Research Article |
Corresponding author: Klaus-Gerhard Heller ( heller.volleth@t-online.de ) Academic editor: Juliana Chamorro-Rengifo
© 2018 Klaus-Gerhard Heller, Claudia Hemp, Bruno Massa, Maciej Kociński, Elżbieta Warchałowska-Śliwa.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Heller K-G, Hemp C, Massa B, Kociński M, Warchałowska-Śliwa E (2018) Paraplangia sinespeculo, a new genus and species of bush-cricket, with notes on its biology and a key to the genera of Phaneropterinae (Orthoptera: Tettigonioidea) from Madagascar. Journal of Orthoptera Research 27(2): 143-153. https://doi.org/10.3897/jor.27.24243
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Madagascar is a well-known hotspot of biodiversity. However, many Orthoptera, and especially the Tettigonioidea, belong to little-studied groups. Here we describe a new genus and species of bush-cricket reared from field-collected eggs. Paraplangia sinespeculo gen. nov., sp. nov. belongs to Phaneropterinae and shares diagnostic characteristics with members of the tribe Amblycoryphini and its African subtribe Plangiina stat. nov. Paraplangia, which has a chromosome number of 31 X0, differs from other African members of the tribe and subtribe such as Eurycorypha and Plangia, which both have 29 X0. In addition to morphology, we describe the male calling song, female acoustic response, and mating behavior. As calling song, the male produces two series of short syllables. At the end of the second series the female responds with signals of similar duration and spectral composition as the male sounds (peak about 8-9 kHz). To make future identification easier, a key to all genera of Phaneropterinae found in Madagascar is presented.
Amblycoryphini, bioacoustics, chromosomes, duetting, Orophus
The island of Madagascar is well known for its richness of endemic species. It is considered to be a very important hotspot of biodiversity due to several factors. Madagascar is a very large island, which is large enough to minimize the risk of extinction once a species has established. It is close enough to the mainland to receive occasional continental immigrants, but is far enough to allow a long, independent evolution of the fauna (
One of these groups is bush-crickets (katydids; Tettigonioidea). Most studies on this group in this region were conducted before 1914, when the last comprehensive paper appeared (
The largest Malagasy subfamily is Phaneropterinae (at present ca. 40 species; Table
Considering the low intensity of research, the number of species and genera known at present is certainly only a small percentage of that actually occurring there or which had, sadly, occurred there in the past (see
List of the known phaneropterine species from Madagascar and some islands in the Western Indian Ocean together with some selected measurements. Abbreviations: Mad = Madagascar, Mau = Mauritius, Com = Comoros, Sey = Seychelles, B.v.W. = Brunner von Wattenwyl. Measurements in mm (taken from original sources).
Species | Distribution | Length of | Width of tegmen | Ratio width tegmen/ length pronotum | Ratio length/width of tegmen | ||||
---|---|---|---|---|---|---|---|---|---|
Mad. | Mau. | Com. | Sey. | pronotum | tegmen | ||||
Agennis parallelinervis B.v.W., 1891 | ● | 4.8 | 29 | 4.5 | 0.9 | 6.4 | |||
Anchispora appendiculata B.v.W., 1891 | ● | 6.2 | 46 | 13 | 2.1 | 3.5 | |||
