Research Article Journal of Orthoptera Research 2018, 27(2): 143-153 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 KLAus-GERHARD Hetter!, CLaupiA Hemp2, BRUNo Massa3, MAciey Kocinski, ELzBieta WARCHALOWSKA-SLIWA4 1 Grillenstieg 18, D-39120 Magdeburg, Germany. 2 Senckenberg Biodiversity and Climate Research Centre (BiK-F), Frankfurt (Main), Germany. 3 Department of Agricultural, Food and Forest Sciences, University of Palermo, Italy. A Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakéw, Poland. Corresponding author: Klaus-Gerhard Heller (heller.volleth@t-online.de) Academic editor: Juliana Chamorro-Rengifo | Received 5 February 2018 | Accepted 19 March 2018 | Published 23 July 2018 http://zoobank.org/49DE9A41-355 D-40B0-8612-949CF113449B Citation: Heller K-G, Hemp C, Massa B, Kociriski M, Warchatowska-Sliwa 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 Abstract Madagascar is a well-known hotspot of biodiversity. However, many Or- thoptera, 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 Phanero- pterinae and shares diagnostic characteristics with members of the tribe Am- blycoryphini and its African subtribe Plangiina stat. nov. Paraplangia, which has a chromosome number of 31 XO, differs from other African members of the tribe and subtribe such as Eurycorypha and Plangia, which both have 29 XO. In addition to morphology, we describe the male calling song, female acoustic response, and mating behavior. As calling song, the male produ- ces 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. Key words Amblycoryphini, bioacoustics, chromosomes, duetting, Orophus Introduction 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 (MacArthur and Wilson 1967). Madagascar was formerly part of a continent (for a review see Vences et al. 2009). It separated from Gondwana about 135 million years ago (e.g. Briggs 2003, Yoder and Nowak 2006, McIntyre et al. 2017), so parts of its fauna and flora may be of Gondwanan origin. Then, for some time Madagascar and India drifted north-eastwards together, until their connection also broke (about 90 million years ago) and Madagascar remained isolated in the Indian Ocean. It was, however, never very far from Africa, and even non-flying mammals seem to have crossed this channel sev- eral times (Ali and Huber 2010). Therefore, the time at which the Malagasy taxa diverged from their closest relatives varies greatly, and there are many publications dealing with this question for dif- ferent groups of animals and plants. However, even more groups, especially insect groups, are still nearly completely unstudied. 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 (Carl 1914). Only very recently have some papers on selected groups been pub- lished (Massa 2017a,b,c,d, Unal and Beccaloni 2017). There are no published data on the biology of any taxon (except some obser- vations in Unal and Beccaloni 2017). At present, four subfamilies of Tettigonioidea are represented by several species each in Mada- gascar, and Listroscelidinae is represented by a single species (all data according to Cigliano et al. 2018, abbreviated OSFO). The four more common subfamilies have quite different biogeographic patterns in terms of their Malagasy fauna. Pseudophyllinae and Meconematinae are represented with relatively few species each and only endemic species or genera are known. All Pseudophyl- linae (except the phyllophorine-like genus Aspidonotus) belong to the tribe Simoderini which consists of species that occur exclusively in Australia and Madagascar. If this tribe was confirmed as a mono- phyletic unit, it would be a strong indication for a Gondwanan origin of the group. The subfamily Conocephalinae has many spe- cies in Madagascar and most genera are endemic, but there are also a few endemic species of the widespread genera Conocephalus and Ruspolia and even non-endemic species [Conocephalus maculatus (Le Guillou, 1841) (“Madagasian Region” Pitkin 1980), Pseudorhynchus hastifer (Schaum, 1853) and the swarm-producing (see Bailey and McCrae 1978) Ruspolia differens (Serville, 1838)]. Most conocepha- line genera belong to the tribe Euconchophorini, which is endemic to Madagascar and some surrounding islands. Gorochov (1988) considered this group of short-winged species as a sister group to all other world-wide distributed Conocephalinae. However, in a JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 144 recent molecular study (Hemp et al. 2015) the Euconchophorini representative appeared basally to the studied Agraeciini from Af- rica, India and Australia, but within Conocephalinae. In any case, a member of this Agraeciini/Euconchophorini-complex seems to have arrived in Madagascar relatively long ago. The largest Malagasy