New species of the genera Limentinus Distant, 1917 and Calodia Nielson, 1982 (Hemiptera, Auchenorrhyncha, Cicadellidae, Coelidiinae) from the Makay Massif of Madagascar, with a key to Malagasy species

ABSTRACT Three new species of the genus Limentinus Distant, 1917 and one new species of the genus Calodia Nielson, 1982 are described from the former Toliara Province of southwestern Madagascar: Limentinus oryx n. sp., L. nielsoni n. sp., L. nigrifacies n. sp., Calodia makayensis n. sp. The genus Calodia Nielson, 1982 is recorded for the first time from Madagascar. Coelidia perineti Evans, 1953 is redescribed and transferred to the genus Limentinus with a new combination formed, Limentinus perineti (Evans, 1953) n. comb. Key to species of the tribe Coelidiini Dohrn, 1859 known currently from Madagascar is given.


INTRODUCTION
The cicadellid tribe Coelidiini Dohrn, 1859 of the subfamily Coelidiinae Dohrn, 1859 was known so far from Madagascar only after the genus Limentinus Distant, 1917(Nielson 1982, 1991. This genus was erected for a single species, Limentinus aldabranus Distant, 1917, from Aldabra atoll in Outer Seychelles (Distant 1917) and recorded later also from Cosmoledo and Astove atolls (Webb 1980). Two years later this species was recorded from Antsiranana (or Diego-Suarez) in northern Madagascar, with male genitalia illustrated for the first time (Nielson 1982). Subsequently, the synonymy of L. aldabranus with Coelidia perineti Evans, 1953, the latter described from Analamazaotra (or Périnet) in Eastern Madagascar, was suggested by Nielson (1982) and confirmed by Nielson & Zahniser (2012). However, this synonymy cannot be accepted according to the differences in male genitalia structure illustrated below. Thus, I treat Coelidia perineti as valid name and transfer it to the genus Limentinus.
In the same paper by Nielson (1982) Coelidia cambouei Signoret, 1886, originally described after a female from the environs of Tamatave (Atsinana Region in eastern Madagascar), was transferred to Limentinus with new material listed from Mailaka and Antatabe. Nine years later Nielson (1991) added three more new species from Madagascar: L. sagittus Nielson, 1991 -from Toamasina Province (Fampanambo); L. bracchius Nielson, 1991 -from Nosy Boraha Island (or Sainte Marie); L. varius Nielson, 1991 -from Comanarita (the locality name was not deciphered exactly after handwritten label). Finally, L. declinatus Wang, Dietrich & Zhang, 2018 was described from nearby of Fianarantsoa. Thus currently, Limentinus is represented by seven species known mainly from the holotypes and including three species known only after the males. According to Nielson (1982), Limentinus Distant is closely related to African Krosolus Nielson, 1982 andOriental Calodia Nielson, 1982, but well distinguished by its tubular aedeagus with spine-shaped processes.
Up to now the subfamily Coelidiinae was known from Madagascar from just 17 specimens of Limentinus mostly from northern and eastern regions of the island (Signoret 1886;Evans 1953;Nielson 1982Nielson , 1991Wang et al. 2018). However material collected during a field trip to the Makay Massif of former Toliara Province of southwestern Madagascar in 2011, organised by the Association "Naturevolution", revealed three new species of Limentinus and a new species of the genus Calodia Nielson, 1982 -the largest coelidiine genus with Oriental distribution (Nielson 1982; Viraktamath & Meshram 2019) which had not previously been recorded from the island. Thus, the total number of Coelidiini genera and species known from Madagascar increases to two and 11 respectively including new records of this tribe from the former Toliara Province. A key to all Malagasy Coelidiini is given below.

