taxonID	type	description	language	source
147287CE4B651E0CFE73FF55FEB0D985.taxon	description	Bourgoin (1993 b), however, argued that if Meenoplinae form probably a monophyletic group, monophyly of Kermesiinae remains questionable. Obviously the generic concepts in the Meenoplidae still need to be tested more thoroughly and subdivision of the Meenoplidae must therefore be regarded preliminary.	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
147287CE4B651E01FE77FBF8FD6ADE82.taxon	materials_examined	Type species: Tsingya clarkei Hoch and Wessel sp. nov. Type locality: Madagascar.	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
147287CE4B651E01FE77FBF8FD6ADE82.taxon	description	Description Habitus. Large meenoplid (c. 1 cm), with small compound eyes, well-developed tegmina and wings, and long forelegs. Body pigmentation pale and whitish, except light red compound eyes. Head. (Figure 2). Vertex short, c. 7 × wider than medially long, posterior margin convex, separated from frons by a distinct carina concave towards frons. Frons subrectangular, with lateral carinae strongly ridged, with a dense row of sensory pits along outer margin. Frons and postclypeus medially smooth, anteclypeus with a distinct median carina. Frontoclypeal suture nearly straight, slightly arched towards frons. Rostrum elongate, in repose well surpassing hind-coxae. Compound eyes present, comparatively small, lateral ocelli present, median frontal ocellus vestigial. Antennae with scape cylindrical, pedicel subcylindrical, c. 2 × as long as scape, with distinct rows of sensory plaque organs. Thorax. (Figures 2 and 3). Pronotum short, 1.54 × wider than maximum width of head (including eyes), lateral carinae sinuate, running parallel to posterior margin of head, median carina obsolete; posterior margin of pronotum sinuate, distinctly concave. Mesonotum tricarinate, carinae obsolete, c. 1.48 × wider than long. Tegmina well developed; c. 2.7 × longer than maximal width; anterior and posterior margin nearly parallel, distal margin rounded; distal part not notably expanded; fork of Sc + R and M at about the midlength of Sc + R, Sc + R in proximal threequarters with two rows of sensory pits; fusion of claval veins near middle, claval veins proximally of fusion each with two rows of sensory pits; second claval vein only slightly curved, nearly parallel with posterior margin of tegmen. Wings well developed; venation not notably reduced. Legs overall long and narrow, fore femora distinctly longer than middle femora, middle femora slightly longer than hind femora. Hind tibiae laterally unarmed, distally with a row of eight equally small teeth, arranged in two groups (medially three, laterally five; group of five slightly more proximally than group of three); metabasitarsus and second metatarsal segment each with seven small teeth distally, arranged in a slightly arched row. Metabasitarsus about one-third longer than second and third metatarsal joints together. Male genitalia. (Figures 4 – 7). Genital segment bilaterally symmetrical, in caudal aspect highly ovate, outline nearly figure- 8 - shaped; in lateral aspect caudal margin deeply notched in dorsal third; dorsocaudal margin laterodorsally distinctly projected laterally, ear-shaped. Anal segment in dorsal aspect narrow at base, with lateral margins diverging distally; lateral lobes distally well surpassing anal style, converging medially, in lateral aspect distally rounded and slightly bent dorsally. Parameres bifurcate. Aedeagus differentiated into two processes arising from a common base: dorsal process trilobate, with phallotreme located medially at apex, phallotreme exposed ventrally; ventral process tongue-shaped. External female genitalia. (Figure 8). Strongly reduced, ventral valvifer laterodorsally rounded; ventral valvulae slender, narrow, tip rounded, directed mediocaudad, laterally and ventrally with several smaller and one prominent, almost claw-like seta ventrally near apex; VIIth sternite in ventral aspect narrow, c. 1.3 × longer than wide. The wax-plate areas are of paratopic type (= internal area of wax plate shifted in a median position and joining the body sagittal line) (Bourgoin 1997, p. 202). Internal female genitalia. (Figure 9). Bursa copulatrix spherical with few rounded sclerotinized ornamentations limited to its posterodorsal part; ductus bursa long and thin, emerging from aliferum (Bourgoin 1997, p. 209), ventral wings and the spermatheca posterior of the aliferum. Spermatheca divided into a long and thin ductus receptaculi, ampulla-like distally (= diverticulum ductus), a long pars intermedialis (= spermathecal pump) and the very thin ramified glandula apicalis. Aliferum long and robust, bearing distally a pair of laterodorsally pointed, foliaceous, ear-like expansions (laterodorsal wings) and a ventral pair of flattened cupula-like expansions (ventral wings), enclosing between them the bursa copulatrix ductus. Spermathecal ductus receptaculi running anteroventrally to the ventral wings; both ducts running inside the aliferum joining ventrally the dorsal wall of the vestibulum. Vagina opening anteriorly into the vestibulum at the base of the aliferum. Common oviductus as usual, branched anteroventrally but directed caudad. Lateral sides of sternite VII shortly produced posteriorly, not forming any rod-like arm, reaching gonocoxae VIII ventrally.	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