subfamily is Phaneropterinae (at present ca. 40 species; Table 1). Madagascar has a few species in common with continental Africa (Phaneroptera sparsa, Tylopsis bilineolata, T. irregularis, and Eurycorypha cereris), and several genera have species which are endemic to Africa as well as to Madagascar - most of them also with endemics on other islands in the Western Indian Ocean (Eurycorypha, Plangia, Parapyrrhicia, Trigonocorypha). One genus with one endemic species to Madagascar has most of its species distributed in Asia (Holochlora). Thus, two thirds of the 22 phaneropterine genera are endemic to Madagascar. Most phan- eropterine species are long-winged. There have probably been several independent colonization events of Madagascar, mostly from Africa [as with the flying mammals (bats), although some of which seem to have come directly from Southeast Asia; Bates et al. 2006]. For genera endemic to Madagascar, the closest relatives are expected to be found in Africa. Considering the low intensity of research, the number of spe- cies and genera known at present is certainly only a small percent- age of that actually occurring there or which had, sadly, occurred there in the past (see Goodman and Jungers 2014). Therefore, one of the authors (KGH) seized the opportunity to rear nymphs of an unidentified species from Madagascar. After these animals molted successfully to adults, they had to be identified (see key below) and, not completely unexpectedly, they were identified as belonging to a new genus and species. Since they lived for quite some time in the laboratory, we were able collect data which were not previously known from any other Malagasy bush-cricket, and which are hopefully a starting point for similar studies in situ. Methods The animals were held in plastic containers, differing in size depending on the size of the animals, and fed with Taraxacum of- ficinale, replaced daily. Measurements.—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). Acoustics. —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 connect- ed 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.hair- ersoft.com). Oscillograms of the songs were prepared using Tur- bolab (Bressner Technology, Germany). All recordings were made K.-G. HELLER, C. HEMP, B. MASSA, M. KOCINSKI AND E. WARCHALOWSKA-SLIWA 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. Acoustical terminology.—Tettigonioids produce their songs by repeat- ed opening and closing movements of their tegmina. The sound resulting during one cycle of movements is called a syllable; open- ing and closing hemisyllables can often be differentiated (Ragge and Reynolds 1998). Syllable duration: time period measured from the first impulse to the last; syllable period: time period measured from the first impulse to the first impulse of the next syllable; im- pulse: a simple, undivided, transient train of sound waves (here: the damped sound impulse arising as the effect of one tooth of the stridulatory file). Typically, after some time the same or a simi- lar pattern of syllables and pauses is repeated. This grouping, often separated from the next by a silent interval, is called a song unit. Chromosomal analysis.—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 chromo- somal analyses. Preparations were obtained from testes and ova- ries, 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 (AgNO,) for localization of the nucleolus organizer regions (NORs) was performed as previous- ly reported (Warchatowska-Sliwa and Maryariska-Nadachowska 1992). Fluorescence in situ hybridization (FISH) with ribosomal 18S rDNA and telomeric (TTAGG), DNA probes were conducted following the protocol described by Warchatowska-Sliwa et al. (2009). Chromosomes were studied with a Nikon Eclipse 400 mi- croscope with a CCD DS-U1 camera and NIS-Elements BR2. Results The specimens of the new species were identified using the key given below (based on Karsch 1889, Brunner von Wattenwyl 1891, Carl 1914, Ragge 1980, OSFO). See also Table 1 for additional data. Using Ragge’s (1980) key for all African Phaneropterinae with open tympana (excluding Madagascar) one would end up with Plangia. Key to the genera of Phaneropterinae of Madagascar and the islands in the western Indian Ocean 1 Fore Coxe without-spilies: ml 8 eee §...nl eer oes nee ee 2 ~ POLO COXA SVMEMES I) Mile a oa nll ac totaal Bel xl aten let tomas Me hee al otic A ala 4 2 Occurring intMadagascaris wna seore.coptuapersacetacten com depectiadaseacenntyeedenee 3 - Occurring in Mauritius only «0.00.0... eeeeeeneees Arantia (two species) 3 Tegmen shorter than 30 mm (observed 25), hind wings not longer iG UPSON Cored OO VT8 Ts ORR Dal ln Ried Beg a Paracosmophyllum atrodelineatum - Tegmen longer than 30 mm (observed 37), hind wings longer than CETL Ae 5 Faire iain Ae eyaeieds agnanseaund yaphadedanielsyaete det fraioss aves Nesoscirtella polita 4 One or both tympana of the fore tibia closed ...........cceeeeseeteeeeeees 5 - Bothetympana-of the fore bia pews. wert cea ok Meenand gonct 6 5 Bothstympana'closed wicwstieci tela meee. Tylopsis (two species) ~ Onlvaintenority lipid Mutt Closed. Si Mew chcse Soe leWenseeieak Neeata sat abseene Tak al ph ae Prheee Holochlora (mauritiana Mauritius, biloba Madagascar) JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 21 22 23 K.