MATERIAL AND METHODS
Morphological terminology follows Anufriev & Emeljanov (1988) and Dietrich (2005). The terms "anal tube" refers to the X segment and "anal column" refers to the paraproct.
The genital segments of male specimens examined were macerated in 10% KOH and figured in glycerin jelly (Brunel Micro Ltd, UK) using Leica MZ9.5 light microscope with camera lucida attached. The photos were taken using same microscope with camera Leica DFC 290 and Canon EOS 6D camera with macrolens Canon MP-E 65 mm f/2.8 1-5X. Images are produced using the software Helicon Focus and Adobe Photoshop.
The material examined, including the type specimens of the species described below, are deposited in the Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia (ZIN)  diagnosis. -Head short, together with eyes narrower than pronotum; anterior margin rounded. Face long; frontoclypeus almost twice as long as wide between the eyes, without carinae; lorae narrowly oval; anteclypeus distinctly (mushroomlike) enlarged apically, with straight basement (Fig. 6A); antennal pedicel small, cylindrical; ocelli prominent, on margin of face and crown; rostrum with short 3rd segment, reaching middle coxae. Crown with lateral margins diverging anteriorly with weak median carina. Pronotum without carinae, anterior margin convex; posterior margin concave. Mesonotum large, with scutellum well separated by transverse depression; disk of pronotum separated by carina from its lateral parts (paranotal lobes). Forewings elongate, with five apical and three anteapical cells, outer one closed; appendix well developed. Hind femora with 2 + 2 macrosetae apically. First metatarsomere as long as second and third ones combined. First metatarsomere with 2 + 1 macrosetae dorsally -two apically and one laterally. First metatarsomere with two spines laterally and two platellae between them ventroapically (Fig. 5A). Second metatarsomere with two spines laterally and one or two platellae between them ventroapically. Claws wide, each with single long lateral seta. Aedeagus tubular, asymmetrical, with long and narrow shaft bearing spiny processes. Connective short and broad, arcuated.
Limentinus perineti (Evans, 1953)  diagnosis. -Aedeagal shaft with three spine-shaped straight processes subapically on right side and one spine-shaped straight subapical process on left side and with two short and two long spine-shaped slightly curved processes below on right and left sides (Fig. 5E, F).
desCripTion Structure As mentioned for the genus. Male genitalia (Fig. 5) Anal tube 1.5 times as long as wide, flattened ventrally. Anal column short. Pygofer lobe triangularly elongate, with an additional small lobe on apical angle under anal tube and with ventral lobe in its middle (Fig. 5B); apical lobe with seven macrosetae subapically on its inner side (Fig. 5C). Subgenital plate long and narrow, not narrowing apically, with microsetae apically and with long setae on main surface, including several subapical macrosetae (Fig. 5D). Aedeagus asymmetrical, shaft long and narrow, slightly curved, with small hook apically bearing marginal denticles ( Fig. 5E-G), with three (2 + 1) spine-shaped straight processes subapically on right side and one spine-shaped straight subapical process on left side and with two short and two long (one is damaged) spineshaped slightly curved processes below on right and left sides respectively (Fig. 5E, F); gonopore probably subapical. Styles elongate, narrowing apically, with rounded apices (Fig. 5H). Connective arcuate (Fig. 5I).

Measurement
Total length from the apex of head to the apices of forewings: 7.0 mm. diagnosis. -Aedeagal shaft with two spine-shaped processes subapically on right side and one spine-shaped subapical process on left side and with two longer spine-shaped processes below on left side (Fig. 6D, E).
TaxonomiC relaTionships. -Very close to L. declinatus Wang, Dietrich & Zhang, 2018 in coloration, but well distinguished by lighter forewings and dense brown dots on pronotum and by fewer spiny processes of aedeagus -five processes in comparison to 12 in L. declinatus.
desCripTion Structure Structure as mentioned for the genus. Crown narrow and long, 1.5 times as long at midline as wide basally. Pronotum nearly as long as crown medially. Mesonotum 1.5 times as long as pronotum medially. Second metatarsomere with two spines laterally and one platella between them ventroapically. Apodemes of second abdominal segment rather long and wide (Fig. 6C).
Coloration (Fig. 4A Male genitalia (Fig. 6) Anal tube 1.5 times as long as wide, flattened ventrally (Fig. 6F). Anal column short. Pygofer lobe triangularly elongate, with an additional small lobe on apical angle under the anal tube and with ventral lobe in its basal half (Fig. 6B). Subgenital plate long and narrow, with around 12 microsetae apically and with long setae on the main surface. Aedeagus asymmetrical, shaft long and narrow, curved, with small hook apically (Fig. 6D), with two spine-shaped slightly curved processes subapically on right side and one spine-shaped slightly curved subapical process on left side and with two longer slightly curved spine-shaped processes below on left side (Fig. 6E), gonopore probably subapical. Styles elongate, narrowing apically, with rounded apices (Fig. 6G). Connective arcuate (Fig. 6H).

Measurement
Total length. 6.0 mm. diagnosis. -Aedeagal shaft with three straight short processes apically, two spine-shaped slightly curved processes subapically on right and left sides (right processes longer) and two spine-shaped slightly curved processes (longest ones) below on left side (Fig. 7D-F).
desCripTion Structure In structure and coloration very similar to L. oryx n. sp. Second metatarsomere with two spines laterally and two platellae between them ventroapically.
Coloration (Fig. 3A, B) Scutellum with brown yellowish margins. Abdominal tergites black, each with pair of large orange yellow spots laterally running from its anterior margin to its yellow posterior margin. Abdominal sternites black, with yellow hind margins.
Male genitalia (Fig. 7) Anal tube 1.5 times as long as wide, flattened ventrally (Fig. 7G, H). Anal column short. Pygofer covered by long setae; pygofer lobe triangularly elongate, with an additional small lobe on its apical angle under the anal tube and with ventral lobe in its basal half (Fig. 7A, B); apical lobes of pygofer each with eight and more setae on its inner side (Fig. 7B). Subgenital plate long and narrow, with microsetae apically and with long setae on the main surface. Aedeagus asymmetrical, shaft long and narrow, slightly curved (in ventral and lateral views), with small hook apically (in lateral view) (Fig. 7D), with three straight short processes apically, two spine-shaped slightly curved processes subapically on right and left sides (right processes longer) and two spine-shaped slightly curved processes (longest ones) below on left side (Fig. 7D-F), gonopore subapical, large (Fig. 7F). Styles small, elongate, with rounded apices (Fig. 7D). Connective arcuate (Fig. 7C).

