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A revision of the subgenus Parasphingonotus Benediktov & Husemann, 2009 (Orthoptera: Oedipodinae: Sphingonotini)

Authors:
  • Leibniz Institute for the Analysis of Biodiversity Change

Abstract

The recently erected subgenus Parasphingonotus is revised. The diagnostic characters are given and the species Sphin-gonotus (Parasphingonotus) radioserratus comb. n., Sphingonotus (Parasphingonotus) femoralis comb. n. and Sphin-gonotus (Parasphingonotus) turkanae comb. n. are assigned to the subgenus. Sphingonotus (Parasphingonotus) airensis is synonymized with S. (P.) femoralis, making the latter species the new type species of Parasphingonotus. Keys to the subgenera (Sphingonotus, Neosphingonotus, and Parasphingonotus) of the genus Sphingonotus and to the species of Parasphingonotus are presented. All Parasphingonotus species are re-described and further information on the distribu-tion of the species are given.
Accepted by H. Song: 4 May 2011; published: 14 Jun. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2916: 5161 (2011)
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51
A revision of the subgenus Parasphingonotus Benediktov & Husemann, 2009
(Orthoptera: Oedipodinae: Sphingonotini)
MARTIN HUSEMANN1,3, JESSE RAY1 & AXEL HOCHKIRCH2
1Baylor University, Department of Biology, Waco, Tx-76706, USA
2Trier University, Department of Biogeography, D-54286 Trier, Germany
3Corresponding author. E-mail: Martin_Husemann@Baylor.edu
Abstract
The recently erected subgenus Parasphingonotus is revised. The diagnostic characters are given and the species Sphin-
gonotus (Parasphingonotus) radioserratus comb. n., Sphingonotus (Parasphingonotus) femoralis comb. n. and Sphin-
gonotus (Parasphingonotus)turkanae comb. n. are assigned to the subgenus. Sphingonotus (Parasphingonotus) airensis
is synonymized with S. (P.) femoralis, making the latter species the new type species of Parasphingonotus. Keys to the
subgenera (Sphingonotus, Neosphingonotus, and Parasphingonotus) of the genus Sphingonotus and to the species of
Parasphingonotus are presented. All Parasphingonotus species are re-described and further information on the distribu-
tion of the species are given.
Key words: Sphingonotus, Sphingonotini, Africa, stridulatory organs, intercalary vein, radial vein
Introduction
The genus Sphingonotus Fieber, 1852 is a highly diverse and widely distributed group of band-winged grasshop-
pers, currently including ca. 140 species (Eades et al. 2011). The species differ in many characters (e.g. pronotum
shape, wing venation, wing shape and coloration), but the most intriguing trait is the diversity of their stridulatory
mechanisms (Hochkirch & Husemann 2008). While most Sphingonotus species possess a serrate intercalary vein
on the fore wing, which is the usual type of stridulatory apparatus in Oedipodinae (Oedipodinae type), there are
two apomorphic stridulatory mechanisms found in two well defined subgenera. One group (Neosphingonotus
Benediktov, 1998) has thickened cross-veinlets between medial vein and radius of the fore wing and the other sub-
genus (Parasphingonotus Benediktov & Husemann, 2009) has a serrated radius instead of a serrated intercalary
vein (Johnsen & Schmidt 1982; Johnsen 1985). Due to its high significance for species recognition, the stridulatory
apparatus has a great value for taxonomy and phylogeny in the genus Sphingonotus (Hochkirch & Husemann
2008).
Discussions about the taxonomic and systematic status of Sphingonotini with specialized stridulatory apparati
reach back to Bey-Bienko’s (1948) description of Sphingonotus paradoxus Bey-Bienko, 1948. This species shares
most of its general morphology with Sphingonotus savignyi Saussure, 1884, but differs in the structure of its strid-
ulatory apparatus (Johnsen 1985). While S. savignyi shares the serrate intercalary vein with most other Oedipodi-
nae, S. paradoxus has elevated cross veinlets between radius and media (Neosphingonotus type). After more than
half a century of repeated systematic reorganization (reviewed in Benediktov 1998, 2009), twelve species with this
stridulatory structure are meanwhile grouped in the subgenus Neosphingonotus (Benediktov 2009).
