges. Ovipositor
(Fig. 96) straight, 1.5–1.6 times longer than hind femur and slightly longer than abdomen; apical part of ventral
valve with small teeth.
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FIGURES 87–97. Colossopus redtenbacheri. Male: 87 forelegs, head and pronotum, dorsal view, 88 head and pronotum,
lateral view, 89 habitus, dorso-lateral view, 90 prothoracic auditory spiracle and mesothoracic auricle, 91 last abdominal tergite,
92 cercus, 93 subgenital plate. Female: 94 forelegs, head and pronotum, dorsal view, 95 head and pronotum, lateral view, 96
habitus, dorso-lateral view, 97 subgenital plate. Scale bars 1 mm.
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Colour (dry specimens). Body with different shades of brown, mostly milky and light brown. Face darker,
reddish brown; clypeus and labrum yellowish brown. Metazona of pronotum darker with reddish brown, prozona
light brown. Fore tibia, anterior part of fore femur, knees of mid and hind legs and tarsi reddish brown, remaining
parts of legs milky brown. Male tegmina yellowish brown. Abdominal tergites reddish brown, with a very narrow
black stripe on hind margin in female. Basal part of ovipositor milky brown, its apical part light reddish brown.
Diagnosis. This species differs from C. grandidieri in the structure of the pronotum, male cercus, shorter
ovipositor, the ratios of ovipositor/hind femur and ovipositor/abdomen, clearly lighter body colour and smaller
size. It differs from C. parvicavus sp. nov. by the shape of the male pronotum, prothoracic auditory spiracle (which
has a wide opening), the presence of ventral spines on the hind femur, and the ratios of ovipositor/hind femur and
ovipositor/abdomen.
Material examined. Madagascar, 1867, 1 male (Lectotype), 1 female (Paralectotype) (leg. M. Grandidier)
(MNHN). Toliara Province, Andohahela National Park: Tsimelahy parcel, (-24.93, 46.65) [24
°
55'48'' S, 46
°
38'60''
E], 1.12.2004, Coll. leg. G.W. Beccaloni, 1 male, 1 female (NHMUK). Madagascar, Toliara Prov., Parc Nat.
d’Andohahela, Forét d’Ambohibory, 1.7 km 61 ENE Tsimelahy, 36.1 km 308 NW Tolagnaro, 16–20.1.2002,
24
°
55'48" S, 46
°
38'44" E, 2 males, 1 female (Fisher, Griswold et al.), tropical dry forest; Castlot 006432 [locality
unknown], 3 males, 3 females, 1 male nymph (CAS).
Measurements (mm). Lectotype (male): Body length: 40; pronotum length: 10.6; width of pronotum: 10; fore
femur length: 12.9; width of fore femur: 4.6; fore tibia length: 10.3; hind femur length: 14.4. See Table 4 for the
range of measurement of the other specimens including females.
Distribution. South-eastern Madagascar (Fig. 111). Toliara Province: Toliara, Andohahela.
Habitat. This species has been found in transitional tropical dry forest. The specimens collected by the second
author were found under loose dead bark.
Phenology. Adults have been found in December and January.
Colossopus parvicavus sp. nov.
(Figs. 98–108, 111)
http://lsid.speciesfile.org/urn:lsid:Orthoptera.speciesfile.org:TaxonName:499692
Type locality. Madagascar, Toliara Prov., W. Marovato. Holotype male (CAS).
Description. Male (Holotype): Body size smallest in the genus (Tab. 4). Fastigium of vertex (Figs. 98, 99)
with three-pronged horn; above 2 arms equal and shorter, directed slightly upwards; lower one placed between the
above arms, longer and slightly directed downwards; 1.5 times narrower than diameter of eye; its basal part as wide
as antennal scape. Fastigium of frons (Fig. 99) with a large simple spiniform tooth. Face slightly convex, with
tubercles some of which look like short spiniform teeth. Pronotum (Figs. 98, 99) slightly longer than wide, 3 times
longer than its height; anterior margin convex, posterior margin with a shallow rounded median incision; prozona
2.3 times longer than metazona. Tegmina (Figs. 98, 99) wavy, clearly protruding from under pronotum, reaching
beyond middle half of metanotum (in other male reaching to end of metanotum). Prothoracic auditory spiracle (Fig.
