Carinetini Distant, 1905a

Tribe Carinetini Distant, 1905a View in CoL

Carinetaria Distant 1905a: 483.

Type genus. Carineta Amyot & Audinet-Serville 1843: 482 .

Remarks. Carinetini was defined using seven general characters when it was erected including some contradictory characters (e.g. hyaline but possibly semi-opaque or infuscated fore wings, and only fore wings variable in coloration but the type species of the type genus has infuscated hindwings and immediately after forming the tribe he described one new species with infuscated hindwings and another with semi-opaque hindwings) (Distant 1905a) (see complete original description below). There were four genera originally included in the tribe, one from China (which is now classified in a different subfamily), one from Africa, and two (including the nominotypical genus) from the Neotropics (Distant 1905a). Since that time, four additional Neotropical genera have been added to the tribe (see summary in Marshall et al. 2018). The inclusion of the African genus is questionable and re-evaluated here based on its morphology.

The African genus Paranistria Metcalf, 1952 (nom. nov. pro Tympanistria Stål, 1861: 619 nec Tympanistria Reichenbach, 1852 : XXV) is currently the only genus of the Carinetini not inhabiting the Neotropical region. Although the two remaining species of Paranistria have general characteristics of the Carinetini as the tribe was originally described, the laterally expanding central abdomen and large lateral angles of the pronotal collar in Paranistria species are distinctly different than the tapering abdomen and non-ampliate lateral pronotal margin used as defining characteristics of the Carinetini (Distant 1905a) . Investigating the placement of the genus, and what appear to be related African genera, leads to a series of conflicting data in determining a final tribal assignment for the genus.

Boulard (1990) reassigned Paranistria fornicata (Linnaeus, 1758) to Zouga Distant, 1906b and incorrectly synonymized Neomuda Distant, 1920 into Zouga . These changes were done without discussion in footnotes to the paper. Stephen (2021) suggested that Paranistria is a junior synonym of Neomuda Distant, 1920 but did not perform a detailed analysis nor make an official change under the Code. This leads to a problem in that the two genera that have been suggested to be closely related to Paranistria are currently assigned to two different tribes.

Zouga is currently classified in Parnisini Distant 1905b while Neomuda is currently assigned to Lamotialnini Boulard, 1976 . Specimens from neither genus (nor any of the related genera) were available for the family level genetic analysis of Marshall et al. (2018) so the tribal assignment of these genera could not be confirmed with the genetic tests. The more recent genetic analysis of Owen et al. (2022) placed Zouga between genera of Chlorocystini Distant, 1905b and Katoini Moulds & Marshall (in Marshall et al. 2018), 2018 and separated from the Parnisini genus included in that analysis. It has already been suggested that Parnisini may be an unnatural grouping of genera and some of the African genera may be incorrectly included in the Lamotialnini (Marshall et al. 2018) .

There are several African genera related to Paranistria that are currently classified in either Parnisini or Lamotialnini . These genera may require a new tribe based on the current data available but genetic analyses appear to be required to determine the final details as to how these genera are related to each other and the tribes to which they are currently assigned (M. Villet, personal communication).

Based on images of a junior synonym ( Carineta leuconeura Walker, 1850 ) of Zouga (formerly Paranistria ) fornicata and Paranistria trichiosoma Walker, 1850 , the general morphology supports the reassignment of Paranistria to the Parnisini . The eyes do not project distinctly beyond the anterior angles of pronotum and the length between the apex of the head and the posterior of the cruciform elevation is longer than the length of the abdomen, all the defining characteristics of Parnisini given by Distant (1905b). The anterolateral vertex is shorter than the supra-antennal plate, fore wing cubitus anterior 1 vein is divided by the crossvein so that the distal portion is longest, the hindwing anal vein 3 does not curve at the distal end, the hindwing anal lobe is narrow, abdominal tergites 2 expands laterally and tergites 2 and 3 are wider than tergites 4–7, all of which contradict the characteristics of Lamotialnini (Marshall et al. 2018) . As a result, the genus Paranistria Metcalf, 1952 is reassigned to Parnisini Distant 1905b here to align the genus with the majority of related African genera and the species ( Zouga fornicata ) that was once assigned to Paranistria .