In contrast to the Neosphingonotus type, the third type of stridulatory apparatus has received relatively little
attention. Johnsen and Schmidt (1982) first noted that Sphingonotus turkanae Uvarov, 1938 possesses an epiproct
unique to the genus, as well as a smooth intercalary vein and a serrate radius. The same authors mentioned that this
trait is shared with a species from Tunisia, which was later described as Sphingonotus radioserratus Johnsen, 1985
(Johnsen 1985). Johnsen (1985) also added Sphingonotus airensis Chopard, 1950 to this species group without for-
HUSEMANN ET AL.
52 · Zootaxa 2916 © 2011 Magnolia Press
mally describing a new genus or subgenus. Benediktov (2009) erected the subgenus Parasphingonotus for Sphin-
gonotus species with a serrate radial vein and assigned S. airensis as type species.
Here, we revise the subgenus Parasphingonotus and provide a key to subgenera of Sphingonotus as well as to
the species within Parasphingonotus.
Material and methods
Morphological analyses. We studied the Sphingonotus material in the collections of the Natural History Museum
London (NHM), the Academy of Natural Sciences Philadelphia (ANSP), the Zöologisch Museum Amsterdam
(ZMA), the Muséum national d’histoire naturelle Paris (MNHN), the Lund Zoological Museum (LZM), the Natu-
ral History Museum Berlin (MfN). We also studied some specimens in the private collections of MH, AH and
Claudia Hemp (Bayreuth) as well as some specimens provided by Tim McNary (Fort Collins, CO). In total, we
examined the stridulatory mechanism of 123 species and subspecies of Sphingonotus. In addition we investigated
specimens of several related genera belonging to the tribe Sphingonotini (Thalpomena Saussure, 1884, Leptop-
ternis Saussure, 1884, Granada Koçak and Kemal, 2008, Trimerotropis Stål, 1873, Cophotylus Krauss, 1902,
Asphingoderus Bey-Bienko, 1950, Eusphingonotus Bey-Bienko, 1950, Eusphingoderus Bey-Bienko, 1950, Sphin-
goderus Bey-Bienko, 1950, Heliopteryx Uvarov, 1914, Helioscirtus Saussure, 1884, Vosseleriana Uvarov, 1924).
Biometric measurements were obtained using a digital calliper (range 0–150 mm). After microscopic inspection,
we examined the wing structures of selected species and specimens using Scanning Electron Microscopy (SEM).
We used a Zeiss DSM 962 (Oberkochem, Germany) at the University of Osnabrück and a Leo 1455 VP (Zeiss,
Oberkochem, Germany) at the NHM (London) to photograph the stridulatory structures. The Leo 1455 VP was
also used for examining type material as the specimens do not need to be dissected.
The epiproct of males and ovipositors of females were photographed using a Canon EOS 450D with a Canon
EF 100 mm macro lens. The genitalia of males of all available Parasphingonotus species were dissected. For this,
either whole specimens were softened in a refreshment box containing water and naphthalene or the abdomen of
dried specimens were soaked in water or in a 0.2 M NaOH solution. We then extracted the phallic complex (aedea-
gus and epiphallus) with a bent pin and forceps. Spare tissue was removed via incubation in 0.2 M NaOH and
removal with a forceps. The sclerotized structures were stored in 100% ethanol and then photographed using a ste-
reo-microscope with attached camera. The epiproct and epiphalli of all species were drawn. In addition to Paras-
phingonotus species, the epiprocts and epiphalli of S. S. caerulans from France and S. N. azurescens from Morocco
were drawn as reference for the other subgenera.