101) almost closed except for a small rounded hole which is visible in lateral view. Mesothoracic auricle (Fig. 101)
visible in lateral view, its inner gap long, oval, with a long ventral lobe. Prosternum with two long, V-shaped
spines. Fore coxa with a short dorsal spine. Fore femur (Fig. 98) with a large inner spine on anterio-lateral margin;
with a large inner and a large outer apical spine ventrally; with 5 additional ventral spines on inner side (in the other
male, inner side of fore femur with 6 ventral spines). Fore tibia (Fig. 98) with 5 inner and 6 smaller outer spines
ventrally (in other male outer side of fore tibia with 4–5 ventral spines). Mid femur with 2 ventral spines on outer
side (in other male with 1 spine); hind femur without ventral spines. Hind tibia with 6–8 dorsal spines on inner
side; with a single apical spur on inner side dorsally; with 4 apical spurs ventrally, inner 2 shorter than outer ones.
Last abdominal tergite (Fig. 102) large, elongated backwards, with a small and narrow median incision. Cercus
(Fig. 103) small, almost 1.5 mm, basal part broad, end of basal part with a small tooth on inner side; apical part
narrow, slightly incurved (in other male apical part more incurved); with a very distinct incurved apical tooth.
Subgenital plate (Fig. 104) slightly longer than wide, its styli longer than depth of posterior incision of plate.
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FIGURES 98–108. Colossopus parvicavus sp. nov. Male: 98 forelegs, head and pronotum, dorsal view, 99 head and
pronotum, lateral view, 100 habitus, dorso-lateral view, 101 prothoracic auditory spiracle and mesothoracic auricle, 102 last
abdominal tergite, 103 cercus, 104 subgenital plate. Female: 105 forelegs, head and pronotum, dorsal view, 106 head and
pronotum, lateral view, 107 habitus, dorso-lateral view, 108 subgenital plate. Scale bars 1 mm.
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Female: Head as in male, including the ratios. Pronotum (Figs. 105, 106) slightly longer than wide (in other
female slightly wider than long); prozona 4.6 times longer than metazona; anterior margin convex, posterior
margin concave. No tegmina. Prothoracic auditory spiracle almost closed except for a small rounded hole.
Mesothoracic auricle as in male, not covered by pronotum; its ventral lobe smaller than that of female. Fore femur
(Fig. 105) with a large inner spine on anterio-lateral margin; with a large inner and a large outer apical spine
ventrally; with 4–5 additional ventral spines on inner side. Fore tibia (Fig. 105) with 5 inner and 5 outer spines
ventrally; mid and hind legs as in male. Last abdominal tergite very broad and narrow, posterior margin with a
distinct median incision which separates the posterior margin into two rounded lobes. Cercus narrow, longer than
last abdominal tergite; its apex slightly incurved. Subgenital plate (Fig. 108) short and broad, hind margin
narrowed and elongated backwards; with a large, almost rectangular median carina, lateral parts of plate with
distinct transverse ridges. Ovipositor (Fig. 107) long and straight, slightly more than twice as long as hind femur
and 1.4–1.5 times longer than abdomen; apical part of ventral valve with small teeth.
Colour (dry specimens). Body milky and light brown with dark brown transverse bands. Face light brown, its
tubercles with black or dark brown tips. Legs milky brown, their spines with black tips; Pronotum light brown hind
margin with a large dark brown transverse band. Male tegmina milky brown. Hind margin of each abdominal
tergite with a thin black transverse band. Basal third of ovipositor milky brown, apical two thirds dark brown.