-G. HELLER, C. HEMP, B. MASSA, M. KOCINSKI AND E. WARCHALOWSKA-SLIWA 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................cceceees 11 Hind wings hidden under (shorter than) tegmina................ cee 8 Mindswinessla niger than teariitia a. eecstacaencsauadticetre ys duaas sauean pusatecl ee 9 Male supra-anal plate not modified................ Cosmozoma (4 species) Male supra-anal plate enlarged and three-lobed.............eeceeeeeees gia y2 su bssqaiedenp layin ts sng std Miebarsen bea N ate adtaziinintagheon eed Sikoriella (bimaculata) Side keels with 4 large tubercles «0.0.0.0... Polygamus (2 species) Side keels of pronotum crenulated or with small, irregular and indis- PINE TUDOR OSE aes topencapeameneatete narenare ans tepte tenenmartm mma er tenet anounanee 10 Side keels of pronotum dark, crenulated ......Trigonocorypha (maxima) Side keels of pronotum not dark, with small, irregular and indistinct GUberGley se Mk Mit MERE AOE Mea ie Mea AeB RD s ste MeN Megotoessa insulana Tegmina narrow [ratio length/width >3.8; observed (3.8 M.p.*-) Tegmina broad (ratio length/width <3.8; observed 2.1-3.6)......... 18 Hind wings less than half of the length of the tegmina................0.. PRT Tess IE ITS LOL RW Te ROLE LOTT SOT Te PRE OINE Eucatopta heringi Hind wwines*loneer thariteemiiti ag sts.c0h. Ohseknk hte sae ha teaedl ck ak nat 13 Dorsal’side of fore tibia TOUNGEe deo foi) casccesagecdhessaseeesciaceratesasgovevens 14 Dortsal-side:of tore tibia flat or fitrowed 2 Saka wecostey basueweaneleans 15 Margins of pronotum rounded (=side keels indistinct)........0...... iy achat Gtunhobe dat dara rattan earantidy apasbacateavnda rutile can Parapyrrhicia (5 species) Margins of pronotum evident (=side keels smooth but distinct)....... Pars iene ter encore la el oh Od Symmetroraggea (2 species) Tegmina long (>25 mm; observed 29-46 MM) ....... ee eeeeeseeteeeteees 16 Tegmina quite short (<25 mm; observed 14-21 mm)... iy Tegmina long (>32 mm; observed 34-46 MM) ou... eeeeeeeeneeeteeeeees le hates Male Piece Pik bee Mids Die beR 3 ee Mele Std cu Mimoscudderia (4 species) Tegmina short (<32 mm; observed 29 mm)...Agennis (parallelinervis) Last tergite of male unmodified, ovipositor short (4.0-4.7 mm), teg- MITA CIEOLOTE stata: co. pets) oes eae Le Phaneroptera (sparsa) Last tergite of male modified, ovipositor long (7.5-8.0 mm), teg- Mita bicGlored wuneeiiiat Avie Maken neanlewen tess Xenodoxus (2 species) Male subgenital plate with long, up- and recurved appendices ......... Peri sclaneaes tuvesn tustatestemtananaisen abana eeeaaacetavesdaeeises Anchispora appendiculata Male subgenital plate without large appendices ...........ececeeeee 19 Fastigium verticis narrower than SCaPUS.........:.:cccesceeseeeeteeeneeteeeeees 20 Fastigium verticis as wide or wider than Scapus ...........ccceseeeseeeteees 21 Occurring in Madagascar, tegmina wide (ratio length/width = 3)..... Pele NCI a tacts et at ee RT en eet te Paraphylloptera relicta Occurring on the Seychelles, tegmina narrow (ratio length/width = 0 een Wi RN Orc Nic A De Pelerinus rostratus shih te MEARE IM ce Pleat bee MEE, Ped THAR se Me Ma nd Phased M Eurycorypha (4 species) Eyes circular or oval, fastigium verticis less than twice as broad as SCAPUS) fe cvicecviences tesules'ed vas cestgndeusvtuceder sed catenseaseuedeedevederapecetdvauudeusdeds tae DD. Fore femora ventrally armed, ventral edge of ovipositor evenly TOUTE GL Was Ati such A Wie Dk nee dale 2 ce Re Bh el 23 Fore femora ventrally unarmed, ventro-posterior edge of ovipositor slightly curved, its dorsal ending bent frontally (horizontal)............. Pe anesteer ee aves tgusleuet unt acdase aubasuntuert sued vesuansiadoseneetat Paraplangia gen. n. Legs flattened laterally 5 i cece teotardnsduecsvsuneeds Plangia (3 species) Legs not flattened laterally... Madagascarantia (albolineata) Only Mimoscudderia paulyi Massa, 2017 with unusually wide tegmina. 