Arantia (Arantia) dentata Saussure, 1899 | ● | 7.5 | 48 | 10 | 1.3 | 4.8 | |||
Arantia (Euarantia) mauritiana Saussure, 1899 a | ● | 8.5 | 60 | 19 | 2.2 | 3.2 | |||
Cosmozoma coelebs Carl, 1914 | ● | 9 | 62 | 18 | 2.0 | 3.4 | |||
Cosmozoma doenitzi Karsch, 1889 | ● | 6 | 37 | 11.5 | 1.9 | 3.2 | |||
Cosmozoma sikorae B.v.W., 1891 | ● | 5 | 30 | 10 | 2.0 | 3.0 | |||
Cosmozoma vespertilio Carl, 1914 | ● | 10 | 65 | 21 | 2.1 | 3.1 | |||
Cosmozoma voluptaria B.v.W., 1891 | ● | 8.2 | 43 | 17 | 2.1 | 2.5 | |||
Eucatopta heringi Karsch, 1889 | ● | 3.5 | 12.5 | 3 | 0.9 | 4.2 | |||
Eurycorypha brevipennis Karsch, 1889 | ● | 5 | 21 | 10.2 | 2.0 | 2.1 | |||
Eurycorypha brunneri Brancsik, 1893 | ● | 5.5 | 35 | 14 | 2.6 | 2.5 | |||
Eurycorypha cereris (Stål, 1857) | ● | 5 | 28 | 9 | 1.8 | 3.1 | |||
Eurycorypha prasinata Stål, 1874 | ● | ● | 5 | 30 | 10 | 2.0 | 3.0 | ||
Holochlora biloba Stål, 1874 b | ● | ●? | 8 | 52 | 14 | 1.8 | 3.7 | ||
Holochlora mauritiana Massa, 2017 | ● | 7.2 | 48.1 | 12.2 | 1.7 | 4.0 | |||
Madagascarantia albolineata (B.v.W., 1878) | ● | 9 | 48 | 18 | 2.0 | 2.7 | |||
Megotoessa insulana Karsch, 1889 | ● | 9 | 47 | 18 | 2.0 | 2.6 | |||
Mimoscudderia modesta Carl, 1914 | ● | 5 | 39 | 7.5 | 1.5 | 5.2 | |||
Mimoscudderia paulyi Massa, 2017 | ● | 4.7 | 45.6 | 12.1 | 2.6 | 3.8 | |||
Mimoscudderia picta Carl, 1914 | ● | 5.5 | 41 | 7 | 1.3 | 5.9 | |||
Mimoscudderia spinicercata Massa, 2017 | ● | 4.5 | 34.2 | 6.8 | 1.5 | 5.0 | |||
Nesoscirtella polita Carl, 1914 | ● | 6 | 37 | 11 | 1.8 | 3.4 | |||
Paracosmophyllum atrodelineatum B.v.W., 1891 | ● | 6.5 | 25 | 9.5 | 1.5 | 2.6 | |||
Paraphylloptera relicta Carl, 1914 | ● | 6 | 39 | 13 | 2.2 | 3.0 | |||
Paraplangia sinespeculo sp. n. | ● | 6.5 | 42.5 | 11.7 | 1.8 | 3.6 | |||
Parapyrrhicia dentipes Saussure, 1899 | ● | 5 | 34 | – | – | ||||
Parapyrrhicia insularis Chopard, 1958 | ● | 4.5 | 23.5 | 5.5 | 1.2 | 4.3 | |||
Parapyrrhicia longipodex Massa, 2017 | ● | 4.2 | 26.6 | 4.2 | 1.0 | 6.3 | |||
Parapyrrhicia madagassus (Karsch, 1889) | ● | 5 | 28.5 | 6.5 | 1.3 | 4.4 | |||
Parapyrrhicia virilis Carl, 1914 | ● | 5 | 35 | – | – | ||||
Pelerinus rostratus (B.v.W., 1878) | ● | 6 | 38 | 11 | 1.8 | 3.5 | |||
Phaneroptera sparsa Stål, 1857 c | ● | 3.4 | 17 | 2.8 | 0.8 | 6.1 | |||
Plangia guttatipennis Karsch, 1889 | ● | 6 | 27.5 | 8 | 1.3 | 3.4 | |||
Plangia ovalifolia Bolívar, 1912 | ●? | ● | 4.5 | 29 | 10 | 2.2 | 2.9 | ||
Plangia segonoides (Butler, 1878) | ● | 8 | 42.5 | 18.4 | 2.3 | 2.3 | |||
Polygamus macropterus Carl, 1914 | ● | 9 | 61 | 16 | 1.8 | 3.8 | |||
Polygamus punctipennis Carl, 1914 | ● | 6 | 50 | 12.5 | 2.1 | 4.0 | |||
Sikoriella bimaculata Carl, 1914 | ● | 5 | 28 | 10 | 2.0 | 2.8 | |||
Symmetroraggea depravata Massa, 2017 | ● | 4 | 26.3 | 5.4 | 1.4 | 4.9 | |||
Symmetroraggea dirempta (Karsch, 1889) | ● | 4.5 | 29 | 4.5 | 1.0 | 6.4 | |||
Trigonocorypha maxima Carl, 1914 | ● | 6 | 53 | 15 | 2.5 | 3.5 | |||
Tylopsis bilineolata (Serville, 1838) d | ● | 3.5 | 20 | 3.5 | 1.0 | 5.7 | |||
Tylopsis irregularis Karsch, 1893 e | ● | 4 | 30.6 | 3,5 | 1.1 | 8.7 | |||
Xenodoxus annulatus (B.v.W., 1891) | ● | 3 | 17.5 | 2.5 | 0.8 | 7.0 | |||
Xenodoxus nobilis Carl, 1914 f | ● | 2.75 | 20 | 3.6 | 1.3 | 5.6 |
The animals were held in plastic containers, differing in size depending on the size of the animals, and fed with Taraxacum officinale, replaced daily.
