Research Article |
Corresponding author: Sigfrid Ingrisch ( s.ingrisch@leibniz-zfmk.de ) Academic editor: Martin Husemann
© 2020 Sigfrid Ingrisch.
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Citation:
Ingrisch S (2020) New subgenera and species of Agraeciini (Orthoptera, Tettigoniidae, Conocephalinae) from South Asia found in historical insect collections. Evolutionary Systematics 4(2): 119-132. https://doi.org/10.3897/evolsyst.4.60525
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Based on specimens collected in historical times and stored in the insect collections of the Natural History Museum London and the Zoological Museum Hamburg two new subgenera are established Robustacca subgen. nov. of the genus Peracca Griffini, 1897 and Indoliara subgen. nov. of the genus Liara Redtenbacher, 1891. Five new species are described: Peracca (Robustacca) nigrifrons sp. nov. from Sumatra, Liara (Acanthocoryphus) durata sp. nov. from Hongkong, Liara (Indoliara) dividata sp. nov. from South India, Pseudosubria assamensis sp. nov. from Assam and Depressacca kinabalu sp. nov. from Sabah. The species Odontoconus robustus Karny, 1923 is newly combined as Peracca (Robustacca) robustus (Karny, 1923) comb. nov. An extended and updated key to the subgenera of the genus Peracca Griffini, 1897 is provided. All specimens are deposited in the original collections.
Biodiversity, BMNH, ZMH, Hongkong, India, Malaysia, Sabah, Singapore, Sumatra
As our knowledge of the insect diversity in number of species increases and the knowledge of their morphology, diversity, distribution and ecology becomes more detailed than in historical times, we may come to different solutions for the arrangement of the lower taxa as species or subspecies into subgenera, genera and higher taxonomic units. A recent example in the tribe Agraeciini is the re-grouping of taxa related to Peracca Griffini, 1897, Liara Redtenbacher, 1891 or Anelytra Redtenbacher, 1891 into genera and subgenera by
The discovery of new species in formerly unworked, historical insect collections kept in museums inspired me to refine the subdivision of Peracca into subgenera proposed by
The current study is based on formerly unidentified specimens that were found in the insect collections of the Natural History Museum London (BMNH) and the Zoological Museum Hamburg (ZMH). Their data had not been published elsewhere. Data of one specimen kept in the Bishop Museum Honolulu (BMH) and published already in
Photographs were done some years ago with a Nikon E4500 (habitus) or the same camera mounted to a Motic M5 stereo microscope. Some of the habitus photos were taken with classical photography. Line drawings were done using a camera lucida mounted to a Wild stereo microscope.
The ID numbers assigned to the specimens listed in this publication refer to the specimen IDs used in my database. These numbers are also printed at the identification labels attached to the specimens. In addition, for specimens in BMNH the serial acquisition number is given. This number is however not unique for a single specimen but refers to the collection event and donator. It consists of the year of acquisition and a serial number. For the specimen in ZMH the collection ID of the specimen is provided.
Geographical coordinates were not available from information attached to the specimens. For most of the specimens studied I found rough estimates for the coordinates of their localities from a search in
The specimens on which the current study is based are housed in the following museums:
BMH Bishop Museum Honolulu, USA
BMNH Natural History Museum London, UK
ZMH Zoological Museum Hamburg, part of the Centrum für Naturkunde, University of Hamburg, Germany
Peracca Griffini, 1897: 1; type species: Peracca conspicuithorax Griffini, 1897 from Malaysia, Perak, Malacca
Species of the genus Peracca Griffini, 1897 occur in Malay Peninsula, Singapore, Borneo, Sumatra, Java, and some smaller islands of that area. Seven species of Peracca have been known during my revision of the SE Asian Agraeciini (
After Odontoconus had been downgraded to a subgenus of Peracca by
(extended and modified from
1 | Face oval. Pronotum in both sexes with lateral lobes rather deep, less than twice as long as high. Male subgenital plate with central area prolonged behind, with narrow apical third, at end upcurved and divided in midline, terminating into a pair of acute apical cones, without moveable styli (Fig. |
subgenus Robustacca subgen. nov. |
– | Face conical, mostly narrow conical. Pronotum in both sexes with lateral lobes low, narrow, more than twice as long as high. Male subgenital plate not prolonged, with apical margin broadly truncate or slightly concave, provided with moveable styli | 2 |
2 | Male pronotum with elongate hind lobe covering only base of tegminal stridulatory apparatus; prosternum with very long spines; anterior part of mesosternum without distinct spines or spine-like projections. Male cerci with almost rounded, not hooked, large proximal lobe directed more or less laterally | subgenus Sumatracca Gorochov, 2011 |
– | Male pronotum of different shape, either longer or shorter; prosternum spines vary between species; anterior part of mesosternum with distinct spines or spine-like projections. Male cerci with a large proximal lobe or two such lobes, and this lobe or one of these lobes more or less hooked and directed medially | 3 |
3 | Fastigium verticis with a short dorsal tubercle or without any. Male pronotum prolonged behind, covering a great part or all of the stridulatory area; prosternum with moderately long or short spines | subgenus Peracca Griffini, 1897 |
– | Fastigium verticis with a rather long dorsal tubercle. Male pronotum less prolonged behind, leaving the stridulatory area largely free, in female with truncate hind margin; prosternum with long spines | subgenus Odontoconus Fritze in Carl, 1908 |
Odontoconus robustus Karny, 1923.