147287CE4B651E01FE77FBF8FD6ADE82.taxon	diagnosis	Diagnosis Tsingya gen. nov. can be readily distinguished from all other meenoplid genera by the combination of the following characters: large body size (c. 10 – 11 mm including tegmina), tegmina with fusion of claval veins near middle, both claval veins with two rows of sensory pits proximally of fusion, and fork of Sc + R and M at c. midlength of Sc + R. In addition to the tegmina venation, the new genus differs significantly from other known meenoplid taxa by the configuration of the head capsule, the male genitalia (shape of genital segment, anal tube and paramere) and female genitalia (aliferum).	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
147287CE4B651E01FE77FBF8FD6ADE82.taxon	etymology	Etymology The genus name honours the Tsingy de Bemaraha National Park in western Madagascar, characterized by its acutely eroded, sharp-bladed “ tsingy ” limestone pinnacle karst.	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
147287CE4B651E01FE77FBF8FD6ADE82.taxon	discussion	Remarks With regard to the internal structures of the female genitalia, the aliferum (Bourgoin 1997) deserves special attention. The aliferum is a complex of sclerotized formation (s) in the dorsal wall of the posterior vagina forming a more or less tubular structure that encompasses the bursa copulatrix ductus, and also in most cases (but not all: e. g. Anorhinosia Bourgoin 1997), the spermatheca, and which separates them from the vagina. It is a particularly common structure observed in all meenoplid genera, of both subfamilies Kermesiinae and Meenoplinae (but also in several kinnarid planthopper genera: Adolenda Distant, 1911, Bashgultala Dlabola, 1957, Emeljanopleroma Koçak, 1986, Nesomicrixia Emeljanov, 1984, Southia Kirkaldy, 1904 and Oeclidius Van Duzee, 1914) and is diverse in its configuration according to the taxa studied. In Tsingy a gen. nov. the sagittal medioventral lamina is separated into two ventral wings. The unpaired dorsal lamina is absent. The morphological origin of this structure is unclear: at least it is formed by the fusion of the basal apodeme of gonapophyses IX with the posterovaginal process, probably also with parts of the gonospiculum (Bourgoin 1993).	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
147287CE4B681E04FE27FA8FFCDADB20.taxon	description	(Figures 1 – 9) Description Habitus. Unusually large meenoplid, tegmina long, steeply tectiform, small compound eyes, pale bodily pigmentation. Body length. Anterior margin of head to distal margin of tegmina. Males. 10.0 – 10.6 mm (n = 2). Female. 11.5 mm (n = 1). Tip of head to distal margin of anal segment. Males. 6.9 – 7.0 mm (n = 2). Colouration (Figure 1). Body pale (white) throughout, except for compound eyes, these pigmented lightly red; tegmina translucent, venation white. Configuration and proportions of head and thorax incl. legs, tegmina and wings as described for the genus (Figures 2 and 3). Male genitalia. (Figures 4 – 7). Genital and anal segments as described for the genus. Parameres with ventral branch more strongly developed, dorsal branch differentiated into a strong, finger-shaped process bent medially, and a short, stout, apically blunt appendage, densely beset with setae, directed laterodorsally. Aedeagus with dorsal and ventral processes arising from a short, wide common base: dorsal trilobate process forming a half-pipe, with median part hood-shaped, distally subacute with phallotreme located medially at apex, phallotreme exposed ventrally; lateral lobes apically rounded, in lateral aspect slightly surpassing median lobe; ventral aedeagal process tongue-shaped, dorsally at base concave, distally taeniform, narrow, apically acute, its tip curved dorsally. Female genitalia. (Figures 8 and 9). As described for the genus.	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
147287CE4B681E04FE27FA8FFCDADB20.taxon	diagnosis	Diagnosis Tsingy clarkei sp. nov. is characterized by a unique combination of characters pertaining to the male genital structures: parameres bifurcate with ventral branch more strongly developed, aedeagus with a dorsal, trilobate process forming a halfpipe, and phallotreme situated medially at apex and a ventral, tongue-shaped, taeniform and apically acute process.	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
147287CE4B681E04FE27FA8FFCDADB20.taxon	etymology	Etymology The species is named in honour of the collector, Arthur Clarke (Honorary Research Associate in the School of Zoology, University of Tasmania) in Hobart, Tasmania (Australia), ardent cave explorer, dedicated cave ecologist and biospeleologist, reliable caving buddy and good friend.	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