145 Tribe Amblycoryphini Brunner von Wattenwyl Brunner von Wattenwyl (1891) combined several genera in Amblycoryphini using the key character “fastigium as wide or wi- der than scapus”. This may well be a convergently evolved character, but as long as no better grouping is available, it is useful for placing and finding similar genera. The group included and still includes (even after the splitting by Cadena-Castanfeda 2014, 2015) African and American genera which were separated in Brunner von Watten- wyl (1891)’s key but not by Cadena-Castaneda (2014). According to the preliminary molecular tree (Mugleston et al. 2013), Ambly- corypha, the type genus, is deeply nested among New World genera from different tribes. In other molecular studies, Eurycorypha and Plangia are closely related (Mugleston 2016), with Isopsera nearby (Liu Cx unpubl.). Indogneta Ingrisch & Shisodia, 2002, which also has a broad fastigium, certainly belongs to Isopserae (see Kang et al. 2014), and Isopsera has ant-like nymphs like Eurycorypha. We also include Corycomina Karsch, 1896 because of its simi- larity to some Eurycorypha (E. flavescens was considered as member of Corycomina Karsch, 1896; see Massa 2017c). Subtribe Plangiina Cadena-Castaneda, 2015, stat. n. (formerly genus group Plangiae) When splitting up Amblycoryphini, Cadena-Castafieda (2015) listed the similarities between Amblycoryphini s.s. and his Plan- giae. They had all generic characters in common except the eye shape. We consider this single trait to be insufficient for exclud- ing Plangia and Monteiroa from the tribe, especially since Plangiae obviously do not have any similarities to other phaneropterine genera, but share e.g. nymphal properties with Eurycorypha, the largest genus of the tribe (see Hemp in preparation). Despite some doubts, we retain the group as a subtribe and include Madagascar- antia Massa, 2017, which is a sister genus to Plangia, Pseudoplangia Massa, 2014, and Paraplangia gen. n. 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. Madagascarantia albolineata (Brunner von Wattenwyl, 1878), comb. n. syn. n. Madagascarantia bartolozzii Massa, 2017 Paraplangia Heller, gen. n. http://zoobank.org/F80FFF6B-DDAB-4836-8F16-045EGEC76F27 urn:lsid:Orthoptera.speciesfile.org:TaxonName:502224 Type species of the genus.—Paraplangia sinespeculo sp. n., here des- ignated. Description.—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 mar- gin straight, posterior margin rounded, with evident lateral exci- sions where wings are inserted. Prothoracic spiracle slit-like, very JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 146 K.-G. HELLER, C. HEMP, B. MASSA, M. KOCINSKI AND E. WARCHALOWSKA-SLIWA Table 1. 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 Ratio width tegmen/ Ratio length/width Mad. Mau. Com. Sey. pronotum tegmen tegmen length pronotum of tegmen Agennis parallelinervis B.v.W., 1891 e 4.8 29 4.5 0.9 6.4 Anchispora appendiculata B.v.W., 1891 e 6.2 46 13 all i he Arantia (Arantia) dentata Saussure, 1899 e 75 48 10 KS 4.8 Arantia (Euarantia) mauritiana Saussure, 1899 * e 8.5 60 19 2.2 ay Cosmozoma coelebs Carl, 1914 e 9 62 18 2.0 3.4 Cosmozoma doenitzi Karsch, 1889 e 6 37 11.5 1.9 i Sd Cosmozoma sikorae B.v.W., 1891 e 5 30 10 2.0 3.0 Cosmozoma vespertilio Carl, 1914 e 10 65 21 oaeal 3.1 Cosmozoma voluptaria B.v.W., 1891 e 8.2 43 17 21 25 Eucatopta heringi Karsch, 1889 e 35 12.5 3 0.9 4.2 Eurycorypha brevipennis Karsch, 1889 e 5} 21 10.2 2.0 2d Eurycorypha brunneri Brancsik, 1893 e 55 35 14 2.6 25 Eurycorypha cereris (Stal, 1857) e 5 28 9 1.8 3.1 Eurycorypha prasinata Stal, 1874 e e 5 30 10 2.0 3.0 Holochlora biloba Stal, 1874 ° & eo 8 52 14 1.8 Set Holochlora mauritiana Massa, 2017 e 72 48.1 12.2 1.7 4.0 Madagascarantia albolineata (B.v.W., 1878) e 9 48 18 2.0 2.7 Megotoessa insulana Karsch, 1889 e 47 18 2.0 2.6 Mimoscudderia modesta Carl, 1914 e 5 39 7.5 1.5 52 Mimoscudderia paulyi Massa, 2017 e 4.7 45.6 12.1 2.6 3.8 Mimoscudderia picta Carl, 1914 e 5.5 41 7 1.3 59 Mimoscudderia spinicercata Massa, 2017 e 4.5 34.2 6.8 14S 5.0 Nesoscirtella polita Carl, 1914 e 6 37 11 1.8 3.4 Paracosmophyllum atrodelineatum B.v.W., 1891 e 6.5 25 95 iS 2.6 Paraphylloptera relicta Carl, 1914 e 6 39 13 22 3.0 Paraplangia sinespeculo sp. n. e 6.5 42.5 11.7 1.8 3:6 Parapyrrhicia dentipes Saussure, 1899 ® 5 34 - - Parapyrrhicia insularis Chopard, 1958 e 4.5 Oke Vas) ety ez 4.3 Parapyrrhicia longipodex Massa, 2017 e 4.2 26.6 4.2 1.0 6.3 Parapyrrhicia madagassus (Karsch, 1889) e 5 28.5 6.5 1.3 4.4 Parapyrrhicia virilis Carl, 1914 ® 5 Ya - - Pelerinus rostratus (B.v.W., 1878) e 6 38 11 1.8 325 Phaneroptera sparsa Stal, 1857 ¢ e 3.4 17 2.8 0.8 6.1 Plangia guttatipennis Karsch, 1889 e 6 XE 8 1.3 3.4 Plangia ovalifolia Bolivar, 1912 e? @ 4.5 29 10 2.2 29 Plangia segonoides (Butler, 1878) e 8 42.5 18.4 2.3 23 Polygamus macropterus Carl, 1914 e 9 61 16 1.8 3.8 Polygamus punctipennis Carl, 1914 e 6 50 12.5 2,1 4.0 Sikoriella bimaculata Carl, 1914 e 5 28 10 2.0 2.8 Symmetroraggea depravata Massa, 2017 e 4 263 5.4 1.4 4.9 Symmetroraggea dirempta (Karsch, 1889) e 4.5 29 4.5 1%) 6.4 Trigonocorypha maxima Carl, 1914 ® 6 Sy) 15 255 3.5 Tylopsis bilineolata (Serville, 1838) ¢ e 35 20 3.5 1.0 Be Tylopsis irregularis Karsch, 1893 ¢ e 4 30.6 i Fa 1.1 8:7 Xenodoxus annulatus (B.v.W., 1891) e 3 1735 25 0.8 7.0 Xenodoxus nobilis Carl, 1914 ' e 215 20 3.6 1.3 5.6 @ status doubtful, perhaps error in type locality (see Hollier and Heads 2015) > not biloba B.v.W = Holochlora indica Kirby ‘data Ragge 1956 4 data B.v.W. 1878 ¢ data from foto in OSFO ‘ data Massa 2017a JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) K.