Results and discussion
The examination of the museum specimens revealed that four species possess a serrated radius and, therefore, have
to be assigned to the subgenus Parasphingonotus: S. airensis Chopard, 1950, S. femoralis Uvarov, 1933,
S. radioserratus Johnsen 1985, and S. turkanae Uvarov 1938. A preliminary phylogenetic analysis of mitochon-
drial and nuclear sequences revealed that the above mentioned species indeed form a monophyletic clade (Hoch-
kirch & Husemann unpublished data). The investigation of the type specimens of S. airensis and S. femoralis, as
well as the original species descriptions, revealed no distinctive differences between these two species. As all spec-
imens of S. airensis studied fell into the range of variation of S. femoralis, we here synonymize S. airensis with
S. femoralis.
The investigated species of the genera Thalpomena, Leptopternis, Jacobsiella, Trimerotropis, Cophotylus,
Asphingoderus, Eusphingonotus, Eusphingoderus, Sphingoderus, Heliopteryx, Helioscirtus and Vosseleriana all
possessed the Oedipodinae type or no stridulatory apparatus on the forewing. The supra-anal plates of Granada
imitans (Brunner von Wattenwyl, 1882) and a hitherto undescribed species from Spain are rather similar to the one
found in Parasphingonotus, but their wings possess the Oedipodinae stridulation type or Neosphingonotus type,
respectively. Only three East Asian species (S. tsilingensis Zheng, Tu and Liang, 1963, S. ningsianus Zheng and
Gow, 1981, S. mongolicus Saussure, 1888) have been included in our analysis so far, all of which belong to the sub-
genus Sphingonotus.
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REVISION OF THE SUBGENUS PARASPHINGONOTUS
Sphingonotus Parasphingonotus femoralis Uvarov, 1933
Synonymy: Sphingonotus airensis Chopard, 1950
Material examined. Type material: Holotype: 1 male, ARABIAN PENINSULA, South Arabian Desert, 1931, leg.
B. S. Thomas, det. B. Uvarov, Depository: NHM, Paratype 1 female, ARABIAN PENINSULA, South Arabian
Desert, 1931, leg. B. S. Thomas, det. B. Uvarov, Depository: NHM, Paratype, 1 female, SOMALIA, Behin, Jan.
1897, leg. F. Gillet, det. B. Uvarov, Depository: NHM.
Additional material examined. 1 male, OMAN, Kabil, 28 Oct. 1970, leg. J. Tunstall, Depository: NHM, 1
female, OMAN, Dhofar, Salalah, Jebel Qara, 23 Oct. 1983, leg. S. D. Gillett, Depository: NHM, 1 female, SAUDI
ARABIA, Nejran, 20 July 1962, leg. G. Popov, Depository: NHM, 1 male, SAUDI ARABIA, upper Wadi Alah-
siba, Tihma, 10 Feb. 1962, leg. G. Popov, Depository: NHM, 1 male, IRAN, Sarbaz, May 1950, leg. Salvatian,
Depository: NHM, 1 female, YEMEN, near Taizz, 8 Oct. 1962, leg. G. Popov, Depository: NHM, 1 male, YEMEN,
El’Asr, about 3 miles W of San’a, 14 Feb. 1938, leg. H. Scott & E. B. Britton, Depository: NHM, 1 male, YEMEN,
Beit Baus, about 5 miles S of San’a, leg. H. Scott & E. B. Britton, Depository: NHM,
Specimens formerly identified as S. airensis: 1 male, 1 , ERITREA, Agra Plain, 07 July 1950, leg. A. J.
Wood, Depository: NHM, 1 male, 1 female (former paratypes of S. airensis), NIGER, Air, Tésuar, 1947, leg. Chop.
Vill., Depository: NHM, 1 male (former Type of S. airensis), NIGER, Air, Tésuar, 1947, leg. Chop. Vill., Deposi-
tory: MNHN, 1 male, 1 female (former Types of S. airensis), NIGER, Air, Tassasset, 1947, leg. Chop. Vill., Depos-
itory: MNHN, 1 female (former Types of S. airensis), NIGER, Air, Tarrouadji, 1947, leg. Chop. Vill., Depository:
MNHN, 1 female, NIGER, Tabourax, 27 Oct. 1985, leg. T. McNary, Depository: Timothy J. McNary, private col-
lection.
Distribution. Records exist from Mauritania, Mali, Niger, Chad (former distribution of S. airensis (Chopard
1950, Dirsh 1965, Mestre & Chiffaud 2006)), Sudan, Western Egypt (Mistshenko 1936), Eritrea, Somalia (Uvarov
1933), Oman, Saudi Arabia, Yemen, Jordan, and Israel, to Iran and Pakistan (Popov 1980, Massa & Fontana 1998,
Ingrisch 1999)
Male. General facies: Habitus as typical for the genus, but fairly slender; medium size: length from fastigium
of vertex to end of fore wings 19.6–21.6 mm (S. femoralis 19.6–21.6 mm (N=5), formerly identified as S. airensis
19.6–21.1 mm (N=3)).