The name of the new subgenus is built from a combination of the root of the name of the type species with the ending of the genus name.
The stout general habitus of the new subgenus and the rather high lateral lobes of pronotum resemble the situation in the genus Liara, but the spine behind the ventro-anterior angle of pronotum, the strongly modified male cerci and the absence of sclerotized titillators agree with the genus Peracca. The new subgenus contains stout, robust species with large head. – Head and pronotum rugose. Fastigium verticis with a dorsal process. Pronotum with lateral lobes rather deep compared to other subgenera of Peracca, less than twice as long as high; at ventral margin with a small spine behind anterior angle. Fully winged, wings surpassing abdomen but not reaching tip of ovipositor in female. Tegmen markedly narrowed in apical third. Prosternal spines long with acute tip. The male cerci have the internal branch prolonged and curved backward at end, the apical process is laterally compressed and pointing straight behind instead of being downcurved as in the subgenus Odontoconus, it is provided with spinules along dorsal margin but is missing a narrow ventral branch that occurs in most species of the subgenus Odontoconus. A so far unique character is the markedly narrowed prolongation of the male subgenital plate that is divided and upcurved at end and has the styli reduced to small and short, acute conical spines while in all other subgenera of Peracca s.l. the males have subgenital plates with truncate or slightly concave apical margin and with distinct, normal styli.
Peracca (Robustacca) robustus (Karny, 1923) comb. nov. and Peracca (Robustacca) nigrifrons sp. nov.
Odontoconus robustus
The Latin word robustus does not accept different gender forms.
1 female (holotype): Singapore: Bukit Timah (1°20'N, 103°47'E), 23.viii.1911 – depository: BMNH (B.M.1937–197); 1 male, Malaysia, Selangor, Ampang (3°10'N, 101°46'E), 1.iii.1948, coll. H.T. Pagden – depository: BMNH (B.M.1955–354); 1 male, Malaysia, Rompin Mining Co. (Rompin 2°48'N, 103°27'E), Railway Track, 50 km, 31.iii.1961, coll. K.J. Kuncheria – depository: BMH Honolulu.
The species was originally described from a single female from Singapore by
Head with fastigium verticis markedly projecting anteriorly, on top in basal area with a roughly cylindrical process that has the apical area narrowed and rounded (Fig.
Peracca (Robustacca) robustus (Karny, 1923) comb. nov. male from Ampang (A–C, E–J), male from SE Pahang (D), and female holotype (K): A–C male subgenital plate and cerci in ventral (A), dorsal (B) and lateral view (C); D basal area of male left cercus in medio-dorsal view; E left tegmen, F stridulatory file on underside of left tegmen; G, H stridulatory area of left (G) and right tegmen (H) in dorsal view; I habitus lateral view; J, K ventral view of body and femora of male (J) and female (K).
Male. Stridulatory file on underside of left tegmen with about 175 teeth; teeth in basal quarter narrow but distinctly spaced, teeth getting wider and more densely packed to about midlength, then becoming very dense and in about apical eighth strongly narrowed (Fig.
Peracca (Robustacca) subgen. nov.: A–C P. (R.) nigrifrons sp. nov. female holotype: A face; B fastigium verticis in ventro-anterior view and C in lateral view. – D–I P. (R.) robustus (Karny, 1923) comb. nov. female holotype (D–H) and male from SE Pahang (I) D Face; E fastigium verticis in lateral view; F female subgenital plate; G habitus dorsal view; H habitus lateral view; I fastigium verticis. Abbreviations: lo lateral ocellus, mo medium ocellus.
Female. The seventh sternite had been cut for filling the specimen. Subgenital plate with converging and upcurved lateral margins; apical margin in ventral view broadly subtruncate with upcurved margin that is slightly excised in middle (Fig.