-G. HELLER, C. HEMP, B. MASSA, M. KOCINSKI AND E. WARCHALOWSKA-SLIWA long, reaching nearly up to metanotum. Ventral edge of paranota rounded. Tegmina much wider than pronotal length. Right teg- men of male without clearly defined mirror. Hind wings longer than tegmina. Fore coxae armed, fore femora unarmed, several spinules posi- tioned ventrally on fore tibiae, furrowed proximally, rounded dis- tally, 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 spi- nules. Hind femora armed ventrally, lower genicular lobe with spine on both sides. Hind tibiae armed ventrally and dorsally, fur- rowed on all sides. Hind tibiae longer than femora. Meso- and metasterna with two lobes each, rounded. Female.—Ovipositor short, curved, but not evenly; ventro-poste- rior 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 (dor- sal area) with transverse veins bearing small teeth. Diagnosis.—Paraplangia differs from most African and Malagasy phaneropterine genera with open tympana by its fastigium. Be- ing 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 (Hemp and Massa 2017), but used by Brunner von Wattenwyl (1891) to differenti- ate genera], which is not observed very often in large species. The paranota are relatively narrow for Plangiina, only about two thirds as wide as high. Paraplangia has other unusual characters, like a stridulatory file with relatively few teeth of varying size, which is quite different from that known in Plangia and Madagascarantia. Its right male tegmen lacking a glossy mirror is not known in any of the similar genera. Derivatio nominis.—Para, Greek = nearby; Plangia, another Plangii- na genus. Paraplangia feminine. Paraplangia sinespeculo Heller, sp. n. http://zoobank.org/03B56614-F8D3-40C1-8652-EFE6F2C80077 urn:lsid:Orthoptera.speciesfile.org:TaxonName:502225 Material examined and depository.—Holotype <, allotype and 1 paratype <@. All pinned, original labels “MADAGASCAR: Mit- sinjo 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 fiir Naturkunde, Berlin, Paratype (CH8240) in Collec- tio 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 1, 2). Measurements.—(In mm) Males. Body length: 27.5-28.2; pro- notum 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 147 femur: 20.9; hind tibiae: 22.0; tegmina: 43.6; length of hind wings: 47.6; tegmina width: 17.2; ovipositor 13.0. Diagnosis.—As for genus (sole species). Description.—Male. Habitus and color: Large bush-crickets, pre- dominantly green with a weak yellow mid line on head and pro- notum (Fig. 1; not visible in dried specimens). Anterior (costal) edge of tegmen in basal half to two thirds with white spots. Head, pronotum and thoracic sternites (Fig. 2) as for genus. The stridulatory area of the left tegmen green, with distinct but not elevated stridulatory vein (Fig. 3A), of right tegmen weakly green or white, with several irregular veins, without glossy mirror (Fig. 3B). The stridulatory vein beginning at the distal end with a series of ca. 15 small teeth, increasing slowly in size. At the same place, the file starting to be elevated above the tegmen level. Ha- ving reached the highest point, there are ca. 10 widely spaced lar- ge teeth. After five of them, tooth size and spacing continuously decrease again. The file then ends in about 10 small to very small teeth (Fig. 3D). Fore coxae armed with a long spine. Fore femora unarmed, fore tibiae with 2-3 inner ventral spines and 3 distal ones, su- periorly 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). Abdomen.—Subgenital plate long, tapering into a deeply incised caudal part, bearing long styli (Fig. 4A). Cerci relatively long, slightly in-curved at tip, with a short, strong, hook-like outer and a rounded inner spine (Fig. 4A-B). No sclerotized titillator. Female.—Color and general habitus like male. Ovipositor as de- scribed for genus (Fig. 5A-B). Subgenital plate short, triangular and at apex slightly incised, side sclerites with strong and incurved lower edge (Fig. 4C), probably the anchor point for the male cerci during mating. Derivatio nominis.