Total body length, lateral aspect, refers to the mid-line length of the insect from fastigium verticis to tip of abdomen including the subgenital plate. In females, the ovipositor is not included in the measurement of the body length. Measurements of ovipositor are taken laterally in a straight line from tip to base disregarding the curvature.
To obtain the mass data, living animals and spermatophores were weighed to the nearest 2 mg (balance Tanita Professional Mini 1210– 100).
The male calling song was recorded in the laboratory using a digital bat detector (Pettersson D1000X) with a sampling rate of 100 kHz. Duets were recorded in stereo using a Sony ECM-121 microphone (frequency response relatively flat up to 30 kHz according to own tests) and an Uher M645 microphone connected to a personal computer through an external soundcard (Transit USB, “M-Audio”; 44.1-kHz sampling rate).
Song measurements and spectrograms were obtained using Amadeus II and Amadeus Pro (Martin Hairer; http://www.hairersoft.com). Oscillograms of the songs were prepared using Turbolab (Bressner Technology, Germany). All recordings were made at temperatures between 22 and 25°C. The singers were caged in plastic tubes or gauze cages with microphone fixed or hand-held at distances between 5 (duet) and 80 cm.
Tettigonioids produce their songs by repeated opening and closing movements of their tegmina. The sound resulting during one cycle of movements is called a syllable; opening and closing hemisyllables can often be differentiated (
Paraplangia sinespeculo (two males CH8239, CH8240) and Orophus cf. tessellatus (Saussure, 1861) [Costa Rica; obtained from https://www.saltatoria.info/arten%C3%BCbersicht-a-z-species-a-z/orophus-tesselatus/] (one male CH7705 and one female CH7707) were used for chromosomal analyses. Preparations were obtained from testes and ovaries, incubated in hypotonic solution (0.9% sodium citrate), fixed in ethanol: acetic acid (3:1), and crushed in 45% acetic acid. C-banding was carried out using the method of Summer (1972) and the silver staining method (AgNO3) for localization of the nucleolus organizer regions (NORs) was performed as previously reported (
The specimens of the new species were identified using the key given below (based on
1 | Fore coxa without spine | 2 |
– | Fore coxa with spine | 4 |
2 | Occurring in Madagascar | 3 |
– | Occurring in Mauritius only | Arantia (two species) |
3 | Tegmen shorter than 30 mm (observed 25), hind wings not longer than tegmina | Paracosmophyllum atrodelineatum |
– | Tegmen longer than 30 mm (observed 37), hind wings longer than tegmina | Nesoscirtella polita |
4 | One or both tympana of the fore tibia closed | 5 |
– | Both tympana of the fore tibia open | 6 |
5 | Both tympana closed | Tylopsis (two species) |
– | Only anterior tympanum closed | Holochlora (mauritiana Mauritius, biloba Madagascar) |
6 | Side keels of the pronotum modified (crenulated or serrate or with tubercles etc.) (possibly all members of Trigonocoryphini) | 7 |
– | Side keels of the pronotum smooth or indistinct | 11 |
7 | Hind wings hidden under (shorter than) tegmina | 8 |
– | Hind wings longer than tegmina | 9 |
8 | Male supra-anal plate not modified | Cosmozoma (4 species) |
– | Male supra-anal plate enlarged and three-lobed | Sikoriella (bimaculata) |
9 | Side keels with 4 large tubercles | Polygamus (2 species) |
– | Side keels of pronotum crenulated or with small, irregular and indistinct tubercles | 10 |
10 | Side keels of pronotum dark, crenulated | Trigonocorypha (maxima) |