Color: Body coloration variable: sandy to medium brown, sometimes with red tint. Antennae alternating light
and dark brown. Outer area of hind femora same color as body, but with dark incomplete fascia 1/3 of the length of
the femora away from the knee. Inner side of hind femora beige with dark knee and one full fascia at the level of
the outer band, a second fascia is of variable width and can fill the whole medial area or can be absent. Hind tibiae
dirty whitish or pale yellowish with dark spines.
Head: Antennae filiform, longer than head and pronotum together. Frontal ridge slightly concave with lateral
carinae slightly widened between basis of antennae. Fastigium of vertex concave, lateral carinae elevated, median
carinula variable, often slightly developed. Temporal foveolae diffuse, elongate triangular.
Thorax: Pronotum almost as wide as long; pronotal disk smooth with three complete transverse sulci; median
carina present in front of first sulcus, often two small elongated tubercles (often of whitish color) between second
and third sulcus; posterior margin rectangular to obtuse angular or rounded; lateral carinae slightly developed in
metazona. Metazona approximately twice as long as prozona (0.37–0.56) (S. femoralis 0.41–0.56 (N=5), formerly
identified as S. airensis 0.37–0.55 (N=3)).
Wings: Hind wings slightly bluish or hyaline without any trace of a fascia but with darkened veins. Fore wings
moderately densely reticulated, relatively narrow: 6.2–7.1 (S. femoralis 6.4–7.1 (N=5), S. airensis 6.2–7.0 (N=3))
as long as wide. Intercalary vein slightly sinuous or curved with the centre being most distant from the radius,
sometimes proximally splitting up into two veins smooth; radius serrated.
Femora: Hind femora 3.2–3.8 (S. femoralis 3.2–3.8 (N=5), S. airensis 3.3–3.5 (N=3)) times longer than wide;
spurs of tibia of normal length for the genus; arolium small (1/3 to 1/2 of length of claws).
Abdomen: Tympanum typical for the genus, as high as long, less than 50% of opening covered by ventral lobe.
Epiproct elongate and trilobate, almost reaches the tip of the cerci, lateral margins slightly elevated, basally with
small tubercles and two triangular and two linear ridges. Cerci slightly longer than epiproct, slightly conical with
blunt rounded tip.
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54 · Zootaxa 2916 © 2011 Magnolia Press
FIGURE 1. Lateral view of male of a) S. P. femoralis, b) S. P. radioserratus, and c) S. P. turkanae.
Inner genitalia: Lophi of epiphallus large and bipartite, internal lobes projecting inside and rounded, external
part broad oval; ancorae in- and down-curved, pointed; epiphallic bridge of medium width; anterior projections
short, spoon-shape, but pointed; posterior projections short, rounded.
Female. The females are mostly similar to males but differ in the following characters:
Body more stout; body size: 27.1–30.7 mm (S. femoralis 28.4–30.7 mm (N=4), S. airensis 27.1–29.1 mm
(N=4)); cerci conical with broad basis and pointed tip; ovipositor variable, short; valves strongly curved, with
rounded or pointed tips; both valves approximately of similar length; lower valve with distinct rounded tooth;
lower valve from seen below shows considerable variation (Fig. 5). Although the appearance of the subgenital
Zootaxa 2916 © 2011 Magnolia Press · 55
REVISION OF THE SUBGENUS PARASPHINGONOTUS
plates of the paratypes of S. femoralis from Somalia and S. airensis from Niger is slightly different, these differ-
ences are in the range of variation found among studied individuals within each species. Lower valve from seen
below with basal plates of oval shape and variable size covering about half of the valve. Proximal part of valves of
medium width, laterally elevated. Subgenital plate with two rounded lobes.
FIGURE 2. Fore and hind wings of a) S. P. femoralis, b) S. P. radioserratus, and c) S. P. turkanae.