—Sinespeculo (Latin = without mirror). To be treated as noun in the nominative singular. Eggs.—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+40.06 mm, n = 4; Fig. 1B). The collector wrote “we found the eggs under leaves”. Nymphs.—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 prepara- tion). 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 Eurycory- pha nymphs (Hemp in preparation; Fig. 1C-E). The pronotum had a middle keel like the adults in Tropidonotacris Chopard, 1954 (Hemp et al. 2014) or Pelecynotum Piza, 1967 (OSFO). The males became adult at 9 and 13" May, the female slightly later, at 20° May. The female died on 30" June due to an infec- tion with Nematomorpha and all were prepared. JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 148 Fig. 1. Habitus. A. Adult male; B. Egg (lateral and ventral view, scale 5 mm); C. Nymph, 3 March; D. Nymph, 30" March; E. Fe- male nymph, 10" May, 10 days before imaginal molt. Fig. 2. Head and pronotum. A. Face (head frontal); B. Pronotum lateral; C. Fastigium verticis and pronotum dorsal; D. Meso- and metasterna, head to the left. K.-G. HELLER, C. HEMP, B. MASSA, M. KOCINSKI AND E. WARCHALOWSKA-SLIWA and C. Female tegmina (scale 5 mm); D. Male stridulatory file (distal end to the left; scale 1 mm). Acoustics. —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 sec- ond only with 4.2+0.9 (range 3-5; Fig. 6). The second series started 9.3 + 0.7 s (range 8.3-10.7 s) after the beginning of the first and both series were separated by a silent interval of 4.2+0.4 s (range 3.7-5.3 s). The intervals between the syllables ranged from 400 to 619 ms (487466 ms), measured in the second half of the first series. In amplitude modulation, both series were decrescendo or without change in loudness. The syllables were very short, less than 5 ms (Fig. 7A), consisting of few, often hardly separable impulses. Like in most phaneropterines (Heller et al. 2015), the fe- male that was ready to mate reacted to the male song with its own acoustic signals. It always answered after or at the end of the second series. The first response syllable of its response was registered 2.7+0.6 s after the beginning of the second male series (range 1.7-4.2 s; n = 74) and 0.55+0.33 s after a male syllable (range 0.04-2.8 s; n = 73). The female response was quite vari- able; the simplest answer consisted of one impulse, but she could also make two impulses at relatively large intervals, long (about 8) series of impulses with short intervals or mixtures with many impulses at varying intervals (Fig. 6B—C; see Suppl. materials 1, 2: duets 1 and 2). The carrier spectrum of male and female song is relatively nar- row-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+40.2 kHz (n = 3; range 10 db below peak 6.9-10.9 kHz; Fig. 7) in the female. Mating behavior.—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 (21% and 24" June). They mated in the morning JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) K.-G. HELLER, C. HEMP, B. MASSA, M. KOCINSKI AND E. WARCHALOWSKA-SLIWA C Fig. 4. Subgenital plate and cerci. A. Male subgenital plate and cerci, ventral; B. Male cercus, dorso-lateral view (dorsum to the left); C. Female subgenital plate with lateral sclerites, ventral view. . os 7 ee ee /% Way. Ato + ae Fig. 5. A, B. Ovipositor of Paraplangia sinespeculo; and other Plangiina for comparison; C. Plangia graminea (Serville, 1838); D. P. guttati- pennis 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; Hemp 2017: C; Hemp et al. 2015: F, H, I. (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%; Hemp et al. 2015). Both spermatophylaces showed relatively irregularly formed central parts protruding anteriorly, in addition to the sym- metrical lateral basal parts (Fig. 8). Chromosomes.—Both analyzed species, the African Paraplangia sine- speculo and Orophus cf. tessellatus from Costa Rica, showed a dip- loid chromosome number of 2n = 31 karyotype in the male with an XO and 32 in the female (O. cf. tessellatus) with XX sex determi- nation system. Fifteen pairs of acrocentric chromosomes gradually decreased in size; the sex chromosome (X) was the largest element in the karyotype (Fig. 9A-D). After FISH with 18S rDNA, silver staining, and C-banding, the results demonstrated coincidence be- tween the localization of major ribosomal genes and active NOR as well as the position of C-bands. In individuals of both species, a single large rDNA cluster per haploid genome was detected, lo- cated on the first pair of autosomes near the distal region (Fig. 9B — left corner) and active NOR (Fig. 9B) as well as C-band (Fig. 9A). JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 150 K.