Diagnosis. This species is similar to C. redtenbacheri in its colouration, male cercus and subgenital plate. It
differs from it in that its prothoracic auditory spiracle opening is reduced to a small hole, that the male pronotum
has a shallow posterior incision, in the absence of ventral spines on the hind femur, in the ratios of ovipositor/hind
femur and ovipositor/abdomen, and in its smaller size. The ratios of ovipositor/hind femur and ovipositor/abdomen
are similar to C. grandidieri, but it differs from C. grandidieri by its much smaller size, its colouration (milky
brown in the new species, distinctly darker in C. grandidieri), its prothoracic auditory spiracle (with a small hole in
the new species, and a large opening in C. grandidieri), in the absence of ventral spines on the hind femur, and that
the male pronotum has a shallow posterior incision.
Material examined. Madagascar, Toliara Prov., Réserve Spéciale de cap Sainte Marie, 12.3 km 262°W
Marovato, elev. 200 m, 11–15 Feb. 2002, 25
°
34'54" S, 45
°
10'6" E, 2 males (including Holotype), 2 females (leg.
Fisher, Griswold et al.), in spiny forest thicket (CAS).
Measurements (mm). Holotype (male): Body length: 34.2; pronotum length: 9.4; width of pronotum: 8.7; fore
femur length: 10.4; width of fore femur: 4.3; fore tibia length: 9.2; hind femur length: 11.4. See Table 4 for the
range of measurements of the other type specimens.
Etymology. The species name is derived from the Latin words “parvus” and “cavum” meaning small and hole
respectively. This refers to the structure of the prothoracic auditory spiracle which has a very small opening
Distribution. Only known from the southern tip of Madagascar (Fig. 111). Toliara Province: Marovato.
Habitat. This species was found in spiny forest.
Phenology. Adults were collected in February.
TABLE 4. Measurements (mm) of the body parts of Colossopus spp. M male, F female.
Body
length
Length of
pronotum
Width of
pronotum
Length of
fore femur
Width of
fore femur
Fore tibia Hind
femur
Ovipositor
C.grandidieri M50.85±7.85
N=7
14±1.1 12.85±1.05 17.6±1.8 5.95±0.55 15.15±1.65 19.1±1.9
F55.45±5.95
N=9
13.5±0.7 14.25±1.25 17.6±0.6 6.4±0.5 15.35±1.35 20.85±0.85 42.25±7.25
C.redtenbacheri M46±6
N=7
12.65±2.05 11.4±1.4 14.45±1.55 5.25±0.65 11.8±1.5 15.95±1.55 ―
F45.35±4.65
N=6
11.75±0.25 12±1 14.65±0.85 5.4±0.3 12.1±0.4 18.05±0.85 27.9±1.9
C.parvicavus M 33.8±0.4
N=2
9.35±0.5 8.6±0.1 10.8±0.4 4.2±0.1 9±0.2 11.45±0.05
F37.45±1.25
N=2
9.3±0.5 9.25±0.15 11.7±0.3 4.45±0.05 9.55±0.15 13.75±0.75 28.5±1.5
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REVISION OF ONCODOPUS AND COLOSSOPUS
FIGURES 109–111. Maps. 109 map of Madagascar, 110 distributions of Oncodopus spp., 111 distributions of Malagasopus
spp. and Colossopus spp.
Discussion
Ecology and Life-cycle. The species discussed in this paper occur in the arid regions of central, western and
southern Madagascar. These regions have a long dry season from May to September, with little or no rainfall, and a
short wet season from December to March (Jury 2003). Madagascar has a north-south, and an east-west gradient in
precipitation, with declining precipitation towards the south and west of the island (Cornet 1974). The driest of the
regions occupied by these insects is the south-west coast, with an average annual rainfall of <400 mm (it is the
most arid part of Madagascar), whilst the wettest is the central highlands, with an average annual rainfall of 1500–
2000 mm (Szab, Raveloson & Székely 2015).