Sphingonotus Parasphingonotus radioserratus Johnsen, 1985
Material examined. Type material: Holotype: 1 male, TUNISIA, Gafsa, 01 Aug. 1951, leg. O. Persson, det. P.
Johnson, Depository: ZML, Allotype: 1 female, TUNISIA, Gafsa, 01 Aug. 1951, leg. O. Persson, det. P. John-
son, Depository: ZML, Paratypes: 2 females, 1 male, TUNISIA, Gafsa, 01 Aug. 1951, leg. O. Persson, det. P. John-
son, Depository: ZML
Additional material examined. MOROCCO: 2 males, 1 , Tarda, 10 km W of Errachidia (31.48 N, 4.36
W), 20 May 2008, leg. M. Husemann, Depository: NHM, 2 males, 2 females, Tarda, 10 km W of Errachidia (31.48
N, 4.36 W), 20 May 2008, leg. M. Husemann, Depository: M. Husemann, private collection.
Distribution. Tunisia (Johnsen 1985), Morocco.
Male. General facies: Habitat as typical for Sphingonotus, but relatively slender; medium size: length from
vertex to end of fore wings 23–24.5 mm (N=7).
Color: Color pattern fairly distinct: body light brownish to sand-colored, abdomen sometimes whitish; head of
the same color with slightly darker eyes; antennae alternating dark and light brown; pronotum variable: similar
color as body or with dark and light x-shaped markings. Outer median area of hind femora light brown; upper
carina often with dark spot about 1/3 of the length of the hind femora away from the knee; inner side of hind fem-
ora light brown with dark knee and two dark spots, one at the position of the dark spot, one at the beginning of the
second third (Fig. 7). Hind tibiae whitish to slightly yellowish with dark spines.
Head: Antennae filiform, longer than head and pronotum together. Frontal ridge slightly concave with lateral
carinae, ridges almost parallel, fastigium of vertex concave, lateral carinulae slightly elevated, median carinula
missing. Fastigial foveolae small, elongate triangular.
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56 · Zootaxa 2916 © 2011 Magnolia Press
FIGURE 3. Supra-anal plates of a) S. P. femoralis, b) S. P. radioserratus, and c) S. P. turkanae; d) S. N. azurescens and e) S. S.
caerulans.
Thorax: Pronotum almost as wide as long; pronotal disk smooth with 3 complete sulci, median carina slightly
elevated in front of first sulcus. Hind margin rectangular to obtuse angular or rounded; lateral carinae slightly
developed in metazona; metazona approximately twice as long as prozona (0.39–0.53, N=7).
Wings: Fore wings with two incomplete dark bands, both starting at the front margin and reaching up to inter-
calary vein, moderately densely reticulated, relatively narrow: 6.7–7.2 (N=7) as long as wide. Intercalary vein
almost straight and parallel to the radius, smooth; radius serrated. Hind wings slightly bluish to hyaline with central
narrow dark band which does not reach the posterior and interior margins.
Femora: Hind femora 3.5–4.7 (N=7) times longer than wide; spurs of tibia of normal length for the genus; aro-
lium small (1/3 to 1/2 of length of claws).
FIGURE 4. Epiphalli from a) S. P. femoralis, b) S. P. radioserratus, and c) S. P. turkanae; d) S. N. azurescens and e) S. S. caer-
ulans.
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REVISION OF THE SUBGENUS PARASPHINGONOTUS
FIGURE 5. View of the lower valve of the ovipositor and the subgenital plate from below: a) a former paratype of S.
airensis from Niger, b) a paratype from S. P. femoralis from Somalia, c) an allotype from S. P. radioserratus from Tunisia, and
d) a paratype of S. P. turkanae from Ethiopia.
Abdomen: Tympanum typical for the genus, as high as long, less than 50% of opening covered by ventral lobe.
Epiproct elongate and trilobate, almost reaches the tip of the cerci, lateral margins elevated, basally with several
small tubercles, apically with horseshoe-shaped ridge. Cerci slightly longer than epiproct, slightly conical with
blunt rounded tip.
Inner genitalia: Epiphallic lophi large and bipartite, internal part bell-shaped, external part oval; ancorae in-
and down-curved, pointed; epiphallic bridge of medium width; anterior projections short, spoon-shaped; posterior
projections short, pointed.