Measurements (1 male, 1 female). Body w/wings: male 42, female 48; body w/o wings: male 33, female 37; pronotum: male 10.2, female 10.5; tegmen: male 28, female 32; hind femur: male 18, female 21; ovipositor: female 21.5 mm.
(female). Indonesia: Sumatra, Riau, Indragiri, Somgei Lalah [Sungai Lala, about 0°24'S, 102°12'E] 1.i.1900–26.viii.1901, leg. W. Burchard – depository: ZMH (HTodosumF01; collection ID: ZMH64336).
The name of the new species refers to its black face; from Latin niger, nigro black and frons face.
The new species differs from the type species, P. (R.) robustus by the black color of the face instead of agreeing with general coloration (Fig.
Peracca (Robustacca) nigrifrons subgen. & sp. nov. female holotype. A Head in dorsal view; B–D habitus in lateral (B), dorsal (C) and ventral view (D); E female abdominal apex in ventral view, the subgenital plate has the anterior area hidden under the 7th sternite in the shrunken specimen; F apical tergites and cerci; G, H pronotum in dorsal (G) and lateral view (H); I thoracic sternites; J ovipositor. Abbreviations: 7th seventh abdominal sternite, m membranes, ov base of ovipositor, sg apical lobes of subgenital plate.
Fastigium verticis subcylindrical, slightly surpassing scapus, apical area conical, apex obtuse; dorsal surface with a large cone at base; lateral ocelli projecting; in frontal view roundly swollen at base (Fig.
Male. Unknown.
Female. Tenth abdominal tergite transverse; angularly excised from apex for almost half of its length (Fig.
Light brown [discolored due to previous conservation in alcohol]. Frons black; mouthparts yellowish to reddish brown; apex of mandibles black. Tegmen with medium brown cells and yellow brown veins and veinlets (pattern little distinct); areas around subcosta and radius and before anal vein of uniform light color; in dorsal area some scattered cells darker than most others.
Body w/o wings: 33; pronotum: 9.8; tegmen: 30; hind femur: 19; ovipositor: 19.5 mm.
1 male, Indonesia: Sumatra, Sungei Kumbang, Korinchi (1°48'S, 101°16'E), 1370 m, 1.–30.iv.1914 – depository: BMNH (Lo042S001). Second record for this species.
Liara Redtenbacher, 1891: 444; type species Liara rufescens Redtenbacher, 1891: 444.
The genus contained so far three subgenera: Liara, Acanthocoryphus Karny, 1907, and Unalianus Koçak & Kemal, 2009 (replacement name for Oxystethus Redtenbacher, 1891 because of homonymy) (
(male). Hongkong [without precision], in or before 1911, coll. F.W. Terry – depository: BMNH (Lo031S001; B.M.1911–359). The type is unique.
The new species is named for the sclerotised central convexity of the male titillators, modified from Latin durare to harden, hardened.
The titillators of the new species are intermediate between those of the subgenera Liara s.str. and Unalianus Koçak & Kemal, 2009 in that they are fused as in Unalianus but of general shape as in Liara; the membranous lateral expansions are sclerotised but the apical areas not as strongly prolonged as in Unalianus. Judging from the shapes of the cerci and titillators, the new species comes close to Liara (Acanthocoryphus) brevis Ingrisch, 1998. It differs by the long and upcurved fastigium verticis which is almost twice as long as the scapus instead of only little longer and has a strong tubercle at base. Further differences are the fused titillators, the very small styli, and shorter wings. The cerci are similar in both species but in the new species less wide at base and the apical branches more curved.
A short and stout, strongly brachypterous species (Fig.
Liara (Acanthocoryphus) durata sp. nov. male holotype: A habitus lateral view; B face, C head dorsal view; D fastigium verticis lateral view; E stridulatory file on underside of left tegmen; F thoracic sternites, the arrow points at one of the prosternal spines; G–I titillators of male phallus in ventro-apical view (G), do. as seen from base (H), do. in oblique lateral view (I); J, K male cercus in external (J) and internal (K) views.
Male. Tenth abdominal tergite little prolonged; apical margin converging on both sides, broadly truncate in middle. Epiproct triangularly rounded with shallow medial furrow. Paraprocts with short obtuse compressed projections. Cerci short, at apex divided into a broad falcate dorsal and a narrow falcate ventral projection, both curved up; on internal side with a weak furrow from dividing point of apical projections to base (Fig.
Uniformely yellowish brown. Frons reddish brown; mandibles black; tip of fastigium verticis especially on ventral side black. Pronotum with indication of brown lateral bands near hind margin only. Tegmen with dark spots. Meso- and metasternal lobes dark brown with tips of lobes yellowish brown.