This would not be the first example of genera assigned by Distant to a tribe from multiple geographic regions that have been reassigned with morphologically more closely related taxa in a more suitable tribe. For example, the Taphurini Distant, 1905c is now a tribe restricted to the Neotropical region that included genera with a worldwide distribution for most of its history (Marshall et al. 2018; Sanborn 2021a). In fact, Distant (1905c) mistakenly assigned a current member of the Carinetini to the Tibicinini Distant, 1905c of the Tibicininae Distant, 1905d when the genus Ahomana Distant, 1905b was described. It remained in the wrong tribe for more than 100 years before being reassigned to Carinetini (Sanborn 2014a) . Although the assignment of some genera has been updated, Distant is commended for tackling the entire group and forming taxa which significantly helped to organize how cicadas were classified in order to produce the first synonymic catalogue of cicadas (Distant 1906a).

Included genera. Ahomana Distant, 1905c , Carineta Amyot & Audinet-Serville, 1843 , Guaranisaria Distant, 1905e , Dorachosa Distant, 1892b , Novemcella Goding, 1925 , and Toulgoetalna Boulard, 1982 .

Diagnosis. The tribe was erected based on the following set of characters: a pronotum that is shorter than the mesonotum and distinctly narrowed anteriorly with oblique lateral pronotal margins that are not ampliate, a more or less robust body narrowed towards the head and apex of the abdomen, and hyaline fore wings and hindwings although the fore wings may be semi-opaque or infuscated (Distant 1905a). This generalized set of characters is no longer sufficient to distinguish members of the tribe so a more detailed diagnosis is provided here to compliment more recent tribal descriptions (e. g. Marshall et al. 2018).

Head including eyes narrow, less than width across lateral pronotal angles, may be wider or narrower than mesonotum; supra-antennal plates extending about half distance to eye; eyes protruding laterally from head; lateral ocelli widely spaced, distance between lateral ocelli about the distance between each lateral ocellus and eye; postclypeus dorsal length as long as or longer than dorsal vertex; postclypeus apex rounded in lateral view; postclypeus rounded in transverse cross section below head. Pronotum shorter than mesonotum, generally narrowed anteriorly with oblique lateral pronotal margins (lateral margins parallel in Toulgoetalna ); weakly developed paranota on lateral pronotal collar, adpressed to pronotal sclerites; mid lateral tooth absent. Mesonotum and metanotum lacking auxiliary soundproducing structures (a few genital stridulatory organs in Carineta ); scutellum cruciform. Opercula size species specific, ranging from small, finger-like extension covering part of tympanal cavity to inflated structure extending over anterior abdominal sternites completely concealing tympanal cavity; meracanthus well developed, triangular, not encapsulated completely by operculum. Fore leg femora with three well developed spines and small apical spine; hindcoxae lacking a large inner protuberance. Fore wings and hindwings generally hyaline, partially semi-opaque, infuscated or bronzed in some species. Fore wing pterostigma present; costal vein about twice the width of and contiguous to radius + subcostal vein; radius anterior vein 1 diverging from subcostal vein at distal node; fore wing median and cubitus anterior veins unfused and widely separated at basal cell; fore wing cubitus posterior and anal vein 1 partially fused; fore wing cubitus anterior vein 1 divided by mediocubital crossvein so that proximal section shortest. Hindwing distal end of cubital cell 1 wider than distal end of cubital cell 2, ranging from slightly longer to as much as five times longer; hindwing radius posterior and median veins fused at their bases. Male abdominal tergites with sides convex in cross section; anterior tergites parallel (male tergites 2–4 significantly constricted in Toulgoetalna ); tergites 4–7 tapering to genitalia; epipleurites not reflexed to ventral surface. Timbals extend below level of wing bases; timbal covers absent, partially turned back ridge on posterior timbal cavity in some Carineta . Pygofer dorsal beak present; distal shoulder undeveloped; upper lobe present; basal lobe well developed, basal lobe appendages can be elaborate in some species. Uncus absent; claspers present, may be well developed, claspers may meet posterior to the anal tube giving the appearance of an uncus with a median suture. Aedeagus with simple theca, restrained by claspers; conjunctival claws absent; pseudoparameres absent.