– | Side keels of pronotum not dark, with small, irregular and indistinct tubercles | Megotoessa insulana |
11 | Tegmina narrow [ratio length/width >3.8; observed (3.8 M.p.*-) 4.3–7.0] | 12 |
– | Tegmina broad (ratio length/width <3.8; observed 2.1–3.6) | 18 |
12 | Hind wings less than half of the length of the tegmina | Eucatopta heringi |
– | Hind wings longer than tegmina | 13 |
13 | Dorsal side of fore tibia rounded | 14 |
– | Dorsal side of fore tibia flat or furrowed | 15 |
14 | Margins of pronotum rounded (=side keels indistinct) | Parapyrrhicia (5 species) |
– | Margins of pronotum evident (=side keels smooth but distinct) | Symmetroraggea (2 species) |
15 | Tegmina long (>25 mm; observed 29–46 mm) | 16 |
– | Tegmina quite short (<25 mm; observed 14–21 mm) | 17 |
16 | Tegmina long (>32 mm; observed 34–46 mm) | Mimoscudderia (4 species) |
– | Tegmina short (<32 mm; observed 29 mm) | Agennis (parallelinervis) |
17 | Last tergite of male unmodified, ovipositor short (4.0–4.7 mm), tegmina unicolored | Phaneroptera (sparsa) |
– | Last tergite of male modified, ovipositor long (7.5–8.0 mm), tegmina bicolored | Xenodoxus (2 species) |
18 | Male subgenital plate with long, up- and recurved appendices | Anchispora appendiculata |
– | Male subgenital plate without large appendices | 19 |
19 | Fastigium verticis narrower than scapus | 20 |
– | Fastigium verticis as wide or wider than scapus | 21 |
20 | Occurring in Madagascar, tegmina wide (ratio length/width = 3) | Paraphylloptera relicta |
– | Occurring on the Seychelles, tegmina narrow (ratio length/width = 3.5) | Pelerinus rostratus |
21 | Eyes elongate, fastigium verticis at least twice as broad as scapus | Eurycorypha (4 species) |
– | Eyes circular or oval, fastigium verticis less than twice as broad as scapus | 22 |
22 | Fore femora ventrally armed, ventral edge of ovipositor evenly rounded | 23 |
– | Fore femora ventrally unarmed, ventro-posterior edge of ovipositor slightly curved, its dorsal ending bent frontally (horizontal) | Paraplangia gen. n. |
23 | Legs flattened laterally | Plangia (3 species) |
– | Legs not flattened laterally | Madagascarantia (albolineata) |
* | Only Mimoscudderia paulyi Massa, 2017 with unusually wide tegmina. |
We also include Corycomina Karsch, 1896 because of its similarity to some Eurycorypha (E. flavescens was considered as member of Corycomina Karsch, 1896; see
When splitting up Amblycoryphini,
After having seen photos of one of the syntypes of Plangia albolineata (described as Turpilia albo-lineata), it turned out that Madagascarantia bartolozzii is conspecific with this species. Since Madagascarantia is considered as generically distinct from Plangia, we propose the following taxonomical corrections.
Paraplangia sinespeculo sp. n., here designated.
Large size, short head, wide round eyes, fastigium verticis as wide as or slightly wider than scapus of antennae, in contact with fastigium frontis, fronto-genal carinae very indistinct. Antennae shorter than tegmina. Pronotum without lateral carinae, length shorter than height, two small pits at two-thirds point on midline; prozona hardly separable from metazona, anterior margin straight, posterior margin rounded, with evident lateral excisions where wings are inserted. Prothoracic spiracle slit-like, very long, reaching nearly up to metanotum. Ventral edge of paranota rounded. Tegmina much wider than pronotal length. Right tegmen of male without clearly defined mirror. Hind wings longer than tegmina.