147287CE4B681E04FE27FA8FFCDADB20.taxon	distribution	Distribution Known only from Anjohy Manitsy (translates as “ Cold Cave ”) in the southernmost Petit Tsingy (“ Little Tsingy ”) section of the Tsingy de Bemaraha National Park east of Bekopaka, just north of the Manambolo River, in western Madagascar. Likely to be a narrow range endemic, T. clarkei is known only from Madagascar.	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
147287CE4B681E04FE27FA8FFCDADB20.taxon	biology_ecology	Ecology Clarke (2003) reported that some substantial trees occur in the dense thickets of vegetation formed in the canyons and fault grabens between the elevated blocks of pinnacle karst. However, above the caves the karst surface is generally devoid of large trees and almost barren. Among the perilously razor sharp karren flutes, the surface vegetation in the tsingy itself is unique with many endemic xerophytic and / or water storage plants. Although the Bemaraha tsingy receives torrential downpours in the wet season, very little water remains on the surface and in the dry season the only moisture for 6 – 7 months is the nightly dew, so the karst is essentially an arid environment for much of the year. The “ arid ” flora of the Bemaraha karst surface typically includes cactiform succulents such as Euphorbia and Cynanchum, numerous bulbous stemmed plants or trees that hold water in their internal pith (e. g. Pachypodium, Adenia and the occasional baobab), the many thorned, narrow or spiny leafed xerophytic or xerophilous species such as Pandanus and Dracaena and the unusual leafless plants such as Commiphora with its photosynthesizing trunks and branches (Clarke 2003). Access to the enclosed cave collection site involved passing through a lushly forested canyon, before commencing a traverse of 40 – 50 m along a narrow 1.0 – 1.5 m wide, 30 – 40 m high diaclase (roofless cave) rift with abundant tree roots. The entrance elevation (altitude) of the diaclase is approximately 130 m above sea level. Among the green-leafed foliage in the canyon and in the outer extremities of the diaclase, a number of vertebrate species were observed, including three smaller species of lemur: the common brown lemur Eulemur fulvus, the grey mouse lemur Microcebus murinus and a large nocturnal species the Milne-Edwards’ weasel lemur Lepilemur edwardsi and several reptiles. Along with one of the leaf-tailed Uroplatus geckos, three species of chameleons were noted; two species of Brookesia, Brookesia ebenaui and Brookesia perarmata, and the much longer Malagasy Giant Chameleon Furcifer oustelati. A colubrid snake Madagascarophis colubrinus occidentalis was also spotted in a Pandanus tree (A. Clarke, unpublished records, pers. comm.). In the subdued daylight / twilight inner reaches of the Anjohy Manitsy diaclase rift heading into the roofed cave, several specimens of a locally known so-called nocturnal butterfly (a noctuid moth) were found on the rift walls. In the outer twilight section of the cave, two species of land snails were seen among tree roots and rootlets on the moist sandy floor: Kalidos griffithshauchleri (Ariophantidae) and Acroptychia bathiei (Cyclophoridae). Two frogs were observed in this same outer part of cave: the grey-legged, red and brown-backed Mantella betsileo (Mantelidae) and a speckled brown frog, possibly the Mascarene Ridged Frog Ptychadena mascareniensis (Ranidae). In this outer twilight zone, there was also a dark brown coloured flatworm 12 – 14 cm long, with broadly flattened bulbous head, which may possibly predate on the cavernicolous species including the planthoppers. All three specimens of Tsingya clarkei were found in the inner high humidity dark transition zone of the cave, collected from the subterranean roots of an unidentified surface tree and from smaller rootlets in clay soil adjacent to the cave wall. Among the other cave-dwelling invertebrates in Anjohy Manitsy, there were unidentified species of depigmented isopods, entomobryid springtails and millipedes.	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
147287CE4B681E04FE27FA8FFCDADB20.taxon	discussion	Remarks With 10.0 – 11.5 mm in length, Tsingya clarkei sp. nov. is the largest known meenoplid to date.	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
147287CE4B681E04FE27FA8FFCDADB20.taxon	materials_examined	Material examined Holotype male: MADAGASCAR, Anjohy Manitsy, Petit Tsingy, Tsingy de Bemaraha, east of Bekopaka, western Madagascar, May 2003, Arthur Clarke leg., MNHN. Paratypes. 1 male, 1 female, same data as holotype, MNHN.	en	Hoch, Hannelore, Bourgoin, Thierry, Stelbrink, Björn, Wessel, Andreas (2013): Small giants from Madagascan caves: autapomorphic giantism in the new cave-dwelling planthopper Tsingya clarkei gen. nov., sp. nov. (Hemiptera: Fulgoroidea: Meenoplidae). Journal of Natural History 48 (15 - 16): 919-935, DOI: 10.1080/00222933.2013.840399, URL: http://dx.doi.org/10.1080/00222933.2013.840399