Adults of eight of the ten species discussed in this paper have only been observed from November to March.
The single known specimen of M. meridianus was collected as an adult in July and the collecting dates of adults of
O. soalalaensis were not recorded. Nymphs of six species have been observed and these were found from January
to November. From these data it would appear that the life-cycle of these tettigoniids is an annual one, timed so that
the nymphs reach adulthood at the beginning of the wet season. Eggs of C. grandidieri take about 3 months to
hatch (Röhrs 2017), so if this is the case for the other species as well it would mean that the eggs hatch towards the
end of the wet season when the vegetation is at its most luxuriant (during the dry season the majority of the plants
are leafless). Hatching appears to be triggered by alternating dry and wet conditions (Röhrs 2017), which is what
they would experience in their natural habitat during the wet season (the rainfall being low and erratic). A possible
reason why these insects spend the dry season as nymphs rather than as eggs, is that the dry season is very long (5+
months) and eggs may desiccate and die in the dry soil.
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FI G URE S 112 119 . Living specimens. 112 Oncodopus zonatus, adult male, natural resting posture, Zombitse–Vohibasia
National Park, Toliara Province. 17.11.2004. © G.W. Beccaloni, 113 Oncodopus janetae, adult female, Mangatsiaka parcel,
Andohahela National Park, Toliara Province. 30.11.2004. © S. Stammers, 114 Oncodopus brongniarti, adult female, near
village of Mangily, Toliara Province. 11.11.2004. © G.W. Beccaloni, 115 Oncodopus brongniarti, threat display of adult
female, near village of Mangily, Toliara Province. 30.12.2015. © M. Bader, 116 Oncodopus brongniarti, adult pair, natural
resting postures, Belo Sur Mer, Toliara Province. 22.11.2007. © J. Beccaloni, 117 Oncodopus saussurei, adult female, beside
R.N. 7 south of Ambohimahasoa, Fianarantsoa Province. 13.11.2004. © S. Stammers, 118 Malagasopus desutterae, male
nymph, 80 km north of Tulear, near village of Ankililoaka, Toliara Province. 6.2008 © O. Pronk, 119 Colossopus grandidieri,
eggs: newly laid eggs lower right; older (swollen) eggs upper left. Captive stock. © G.J.N. González.
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FIGURES 120–129. Living specimens. 120 Colossopus grandidieri, older (swollen) egg. Captive stock. © G.J.N. González,
121 Colossopus grandidieri, recently hatched 1st instar nymph. Captive stock. © G.J.N. González, 122 Colossopus grandidieri,
late instar male nymph. Captive stock. © G.J.N. González, 123 Colossopus grandidieri, final instar female nymph. Captive
stock. © G.J.N. González, 124 Colossopus grandidieri, adult female. Captive stock. © G.J.N. González, 125 Colossopus
grandidieri, threat display of adult female, near village of Mangily, Toliara Province. 15.3.2015. © M. Bader, 126 Colossopus
grandidieri, adult male, near village of Mangily, Toliara Province. 19.11.2004. © G.W. Beccaloni, 127 frontal view of head of
the previous individual. © G.W. Beccaloni, 128 Colossopus redtenbacheri, adult female, Berenty Private Reserve, Toliara
Province, 4.12.2004. © G.W. Beccaloni, 129 Spiny forest near village of Mangily, Toliara Province in November 2004, close to
where the second author collected Colossopus grandidieri and Oncodopus brongniarti. © G.W. Beccaloni.
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Harbourages and possible mate-guarding. The second author G. Beccaloni (GWB) collected a total of 27
specimens of 6 species of Oncodopus and Colossopus (i.e. O. zonatus, O. janetae, O. brongniarti, O. saussurei, C.
grandidieri and C. redtenbacheri) during his two trips to Madagascar at the end of the dry season in November and
December 2004 and November 2007. All were adult and they were collected during the day when they were
inactive and resting, apart from one female of C. redtenbacheri which was climbing a bush at night (Fig. 128). The
four O. saussurei individuals were found under small rocks which were resting on the soil surface, whilst all others
were discovered under bark loosely attached to dead trees, or in cavities in rotting wood, except for two O. janetae
specimens which were found in the natural cavities which occur at the base of living Dypsis decaryi palm fronds.