Measurement
Total length. 6.0 mm. diagnosis. -Aedeagal shaft with one short subapical process on left side, two long spine-shaped processes below it, and two long spine-shaped processes on opposite right side (Fig. 8D, E).

Limentinus nigrifacies
TaxonomiC relaTionships. -In general the male genitalia structure is close to L. aldabranus Distant sensu Nielson (1982), but is well differentiated by its black face and details of the aedeagus.
desCripTion Structure General structure as mentioned for the genus. Crown wider than in other species described, as long as wide basally. Second metatarsomere with two spines laterally and two platella between them ventroapically.
Coloration (holotype) (Fig. 3C, D Coloration (paratype) (Fig. 3E, F ible segment and brown yellowish 2 nd one. Crown brown yellowish, with black longitudinal areas. Pronotum brown yellowish, except black paranotal lobes. Mesonotum brown yellowish. Forewings brown yellowish, with dark brown veins. Episternae and epimerae with yellow margins. Katepisternae yellow, with dark brown to black central part. Fore and middle coxae dark brown. Hind coxae black. Fore and middle femora and tibiae dark brown, with red tint. Hind femora and tarsomeres brown yellowish. Male genitalia (Fig. 8) Anal tube 1.4 times as long as wide, flattened ventrally (Fig. 8B). Anal column short. Pygofer covered by long setae; pygofer lobe triangularly elongate, with an additional small lobe on apical angle under anal tube and with ventral lobe in its basal half (Fig. 8A, C); apical pygofer lobe apically with macrosetae on its inner side (Fig. 8C). Subgenital plate long and narrow, with long setae on the main surface. Aedeagus asymmetrical; shaft long and narrow, weakly curved, with large hook apically bearing five small lateral denticles on each side (in ventral and lateral views) (Fig. 8D-G), with one short subapical process on left side, two long spine-shaped slightly curved processes below it, and two long spine-shaped slightly curved processes in opposite on right side, gonopore situated below long processes of shaft, small (Fig. 8D). Styles small, rounded (Fig. 8D, E). Connective arcuate.
diagnosis. -As given by Nielson (1982Nielson ( , 2015. In general characters close to Limentinus mentioned above, but well differentiated in male genitalia structure by the aedeagal shaft with fewer processes and larger styles (Fig. 9C, F-H). diagnosis. -Aedeagal shaft with only pair of different in shape and length spine-shaped subapical processes (Fig. 9G, H).
TaxonomiC relaTionships. -According to the shape of aedeagus with right subapical process longer then left one and aedeagal apex with carina below gonopore (Fig. 9G, H) (Fig. 9A, C). The peculiar long style, with median process (Fig. 9F), separates the new species from other Calodia species as well as from all other Coelidiini. Long and narrow styles, but without median processes, are known also for the African genera Africocoelidia Nielson, 1982and Amplicoelidia Nielson, 1991(Nielson 1991.
desCripTion Structure Crown slightly longer than wide basally. Pronotum 1.3 times as long as coryphe medially. Mesonotum 1.5 times as long as pronotum. Second metatarsomere with two spines laterally and two platellae between them ventroapically.
Coloration (Fig. 4C, D) Face with frontoclypeus light yellow, with dense brown dots; genae light brownish yellow; lorae, anteclypeus, legs, and body from below light yellow; ocelli burgundy; crown light yellow. Pronotum and mesonotum dark brown, with dense light yellow dots; paranotal lobes of pronotum light yellow. Forewings ochre, with dark brown to black veins, bearing light dots. Hindwings cinereous, with dark brown veins. Claws dark brown to black. Hind tibiae with brown to dark brown apices of spines. Abdominal tergites brown to dark brown, with light yellow hind margins. Pygofer lobes with light brown yellowish basal and apical angles and dark brown medially and from above; subgenital plate light brown yellowish basally and dark brown laterally and apically; anal tube dark brown.
Male genitalia (Fig. 9) Anal tube 1.5 times as long as wide, flattened dorso-ventrally, slightly enlarged from base to apex (in dorsal view) (Fig. 9A, B). Anal column short. Pygofer covered by long setae; pygofer lobe triangularly elongate, with an additional small lobe on apical angle under anal tube, without ventral lobe (Fig. 9A); pygofer lobe apically with 4-5 macrosetae on its inner side (Fig. 9D). Subgenital plate long and narrow, rounded apically (Fig. 9C). Aedeagus asymmetrical; shaft long and narrow, straight (in ventral view), wider in its basal third, with As many species mentioned above are described after a single specimen, in particular, occasionally swept in the forests during the trip to the Makay Massif, I expect many more undiscovered taxa to be found in other regions of the island. Thus, Olidiana indica (Walker, 1851) recorded by Distant (1917) from Seychelles may still be found in Madagascar. Further discovering and description of these taxa will considerably increase our knowledge of Malagasy Coelidiini and may bring more genera to the list.