-G. HELLER, C. HEMP, B. MASSA, M. KOCINSKI AND E. WARCHALOWSKA-SLIWA —}— + 5S Fig. 6. Oscillograms of: A. the spontaneous male calling song, and B, C. of two-channel recordings of male-female duets (B: Female response with few impulses; C: With many impulses). > Intensity 188) Intensity | a 0 10 20 30 Frequency 40 k4z 50 Fig. 7. Oscillograms of: A. A male syllable and; B. A female re- sponse together with power spectra (Hanning window, 512 points, mean of 12-ms-section of the song). In contrast, in Orophus cf. tessellatus, FISH revealed a paracentro- meric signal on the fourth pair of autosomes (Fig. 9D) coincident with thick C-bands (Fig. 9C) and active NOR (not shown). The signals produced by FISH with the (ITTAGG), probe were stronger in O. cf. tessellatus (Fig. 9D) than those observed in P. sinespeculo (hardly visible). Discussion Paraplangia sinespeculo belongs to a phaneropterine morpho- type 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; Fontana et al. 2008, and Fig. 8. Female with spermatophore after mating. Syntechna; Cadena-Castafieda 2014), Australia (e.g. Paracaedicia; Rentz 2010), China (e.g. Sinochlora; Kang et al. 2014), and Africa (e.g. some Arantia; Hemp and Massa 2017). They are unlikely to be closely related and do not seem to possess any other special characters in common. Their specific ecological adaptations are unknown for all of them. They may live in trees with similar leaf shape or spend the day at special roosting sites where the shape of the tegmina essentially improves the camouflage among the leaves. However, some morphological similarities may have been overlooked. Many species do not have any or have only weakly de- veloped mirrors on the right wing (checked in OSFO for members JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) K.-G. HELLER, C. HEMP, B. MASSA, M. KOCINSKI AND E. WARCHALOWSKA-SLIWA 151 Fig. 9. 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 chromo- some (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. of the above-mentioned genera; only the morphologically very diverse Arantia seems to also be variable in this character: A. fas- ciata 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; Hemp and Massa 2017). The mirror is an important structure for sound radiation and is considered as typical for bush- crickets (e.g. Montealegre-Z et al. 2017). However, its systematic distribution and size have never been studied in detail. In some species it is a typical ‘mirror’, a large, thin, transparent membrane covering a significant portion of the tegmen’s width, while in oth- ers it is hardly discernible at all (for examples see e.g. fig. 5 in Heller et al. 2014). In Amblycoryphini (Old and New World spe- cies) and some other phaneropterines (see above), it is often, if not always, weakly developed or even missing. On the contrary, in many Poreuomenini, another African tribe which occurs together with Amblycoryphini and Phaneropterini (according to OSFO), it is large and transparent. Correlations with song characteristics are unknown. Unfortunately, in many descriptions of new species, even in recent ones, size and shape of the mirror are often not mentioned nor illustrated. Amblycoryphini are well known for the complexity of their songs, as in the type genus Amblycorypha (e.g. Walker 2004). Some species can produce up to four different syllable types. This com- plexity results obviously from different neuro-muscular programs used for the stridulatory movements. The stridulatory files can differ in tooth number between species (Walker 2004), but do not show any irregularities (see front page of Science, Walker and Dew 1972). Also the song of another New World species, Orophus conspersus (Brunner von Wattenwyl, 1878), contains several different song ele- ments (Taliaferro et al. 1999), produced with a simple file (see Ca- dena-Castaneda 2014). The calling songs of two African Plangia spe- cies, however, are relatively simple (Hemp et al. 2015). They consist of short di- or tri-syllabic echemes or of single syllables, sometimes combined into small series. In contrast, Paraplangia males produce a relatively long song which is answered by the female at its end, suggesting that the latter evaluates the preceding syllable pattern. The very short syllables would seem to be suitable as trigger for a very fast female response (e.g. Heller et al. 2018), but there is no strict temporal correlation between the female response and any parameter of the male song. The female responded with a relatively long and variable delay of about 0.5 s after a syllable (mostly the last) of the second series of the male, so the female signal may be in- terpreted by eavesdropping male rivals as an additional male sylla- ble. Also the carrier frequency and the intervals between