The same types of harbourages were also used by hissing cockroaches (Blaberidae: Oxyhaloinae: Gromphadorhini)
which GWB was also collecting (these were the only other large insects found in such harbourages). In two cases
Oncodopus were observed sharing a harbourage with hissing cockroaches: five adult O. zonatus were found under
a large piece of bark near Belo Sur Tsiribihina which also concealed groups of adult Elliptorhina sp. and
Gromphadorhina sp. cockroaches; and the four O. brongniarti from Belo Sur Mer were found under one large
piece of dead bark together with a group of adult Elliptorhina sp. cockroaches, which were aggregated not far from
where the O. brongniarti were resting. Given that the species discussed in this paper have raptorial forelegs and are
probably predacious (as C. grandidieri is known to be), it seems likely that they would prey upon hissing
cockroaches sharing their harbourages (at least on the nymphs). However, the primary reason why these
tettigoniids and hissing cockroaches probably share the same types of harbourages is simply because they are the
only semi-enclosed cavities large enough to accommodate such big insects present in the areas where the insects
are found. Such harbourages are scarce (GWB, pers. obs.) and it is likely that both the tettigoniids and the hissing
cockroaches require them for protection against the intense daytime heat and predators. Note that most of GWB's
collecting localities had sandy soils and rocks were absent from the soil surface, so harbourages in these areas were
limited to bark on dead trees, cavities in rotting wood, or the leaf bases of palms. It is likely that, as with C.
grandidieri in captivity, these tettigoniids rest in harbourages during the day and come out at night in order to feed,
lay eggs etc. GWB observed a female of C. redtenbacheri on a small bush at night in Berenty Reserve (Fig. 128),
but other observations of these insects at night in their natural habitat are currently lacking.
Curiously, 10 of the 27 specimens of Oncodopus and Colossopus collected by GWB were observed resting in
their harbourages very close together as adult male-female pairs. The other 17 individuals he observed were found
singly (if one or more other individuals were present they went unseen), except for the 3 adult males and 2 adult
females of O. zonatus found near Belo Sur Tsiribihina which were all under a one very large piece of bark on a
dead standing tree. When the bark was pulled off the trunk the insects fell to the ground, so it is unknown how they
were originally resting. The specimens observed as pairs were as follows: the female (Fig. 113) and (holotype)
male of O. janetae collected in a cavity in a decaying tree branch in the Mangatsiaka parcel of Andohahela
National Park; a male (Figs. 126–127) and female of C. grandidieri found inside a rotten tree trunk in forest near
Mangily village; the female (Fig. 117) and (holotype) male of O. saussurei found under a stone near
Ambohimahasoa; and the two pairs of O. brongniarti collected under the same piece of dead bark at Belo Sur Mer
(Fig. 116 shows a pair resting naturally as found). This phenomenon of resting in male-female pairs appears to be
mate-guarding behaviour of a type not previously documented in the Tettigoniidae (Karim Vahed, pers. commun.,
2017). Given that four species were observed exhibiting this behaviour, and that they were found in widely
separated localities in different habitat types (sclerophyllous woodland, spiny forest (Fig. 129), and littoral forest
bordering on to dry deciduous forest), it seems likely that the same behaviour is exhibited by the other species
discussed in this paper.