Body w/o wings: 19; pronotum: 7.6 ; tegmen: 8.2, hind femur: 14 mm.
Liara (Indoliara) dividata Ingrisch sp. nov.
The new subgenus resembles the genus Liara sensu stricto in general shape, basic coloration, and the shape of the long, substraight and laterally compressed female ovipositor that is dorso-ventrally widened in about mid-length. It differs by the strongly modified shape of the male cerci that are shorter and hood-shaped instead of laterally compressed and the ventral branch is rather thin, rounded, and inserted at base of the internal side of the main branch, it is angled about in mid-length and reaching only about the middle of the main branch instead of surpassing it in the nominate subgenus. Moreover, a small spine that is found at tip of the main cercus branch in some species of the nominate subgenus moved to about mid-length of the internal side of the main branch in the new subgenus and species. The male titillators are simplified in Indoliara and consist of long but simple, elongate bases that are bent lateral at tip terminating into a long-narrow oval, granular endplate while in Liara s. str. there are often lateral expansions and the tip of the titillators is serrate.
The tip of the fastigium verticis is markedly divided at end into two short lobes; but this might be a species-specific character since a very fine, inconspicuous notch at tip of the fastigium verticis can rarely be also found in species of Liara s. str., e.g. in L. alata Ingrisch, 1998)
(male). India: South India, Thekkadi, Periyar Dam (9°35'N, 77°9'E), 6.–10.v.1937, coll. B.M.-C.M. Expedition to South India – depository: BMNH (Lo032S001; B.M.1939–205).
(1 female). India: South India, Travancore, Pirmed (9°40'N, 76°59'E), elev. 1036 m, 4.–6.v.1933, coll. B.M.-C.M. Expedition to South India – BMNH (Lo032S002; B.M.1939–205).
The name of the new species refers to the shape of the fastigium verticis that is divided into two short apical cones; from Latin dividere to divide.
This is the so far only species in the subgenus. Its differences to the species of the other subgenera of Liara are discussed above under Indoliara subgen. nov.
Fastigium verticis longer than scapus, with a minute tubercle at top; apex bifurcate and slightly S-shaped, separated from fastigium frontis by a shallow furrow, but connected in middle by a lamella (Fig.
Liara (Indoliara) dividata sp. nov. A–G male holotype: A habitus dorsal view; B face; C stridulatory file; D abdominal apex with cerci in ventro-apical view; E titillators of male phallus; F tip of titillators; G subgenital plate. – H–K female: H habitus lateral view; I head in dorsal view; J thoracic sternites; K subgenital plate.
Male. Tenth abdominal tergite with short triangular projections above bases of styli; apical margin broadly concave in between. Epiproct wide with rounded lateral lobes [apex hidden]. Paraprocts with compressed, conical projections. Cerci conical, compressed, apex rounded; dorso-external surface concave; ventro-internal surface convex in apical half, with a broad, oblique, transverse furrow in basal half; with an acute spinule behind furrow and a thin process at very base; process conical at base, rectangularly bent dorsad in about middle, strongly compressed thereafter, apex with acute tooth on proximal side (Fig.
Female. Tenth abdominal tergite with apical margin concave in middle. Epiproct triangular, with shallow dorsal groove. Cerci conical, apex slightly curved mediad. Subgenital plate wider than long, roughly triangular with apex roundly excised. Ovipositor compressed blade-shaped, highest in middle.
Uniformly light brown (ochre). Frons reddish brown in male; labrum yellow, mandibles black. Tegmen speckled with small brown spots (Fig.
Body w/wings: male 36, female 42; body w/o wings: male 36, female 44; pronotum: male 7.5, female 9.5; tegmen: male 25, female 26.5; hind femur: male 16, female 19.5; ovipositor: female 28 mm long, greatest height 3.2 mm.
Karny, 1926: 147; type species: Pseudosubria decipiens Karny, 1926
(male). India: Assam, Imphal (24°51'N, 93°54'E), 22.iii.1946, coll. T.J. Lawrence – depository: BMNH (Lo037S001, B.M.1946–228).
(1 female). India: Assam, Imphal, 15.iii.1946, coll. T.J. Lawrence – depository: BMNH (Lo037S002, B.M.1946–228).
The name of the new species refers to the type locality.
The shape of the male cerci is similar to that in P. bispinosa obtusa Ingrisch, 1998 but has the ventral branch very narrow, regularly upcurved throughout instead of substraight behind the basal bent and reaches the top of the cercus stem (Fig.