Female. The females are mostly similar to males but differ in the following characters:
Body more stout; body size: 28.4–31.3 mm (N=5); antennae only slightly longer than head and pronotum
together; cerci conical with pointed tip. Ovipositor displays some variation; relatively short, valves strongly
curved, with rounded or pointed tips, both valves approximately of similar length, lower valve with distinct
rounded tooth. Lower valve from seen below with basal plates of oval shape, internal site proximally with little
rounded appendix (Fig. 5), the plate covers slightly more than half of the valve; proximal part of valves of medium
width, laterally elevated. Subgenital plate with two lobes of almost rectangular shape.
Sphingonotus Parasphingonotus turkanae Uvarov, 1938
Material examined. Type material: Holotype: 1 male, KENYA, Turkana, Komogin R., 05 Mar. 1934, leg. D. R.
Buxton, det. B. Uvarov, Depository: NHM, Paratype: 1 male, KENYA, Lokitang, Turkana Nord, 1932-33, leg. C.
Arambourg, P.-A. Chappuis & R. Jeannel, det. B. Uvarov, Depository: NHM, Paratype: 1 female, ETHIOPIA,
Bourié, Shore of river Omo, 1932-33, leg. C. Arambourg, P.-A. Chappuis & R. Jeannel, det. B. Uvarov, Deposi-
tory: NHM.
Additional material examined. 1 female, ETHIOPIA, El Oha, 10 Nov. 1968, leg. J. Tunstall, Depository:
NHM, 1 male, ETHIOPIA, Diré-Daoua, Dec. 1934, leg. H. Uhlenhuth, Depository: NHM, 1 male, 1 female,
YEMEN, Socotra, R. A. F. Camp, 26–27 Jan. 1953, leg. G. Popov, Depository: NHM , 1 female, YEMEN, San’a,
10–15 Oct. 1937, leg. Dr. C. Rathjens, Depository: NHM, 1 male 1 female, SOMALIA, near Borama, 1934, leg.
Capt. R. H. T. Taylor, Depository: NHM, 1 male, 1 female, TANZANIA, Tanganyika Terrain, Mkomasi Station, 60
miles W of Amani, 20 June 1934, leg. E. Burtt, Depository: NHM, 1 male, TANZANIA, Mt. Meru, 26 May 2005,
leg. C. Hemp, Depository: M. Husemann, private collection.
Distribution. Ethiopia, Kenya, Tanzania, Somalia (Uvarov 1938, Dirsh 1965, Hemp 2009), Socotra, Yemen
(Uvarov & Popov 1957, Johnsen & Schmidt 1982)
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FIGURE 6. a) Fore wing of S. P. femoralis, SEM photographies of the stridulatory apparati of b) S. P. turkanae, c) S. P. turka-
nae in detail, d) S. S. caerulans, and e) S. N. azurescens.
Male. General facies: Habitus as typical for the genus, but fairly slender; medium size: length from fastigium of
vertex to end of fore wings 20.7–23.3 mm (N=9).
Color: Body coloration variable: light to medium brown. Antennae alternating light and dark brown. Outer area
of hind femora same color as body, but with dark incomplete fascia 1/3 of the length of the femora away from the
knee. Sometimes with smoky spot in the middle of the medial area. Inner side of hind femora beige with dark knee
and one full fascia at the level of the outer band, in addition the whole medial area is dark. Hind tibiae dirty whitish
or pale yellow with dark spines.
Head: Antennae filiform, longer than head and pronotum together. Frontal ridge slightly concave with lateral
carinae, widened between basis of antennae. Fastigium of vertex concave, lateral carinae almost parallel, elevated,
medial carinula variable, usually distinctly developed, often Y-shaped. Temporal foveolae distinct, elongate trian-
gular.
Thorax: Pronotum almost as wide as long; pronotal disk strongly rugose with elevated tubercles and ridges, 2
complete transverse sulci, the area between the sulci consists of two elevated knobs which form a small plateau and
interrupt the third (median) sulcus; median carina distinct in front of first sulcus; posterior margin right angled and
weakly rounded at the tip; lateral carinae developed in metazona. Metazona 2–3 times as long as prozona (0.35–
0.6, N=9).