A plausible reason for this behaviour is that the males are guarding their mates against other males which may
wish to copulate with them. In order to do this effectively they may need to guard the female for the duration of her
active reproductive life, which in C. grandidieri (and probably other species) is about 6 months. We speculate that
the behaviour of resting in harbourages during the day has facilitated the evolution of this mate-guarding
behaviour, since suitable harbourages are scarce and individuals may need to use the same one for their entire lives,
thus making it easier for males to associate with particular females and guard them. Many questions remain
unanswered, however, including: whether the adults mate once or several times; whether males defend their
harbourages against other males; and whether the males accompany and guard the females when they are outside
their harbourage at night. If the latter is true it could explain why these insects might be monogamous, as it would
be presumably difficult for a male to guard more than one female which is moving about to find food, egg laying
Zootaxa 4341 (2) © 2017 Magnolia Press
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227
REVISION OF ONCODOPUS AND COLOSSOPUS
sites etc. It is likely that the size of a harbourage determines the number of individuals which rest inside it, as is the
case with the New Zealand tree weta Hemideina crassicruris (Moller 1985). Moller found that large harbourages
usually contained a male and his 'harem' of up to 9 females, whereas small harbourages contained single
individuals or male-female pairs. It is possible that most of the harbourages that GWB discovered were small ones,
explaining why they contained only a single individual or a male-female pair, although his observations of two
male-female pairs of O. brongniarti occupying a single large harbourage, and 3 males and 2 females of O. zonatus
in a large harbourage, appears to go against the idea that large harbourages are defended by a single adult male
which is guarding a harem of females (as is the case with H. crassicruris).
Acknowledgements
We would like to thank Judith Marshall (NHMUK, London); Michael Ohl (MfN, Berlin); Laure Desutter-
Grandcolas (MNHN, Paris); and Mercedes París and Vincenta Llorente (MNCN, Madrid), for their assistance to
M. Ünal during his work in their museums and for the loan of many specimens; we also thank Piotr Naskrecki
(Museum of Comparative Zoology, Harvard University, Cambridge) for sending the material collected by CAS;
and Peter Schwendinger and John Hollier (MHNG, Geneva) for detailed digital photographs and information about
specimens of C. grandidieri and O. zonatus preserved in the Geneva Museum. We are also grateful to the late
Alfred Kaltenbach (Naturhistorisches Museum, Vienna); to Ros Urban (National Collection of Insects, Pretoria);
and to Margie Cochrane (Iziko South African Museum, Cape Town) for checking their collections for the presence
of Oncodopus and Colossopus material. Mustafa Ünal's work in European museums was supported by the
European Commission's Research Infrastructure Action via the following SYNTHESYS Projects: GBTAF-561,
AT-TAF-562, DE-TAF-558, FR-TAF-1955 and ES-TAF-3897. George Beccaloni would like to thank Janet
Beccaloni (NHMUK, London) for her considerable help with logistics and collecting in Madagascar; Sinclair
Stammers (UK) for permission to use still images from video footage he took; Martin Bader (Germany), Guillermo
J. Navarro González (Spain) and Olaf Pronk (Madagascar) for useful information and permission to use their
photographs; and David Lees (NHMUK, London) for his comments on this manuscript.
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... Most studies on this group in this region were conducted before 1914, when the last comprehensive paper appeared (Carl 1914). Only very recently have some papers on selected groups been published (Massa 2017a,b,c,d, Ünal andBeccaloni 2017). There are no published data on the biology of any taxon (except some observations in Ünal and Beccaloni 2017). ...
... Only very recently have some papers on selected groups been published (Massa 2017a,b,c,d, Ünal andBeccaloni 2017). There are no published data on the biology of any taxon (except some observations in Ünal and Beccaloni 2017). At present, four subfamilies of Tettigonioidea are represented by several species each in Madagascar, and Listroscelidinae is represented by a single species (all data according to Cigliano et al. 2018, abbreviated OSFO). ...
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Revision des Salomonitae
  • C Brongniart
Brongniart, C. (1897) Revision des Salomonitae, Locustidae de la Tribu des Conocephalinae. Bulletin de la Sociét philomatique de Paris, 8 (8), 120-212.