Fore coxae armed, fore femora unarmed, several spinules positioned ventrally on fore tibiae, furrowed proximally, rounded distally, dorsal side rounded or flat or very slightly furrowed (rounded and slightly furrowed on left and right leg of the same specimen), without dorsal spurs. Tympana open on inner and outer side. Mid femora ventrally with 1-2 spinules, mid tibiae with about ten spinules. Hind femora armed ventrally, lower genicular lobe with spine on both sides. Hind tibiae armed ventrally and dorsally, furrowed on all sides. Hind tibiae longer than femora.
Meso- and metasterna with two lobes each, rounded.
Ovipositor short, curved, but not evenly; ventro-posterior edge of the lower valve only slightly curved, without teeth, dorso-posterior edge sharply bent inwards, this part and distal half of the upper valve serrated. Proximal quarter of right tegmen (dorsal area) with transverse veins bearing small teeth.
Diagnosis.—Paraplangia differs from most African and Malagasy phaneropterine genera with open tympana by its fastigium. Being about as wide as scapus, the fastigium is wider than in most genera, but clearly narrower than in Eurycorypha and Monteiroa. Its width is similar to that of Plangia and Madagascarantia. These two genera, however, have evenly curved ovipositors. In Paraplangia, the fore femora are also unarmed [a character occasionally found within otherwise armed genera: e.g. Arantia (
Para, Greek = nearby; Plangia, another Plangiina genus. Paraplangia feminine.
Holotype ♂, allotype ♀ and 1 paratype ♂. All pinned, original labels “MADAGASCAR: Mitsinjo Forest Reserve, near Moramanga (18°57’S, 48°13’E), 1 i - 31 xii 2014, coll. Giesse”. “Holotype Paraplangia sinespeculo” [red handwritten label]. Holo- (CH8239) and allotype (CH8241) in Museum für Naturkunde, Berlin, Paratype (CH8240) in Collectio Heller. One hind leg of CH8240-1 separate in pure ethanol in Collectio Heller.
Sound files are deposited at OSFO and bio.acousti.ca (see also Suppl. materials
Measurements.—(In mm) Males. Body length: 27.5–28.2; pronotum length: 6.8–7.4; pronotum height: 6.9–7.0; hind femur: 19.5–20.0; hind tibiae: 21.9–23.5; tegmina: 43.2–43.8; length of hind wings: 46.4–46.6; tegmina width: 14.3–14.7. Female. Body length: 29.9; pronotum length: 7.8; pronotum height: 7.0; hind femur: 20.9; hind tibiae: 22.0; tegmina: 43.6; length of hind wings: 47.6; tegmina width: 17.2; ovipositor 13.0.
As for genus (sole species).
Male. Habitus and color: Large bush-crickets, predominantly green with a weak yellow mid line on head and pronotum (Fig.
The stridulatory area of the left tegmen green, with distinct but not elevated stridulatory vein (Fig.
Fore coxae armed with a long spine. Fore femora unarmed, fore tibiae with 2–3 inner ventral spines and 3 distal ones, superiorly mainly flat (see above). Mid femora with 2 very small spinules, mid tibiae ventrally with about 7 spines each on outer and inner side. Hind femora distally with about 7 outer and 2-4 inner ventral spines, hind tibiae straight, longer than femora, in cross-section square, with many spines on all four edges (about 15 on each ventral, about 25 on each dorsal side).
Subgenital plate long, tapering into a deeply incised caudal part, bearing long styli (Fig.
Color and general habitus like male. Ovipositor as described for genus (Fig.
Sinespeculo (Latin = without mirror). To be treated as noun in the nominative singular.
Mature eggs were taken from the female after her death and preserved in ethanol. They show the flat, ovoid shape, typical for phaneropterines (length 5.7± 0.2 mm, width 2.75±0.06 mm, n = 4; Fig.
The specimens were obtained as relatively small nymphs (probably stage 2 or 3). At that time they were nearly completely green with few brown markings (not looking like the small nymphs of Plangia satiscaerulea; see Hemp in preparation). Soon they showed a yellow midline running from head to mid of abdomen and they developed a brown pattern at the back of the abdomen, similar to that seen in older Eurycorypha nymphs (Hemp in preparation; Fig.