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REVISION OF THE SUBGENUS PARASPHINGONOTUS
FIGURE 7. Inner side of hind femora of a) S. femoralis, b) S. radioserratus, c) S. turkanae.
Wings: Fore wings moderately densely reticulated, relatively narrow: 5.9–7.4 (N=9) as long as wide. Interca-
lary vein often almost straight, sometimes curved, then the central part is most distant from the radius, smooth;
radius serrated. Hind wings slightly bluish without any trace of fascia but with dark veins.
Femora: Hind femora 3.5–3.9 (N=8) times longer than wide; spurs of tibia of normal length for the genus; aro-
lium small (1/3 of length of claws).
Abdomen: Tympanum typical for the genus, as high as long, less than 50% of opening covered by ventral lobe.
Epiproct short and trilobate, about 2/3 the length of the cerci, lateral margins slightly elevated, basally with several
small tubercles. Cerci longer than epiproct, slightly conical with blunt rounded tip.
Inner genitalia: Lophi of epiphallus large and bipartite, internal lobes mushroom shaped, projecting inside,
external part oval; ancorae in- and down-curved, pointed; epiphallic bridge of medium width; anterior projections
short and narrow; posterior projections short, rounded.
Female. The females are mostly similar to males but differ in the following characters:
Body more stout; body size: 25–29.5 mm (N=7); cerci almost cylindrical with rounded tip. Ovipositor displays
some variation; short, valves strongly curved, with rounded or pointed tips; both valves approximately of similar
length, lower valve with distinct rounded tooth. Lower valve from seen below with basal plates of almost round
shape (Fig. 5), covering less than half of the valve. Proximal part of valves relatively narrow, laterally elevated.
Subgenital plate without distinct lobes, lobes replaced by two broad curvatures.
Differential diagnosis. S. radioserratus is easily distinguished from the other two species of the subgenus by its
two incomplete wing bands on the forewing and by the dark band on the hind wing. The males have a very distinc-
tive supra-anal plate, which is longer than in the other species of the subgenus and has a horseshoe-shaped ridge in
the apical part. S. turkanae and S. femoralis are easily distinguished by the sculpting patterns of the pronotum,
which is much more rugose and has a higher and more distinctive medial carina in S. turkanae. The upper carina of
the hind femora has a distinct step in S. turkanae (like in Oedipoda species). The fastigium of vertex and frontal
ridge of S. turkanae are much wider than in S. femoralis and have a more distinct carina which can be bifurcated in
some individuals. Females of all species differ in the shape of the basal lobes of the lower valve of the ovipositor. S.
femoralis have oval basal lobes, while S. radioserratus has an internal small appendix proximal on the basal lobe.
S. turkanae has very short basal lobes which are of almost round shape. The female subgenital plates of all species
differ as well. S. femoralis has two rounded lobes, while those are almost rectangular in S. radioserratus and only
represented by a broad curvature in S. turkanae.
HUSEMANN ET AL.
60 · Zootaxa 2916 © 2011 Magnolia Press
FIGURE 8. Distribution of Parasphingonotus species (grey and black coloration indicate country records derived from the lit-
erature, symbols indicate locations of examined material).
Key to the subgenera
1 Intercalary vein in the medial area of the fore wings serrate, in females sometimes only slightly serrate or smooth (Fig. 6d). . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sphingonotus s.s.