additional female responses would fit into this pattern. However, the female sometimes produced quite fast response series with unknown func- tion. The male file with its low teeth number and abrupt change in tooth density differs from all genera mentioned above. Paraplangia has a relatively broad fastigium verticis. This cha- racter is generally rare in long-winged phaneropterines world- wide as well as in Africa (e.g. in Monteiroa and Pseudoplangia; Ragge 1980), but surprisingly frequent in Madagascar (genera Eurycorypha, Madagascarantia, Plangia, Paraplangia). It is unknown if the reasons are phylogenetic, biogeographical, ecological or ac- cidental. In any case, the commonness is not based on a Malagasy radiation. All these genera are currently included in Amblycor- yphini. Surprisingly, they also share a flat (only in some Plan- gia and a few Eurycorypha very slightly sulcate; own unpublished observations) or rounded dorsal surface of the fore tibiae, which is also uncommon among Phaneropterinae. In the amblycoryph- ine genera Amblycorypha and Orophus, fore and middle tibia are dorsally sulcate (Brunner von Wattenwyl 1878). In size, habitus JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) 152 of adults and nymphs, and shape of the male subgenital plate, Paraplangia shows surprising similarities to Gonatoxia (Hemp in preparation), but differs in width of fastigium and structure of the tympana. Representatives of the two Amblycoryphini genera examined in this study, Paraplangia and Orophus, have a karyotype includ- ing 31 (male) or 32 (female) acrocentric chromosomes with XO (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 Ameri- ca, East Africa and India are characterized by such basic/ancestral karyotypes (e.g. for review see Warchalowska-Sliwa 1998, Hemp et al. 2014). It should be noted that this chromosome number, mor- phology (except for the bi-armed X chromosome) and sex deter- mination system was found also in Canadian specimens of Ambly- corypha oblongifolia (De Geer, 1773; Beaudry 1973). However, this plesiomorphic chromosome number is reduced to 29 (XO) in the African genera Plangia (Hemp et al. 2015) and Eurycorypha (Hemp et al. 2013 and unpublished data Warchatowska-Sliwa) as a result of a tandem fusion between two pairs of autosomes. Addition- ally, in comparison to Paraplangia and Orophus with an ancestral acrocentric X chromosome, pericentric inversions modified the position of the centromere and changed the morphology of the X chromosome from acrocentric to bi-armed in A. oblongifolia and some species of Plangia and Eurycorypha. In the Amblycoryphini chromosomes described in this paper as well as in that of Euryco- rypha and Plangia (Hemp et al. 2013, 2015, respectively) one 18S tDNA FISH locus (per haploid genome) coincides with a single active NOR and C-band segment, independently from the number of chromosomes in the set. However, in these species, different patterns of the location of rDNA/NOR were observed. The single 18S rDNA loci revealed by FISH are located distally on the largest autosome in the Malagasy Paraplangia (2n = 31, present study) and the African Plangia (2n = 29, Hemp et al. 2015), but were found in the paracentromeric region of a medium-sized chromosome in the Costa Rican Orophus (2n = 31; present paper) and in an inter- stitial region on a short autosome in African species of Eurycorypha (Hemp et al. 2013). The cytogenetic study presented here consti- tutes another step to better understanding of chromosome organi- zation and the evolution of Phaneropterinae species and genera. 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Annual Review of Ecology, Evolution, and Systematics 37: 405-31. https://doi.org/10.1146/annurev.ecolsys.37.091305.110239 Supplementary material 1 Authors: Klaus-Gerhard Heller, Claudia Hemp, Bruno Massa, Maciej Kocinski, Elzbieta Warchalowska-Sliwa Data type: WAV file Explanation note: Male-female-duet 1 (2-channel-recording) with female response consisting of few impulses (microphone male UHER M645, female SONY ECM-121, T=25°C). Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/ odbl/1.0/). The Open Database License (ODDbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/jor.27.24243.suppl1 Supplementary material 2 Authors: Klaus-Gerhard Heller, Claudia Hemp, Bruno Massa, Maciej Kocinski, Elzbieta Warchalowska-Sliwa Data type: WAV file Explanation note: Male-female-duet 2 (2-channel-recording) with female response consisting of many impulses (microphone male UHER M645, female SONY ECM-121, T=25°C). Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/ odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/jor.27.24243.suppl2 JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2)