The male calling song consisted of song units, repeated in interval of many minutes as long as the female did not respond. Each song unit (163 recorded) contained two series of syllables, the first with 10.7±1.0 syllables (mean±SD; range 8-12; n = 21), the second only with 4.2±0.9 (range 3-5; Fig.
Like in most phaneropterines (
The carrier spectrum of male and female song is relatively narrow-banded with its maximum at about 8.4±0.7 kHz (n = 3; range 10 db below peak 7.3-10.2 kHz) in the male and 8.7±0.2 kHz (n = 3; range 10 db below peak 6.9-10.9 kHz; Fig.
Both males (body mass 1178 mg and 1236 mg) copulated with the female (body mass 2536 mg, 2528 mg) at an interval of 3 days (21st and 24th June). They mated in the morning (about 9:00–10:00; they were placed together the evening before, but did not mate) with mating durations of a few minutes and transferred spermatophores of 130 and 151 mg (mean of male loss and female gain), thus about 12% of the male body mass. They were slightly smaller than in Plangia multimaculata (17%;
Both analyzed species, the African Paraplangia sinespeculo and Orophus cf. tessellatus from Costa Rica, showed a diploid chromosome number of 2n = 31 karyotype in the male with an X0 and 32 in the female (O. cf. tessellatus) with XX sex determination system. Fifteen pairs of acrocentric chromosomes gradually decreased in size; the sex chromosome (X) was the largest element in the karyotype (Fig.
A, B. Ovipositor of Paraplangia sinespeculo; and other Plangiina for comparison; C. Plangia graminea (Serville, 1838); D. P. guttatipennis Karsch, 1889; E. P. karschi Chopard, 1954; F. P. multimaculata Hemp, 2017; G. P. nebulosa Karsch, 1890; H. P. satiscaerulea Hemp, 2017; I. P. variacantans Hemp, 2017; J. Madagascarantia albolineata (syntype); K. Pseudoplangia laminifera (Karsch, 1896). Sources OSFO: D, E, G, J, K;
Chromosomes of: A, B. Paraplangia sinespeculo and; C, D. Orophus cf. tessellatus stained using different techniques: C-banding (A, C), silver staining (B), and FISH with both 18S rDNA (green) and telomeric DNA (red) probes (B – left corner and D). Diakinesis and the insert in the left corner mitotic large chromosome (A), diplotene and FISH with 18S rDNA in the left corner mitotic chromosome (B): arrows and the insert chromosome demonstrated distally located thin C-band and single NOR coincide with cluster of 18S rDNA on the first pair of autosome. Arrows in spermatogonial (C) and female metaphases (D) indicate near paracentromeric located C-bands and 18S rDNA loci on the fourth pair of chromosomes. X indicates sex chromosome.
Paraplangia sinespeculo belongs to a phaneropterine morphotype which is widespread in many tropical areas. Such medium to large species with long wings of intermediate width and white markings at the costal edge of the tegmina are found in central America (e.g. Lamprophyllum, Philophyllia;
However, some morphological similarities may have been overlooked. Many species do not have any or have only weakly developed mirrors on the right wing (checked in OSFO for members of the above-mentioned genera; only the morphologically very diverse Arantia seems to also be variable in this character: A. fasciata has a large distinct mirror, while some other Arantia species have small and/or indistinct ones, and A. mauritiana does not have one at all;
Amblycoryphini are well known for the complexity of their songs, as in the type genus Amblycorypha (e.g.
Paraplangia has a relatively broad fastigium verticis. This character is generally rare in long-winged phaneropterines world-wide as well as in Africa (e.g. in Monteiroa and Pseudoplangia;
Representatives of the two Amblycoryphini genera examined in this study, Paraplangia and Orophus, have a karyotype including 31 (male) or 32 (female) acrocentric chromosomes with X0 (male) and XX (female). This chromosome number coincides with previous studies, which revealed that more than 50 genera of phaneropterines found in the Palaearctic region, South America, East Africa and India are characterized by such basic/ancestral karyotypes (e.g. for review see
Our thank goes to Dr. J. Hollier, Muséum d’histoire naturelle (MHN) Genève, for organizing photos of one of the syntypes of Plangia albolineata. For helpful comments on an earlier version of the manuscript we are grateful to Dr. Holger Braun and an anonymous reviewer.