- Intercalary vein not serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2 Thickened cross veinlets between radial and medial vein of the fore wings present (Fig. 6e), radial vein without serration,
supra-anal plate triangular or rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Neosphingonotus
- Without thickened cross veinlets between radial and medial vein, radial vein stronger raised than subcostal vein and serrate
(Fig. 6 c, d), supra-anal plate trilobite (Fig. 3a–c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parasphingonotus
Key to the species of the subgenus Parasphingonotus
1 Hind wings with a dark fascia (Fig. 2b), epiproct elongate, as long as cerci, sides elevated giving it a spoon like appearance
(Fig. 3b), with horse shoe shaped ridge in the apical third (North West Africa) . . . . . . . . . . . . . . . . . . . . . . . S. P. radioserratus
- Hind wings without distinct dark fascia, sometimes with blackish veins, epiproct as long as or shorter than cerci, sides not
raised, no horse shoe shaped tubercle in apical third . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2 Pronotum strongly rugose with raised ridges and tubercles, hind margin rectangular; inner sides of hind femora yellow with
dark knee, distinct dark fascia in apical third and dark medial area (Fig. 7c), epiproct in males much shorter than cerci (less
than 2/3), strongly trilobate with variable numbers of raised tubercles (Fig. 3c) (East Africa, Socotra, Yemen) . .S. P. turkanae
- Pronotum smooth, hind margin right or obtuse angled and rounded; upper carina of hind femora continuously lowered, without
step; inner sides of hind femora yellow or dirty-yellowish with a distinct dark fascia in the apical third (Fig. 7a), medial area
variable, dark spot of variable size can almost fill the whole medial area or can be absent, epiproct more elongate (longer than
2/3 of cerci), trilobate, with two triangular lobes at the basis (Fig. 3a) (Mauritania, Niger, Chad, Northeast and East Africa,
Arabian Peninsula, Iran) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. P. femoralis
Zootaxa 2916 © 2011 Magnolia Press · 61
REVISION OF THE SUBGENUS PARASPHINGONOTUS
Acknowledgement.
We want to thank Alexander Benediktov, David Llucia-Pomares and Bernard Defaut for valuable comments on
Sphingonotus and fruitful discussions. We are grateful to Judith Marshall and George Beccaloni (NHM), Roy Dan-
ielsson (ZML), Christiane Amedegnato (MNHN), Willem Hogenes (ZMA), Michael Ohl (MfN), Daniel Otte and
Jason Weintraub (ANSP) for giving access to museum collections and providing specimens for our study. For addi-
tional material we want to thank Claudia Hemp and Timothy McNary. Parts of the project have been financed by
grants from the Orthopterist’s Society and by the Synthesys project to MH.
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... The wide distribution, the high rates of endemism in some areas, the convergence with North American genera and especially its polymorphic stridulation organs make Sphingonotus an interesting study system for biogeographic and evolutionary questions (e.g. Johnsen 1985;Benediktov 2009;Husemann et al. 2011Husemann et al. , 2012Husemann et al. , 2013Husemann et al. , 2014Husemann et al. , 2015. However, because of the high number of species, the wide distribution, and the often cryptic morphological differences, we still have only a limited understanding of the systematics and local and global species diversity. ...
... Both species are morphologically strikingly similar (Bey-Bienko 1950, Bey-Bienko andMistshenko 1951;Benediktov, 2009) and differ only in a single trait, the structure of their stridulatory apparatus (Johnson 1985). Sphingonotus (Neosphingonotus) paradoxus shares its stridulatory apparatus (elevated cross veinlets between radius and media) with all other species of the subgenus Sphingonotus (Neosphingonotus), whereas Sphingonotus (Sphingonotus) savignyi thus far was known as a species with a serrate intercalary vein (Husemann 2011;Benediktov 1998Benediktov , 2009), like all other species of the subgenus Sphingonotus (Sphingonotus). While this theoretically clearly separates both species the story is more complex. ...
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Contribution a l'etude de l'Air, Orthopteroides. Memoires de l'Institut Francais D'Afrique Noire
  • L Chopard
Chopard, L. (1950) Contribution a l'etude de l'Air, Orthopteroides. Memoires de l'Institut Francais D'Afrique Noire, 10, 127– 145.
Revision of palearctic species of the genus Sphingonotus Fieb. (Orthoptera, Acrididae)
  • L L Mistshenko
Mistshenko, L.L. (1936) Revision of palearctic species of the genus Sphingonotus Fieb. (Orthoptera, Acrididae). Eos, 12, 65– 282.
To the taxonomy and bioacoustics of grasshoppers of the genus Sphingonotus Fieber
  • A A Benediktov
Benediktov, A.A. (2009) To the taxonomy and bioacoustics of grasshoppers of the genus Sphingonotus Fieber, 1852 (Orthoptera, Acrididae, Oedipodinae). Proceedings of the Russian Entomological Society. St. Petersburg, 80, 21–33 [In Russian].