Pseudosubria assamensis sp. nov. A–E female: A habitus lateral view; B head and pronotum; C ovipositor; D, E subgenital plate in ventral (D) and lateral view (E). – F–N male holotype: F left tegmen; G stridulatory file; H base of left tegmen; I mirror of right tegmen; J subgenital plate in lateral view; K, L cercus in external (K) and internal (L) view; M, N phallus with titillators. Abbeviations: 7 seventh sternite, m membrane, ov base of ovipositor, sg lateral lobes of subgenital plate.
A rather small, slender species. Fastigium verticis conical, obtuse, with a dorsal furrow: separated by a concavity and a narrow seam from fastigium frontis. Frons shining with impressed dots, nearly subrugose. Tegmen reaching or little surpassing apex of stretched hind tibiae. Femora with the following number of spines on ventral margins: profemur 1–3 external, 2–3 internal; mesofemur 3–4 external, no internal; postfemur 5–7 external, no internal. Knee lobes of fore femur obtuse on both sides, on mid femur obtuse on external, acute on internal side, of hind femur obtuse on external, short spinose on internal side.
Male. Tenth abdominal tergite prolonged; apical margin in lateral areas nearly straight but converging from both sides; in middle obtuse-angularly excised; with a weak medial carina. Epiproct triangularly rounded with deep medial furrow. Paraprocts with a faint swelling on internal side. Cerci short; external surface convex, at dorso-internal margin with a small triangular expansion; at apex with a large triangular internal expansion covering apex of internal side as a cap; in apical third at ventro-medial margin with a long curved stylate projection curved dorsad behind apex of cercus and narrowing towards apex; internal surface flattened, with a weak fold. Subgenital plate long and narrow; with a medial carina fading towards apex; apical margin slightly concave; styli thin. Titillators X-shaped, fused in middle with basal branches longer than apical branches, for the greatest part hyaline; apex of apical branches darkened, separated by a narrow membranous zone from a dentate apical cap.
Female. Epiproct triangular with medial furrow. Cerci conical with a long styliform apex. Subgenital plate split to base into two conical, cap-like, lateral lobes with obtuse tip that in situ lie in wide emarginations of the base of the ventral ovipositor valves.
Ochre. Head and pronotum with a dark brown medial band.
Body w/wings: male 37.5, female 37.5; body w/o wings: male 21, female 19.5; pronotum: male 5.8, female 5.6; tegmen: male 31.3, female 31; hind femur: male 12.5, female 12.5; antenna: female 50; ovipositor: female 12 mm.
Ingrisch, 1998: 108; type species: Depressacca globosa Ingrisch, 1998
(male). East Malaysia: Sabah, Mt. Kinabalu, Mesilau (6°10'N, 116°39'E), at light, 4.ii.1964, coll. J. Smart, Royal Soc. Exped. – depository: BMNH (Lo033S001; B.M.1964–250). The type is unique.
The name of the new species refers to the type locality; noun in apposition.
So far only one Depressacca species has been known. The new species differs from D. globosa Ingrisch, 1998 by the male tenth abdominal tergite that does not have projections but a slightly concave apex, by the male subgenital plate that does not have a medial process and that has only minute styli, by different shapes of the titillators, by the cerci that have the internal tooth inserted subapical, not apical, and by the fastigium verticis that is provided with a distinct tubercle at base.
Fastigium verticis upcurved, with a tubercle at base; apex obtuse; separated by a shallow furrow from fastigium frontis (Fig.
Depressacca kinabalu sp. nov. male holotype: A habitus dorsal view; B fastigium verticis lateral; C head, pronotum and base of tegmen lateral; D face; E left tegmen; F stridulatory file; G subgenital plate and cerci in ventral view; H left cercus in dorsal view; I tip of abdomen in apical view; J titillators of phallus. Abbreviations: 10 tenth abdominal tergite, c cercus, e epiproct, p paraproct, st stylus.
Male. Tegmen with stridulatory area strongly projecting (Fig.
Discolored light yellowish brown; might have been green when alive.
Body: 17; pronotum: 5.7; tegmen: 10.2; hind femur: 7.8 mm.
For the extended loan of specimens and the patience shown for it or for guidance during visits of the museums I am grateful to the former and current curators of the following collections: The Natural History Museum, London, UK (Judith A. Marshall, George Beccaloni), Zoological Museum Hamburg (Hans Strümpel, Martin Husemann). Thanks go also to the reviewers Claudia Hemp and Ming Kai Tan for useful comments on the manuscript. My stay at the Natural History Museum London was supported by a former call of SYNTHESYS (GB-TAF-4995). Printing was supported by the Centrum für Naturkunde (CeNak), University of Hamburg.