Distinguishing features. Members of the Carinetini can be distinguished from the remaining Cicadettinae tribes by having a combination of the following characteristics: a head not as wide as the lateral angles of the pronotal collar, the supra-antennal plate extends about half the distance to the eye, the lack of a ventrally angularly swollen postclypeus, pronotal margins that are not laterally ampliate, cruciform elevation not very narrow but of normal proportion, the fore wing costal margin is not widest at the node, fore wing radius anterior 1 vein not diverging distally from subcosta, fore wing hind margin not very narrow but of normal proportion, subapical portion of male opercula not enlarged towards the body, the abdominal length is about the same as the distance between the apex of the postclypeus and the posterior cruciform elevation, the male abdomen with parallel sides at the base, is not inflated, does not have a circular crosssection, and lacks a dorsal crest, timbals that extend below the wing bases, the presence of a dorsal beak, the anal tube lacks lateral lobes, the upper pygofer lobe is not superimposed over the basal pygofer lobe and not positioned on the distal half of the pygofer, the basal pygofer lobe does not originate in the distal half of the pygofer, the presence of ornamented basal pygofer appendages, the theca lack pseudoparameres, and the aedeagus that is not S-shaped, not apically bifurcated, or possessing several apical and subapical processes or conjunctival claws.

Distribution. With the reassignment of the South African genus above, the tribe includes only Neotropical genera with species distributed from Mexico in the north to Argentina, Chile, and Uruguay in the south with additional representatives in both the Greater and Lesser Antilles of the Caribbean. Species of Carinetini have been reported from Argentina, Belize, Bolivia, Brazil, Chile, Colombia, Costa Rica, Cuba, Ecuador, El Salvador, French Guiana, Guatemala, Honduras, Martinique, Mexico, Nicaragua, Panama, Paraguay, Peru, Trinidad, Uruguay, and Venezuela (Goding 1925; Boulard 1982; Sanborn 2001, 2006, 2007, 2009, 2010, 2011b, 2014a, 2014b, 2018, 2019a, 2020a, b, c, d, e, 2021b, 2023, 2024a, 2024b; Sanborn & Heath 2014; Sanborn & Maes 2012; Nunes et al. 2023). A lack of collection effort is probably responsible for the only two South American countries (Guyana and Suriname) where representatives of the tribe have not been reported since Carinetini are well represented in neighboring countries (e.g. Sanborn 2020d, f, 2023; Nunes et al. 2023).

Key to the genera of Carinetini Distant, 1905a . The remaining genera comprising the Carinetini include some of the most species rich genera in the New World as well as some monospecific genera.

1 Fore wing with nine apical cells................................................................................................................. Novemcella Goding, 1925

- Fore wing with eight apical cells........................................................................................................................................................2

2 Pronotum with parallel lateral sides, male abdomen significantly constricted in segments 2–4 ............ Toulgoetalna Boulard, 1982

- Pronotum with lateral sides narrowing anteriorly, anterior abdominal segments with parallel sides ................................................3

3 Head as wide as or wider than mesonotum ............................................................................................... Dorachosa Distant, 1892b

- Head narrower than mesonotum.........................................................................................................................................................4

4 Fore wing not longer than body.............................................................................................................. Guaranisaria Distant, 1905e

- Fore wing longer than body................................................................................................................................................................5

5 Postclypeus extending anteriorly well beyond supra-antennal plates, postclypeus distinctly visible from anterior head when viewed from the dorsal side, fore wing cubitus anterior vein arising from about three quarters distant on the basal cell, basal cell distinctly longer than broad.............................................................................................................. Carineta Amyot & Audinet-Serville, 1843

- Postclypeus not extending anteriorly, head smoothly curved anteriorly when viewed from the dorsal side, fore wing cubitus anterior vein arising from the middle of the basal cell, basal cell a little longer than broad........................ Ahomana Distant, 1905c