Description of Tonnoira conistylus sp. nov. from Costa Rica and a new record of Tonnoira distincta Bravo et al. 2008 from Ecuador

ABSTRACT Tonnoira conistylus sp. nov. is described based on a male collected in 1921 from Costa Rica. Illustrations and photographs of the new species are provided. This species is included in the world key of males of the genus Tonnoira Enderlein and an updated distributional map for the species in the genus is provided, as well as a new record of Tonnoira distincta from Ecuador. http://www.zoobank.org/urn:lsid:zoobank.org:pub:FC930575-34FB-4CEE-A456-773F0083B7C5


Introduction
Tonnoira ;Enderlein, 1937 is a Neotropical genus of moth flies (Diptera: Psychodidae) with 26 extant and one fossil species described so far (Azar & Maksoud 2020;Bravo et al. 2020).The genus was first described by Enderlein (1937) based on a single female specimen of T. pelliticornis Enderlein, 1937 from Peru (Santos & Curler 2014), but currently, the extant Tonnoira species have been recorded in ten countries ranging from Nicaragua to Brazil, excluding the Caribbean Islands (Bravo et al. 2008(Bravo et al. , 2020;;Santos & Curler 2014).
Brazil is the country with the highest number of Tonnoira species, with records from 18 out of the 26 described species so far, followed by Suriname (5 species), French Guiana (3 species), Costa Rica, Panama, and Venezuela (2 species), and Bolivia, Ecuador, Nicaragua, and Peru (1 species) (Bravo et al. 2020).
In the present study, a new species is described based on a single male collected in Costa Rica in 1921.Additionally, a new geographical record for Tonnoira distincta Bravo, Alves & Chagas, 2008 from Ecuador is provided, a checklist and an updated distributional map for the species of the genus is provided.

Material and methods
The material examined for this study is deposited at The National History Museum (NHM) in London, UK.Collector, collection method, date, and locality are specified under the material examined after each specimen.In the material examined section, at the end of each record and between square brackets ([]), the holding institution is indicated.In the description of type labels, the contents of each label are enclosed in double quotation marks (' '), italics denote handwriting, and the individual lines of data are separated by a double forward-slash (//).
The specimens were diaphanized, dissected, and mounted on permanent slides.Slides are thought to be prepared by Laurence W. Quate (G.R. Curler, pers. comm. 2021) based on the type of preparation and notes on slides.

Measurements
Head width was taken at the widest part, approximately above the insertion of the antennal scape, whereas the length was taken from the vertex to the lower margin of the clypeus; wing length was measured from the base of the wing at the start of the costal node to the apex of the wingtip, while the width was taken approximately at an imaginary vertical line crossing the radial and medial forks; palpal proportions are given considering the length of the first palpal segment as a unit (1.0).

Genus Tonnoira Enderlein, 1937
Type species: Tonnoira pelliticornis, 1937: 106.Type locality: PERU, Callanga.Bravo et al. 2020: 4 (species list).The distribution of all species can be seen in Figure 1 and Table 1.Santos & Curler, 2014;T. bifida Bravo & Chagas, 2004;T. bifurcata Quate & Brown, 2004;T. bitalea Quate, 1999;T. bitenacula Quate, 1996 Azar &Maksoud, 2020. Azar andMaksoud (2020) present no strong evidence to place Tonnoira sakhalinensis in the genus Tonnoira, the description provided is consistent with placement in several tribes, the wing, on one hand, is consistent with Brunettiini and Maruinini, on the other hand, the genitalia looks more Brunettiini-like rather than Maruinini.The extraordinary number of tenacula present in this species (six compared to 1-3 in extant species) makes it difficult to place it in Tonnoira, therefore I consider this species as incertae sedis.

Differential diagnosis
Tonnoira conistylus sp.nov.can be easily differentiated from the other species in the genus by the combination of the following characters: first flagellomere cylindrical; gonostylus not bifurcate; surstyli with two tenacula; aedeagus straight and with parameres present.This species is similar to Tonnoira cavernicola but it can be easily recognized by the conical shape of the gonostylus in T. conistylus sp.nov.(tapered and curved in T. cavernicola) and the aedeagus with two parameres in T. conistylus sp.nov.(one paramere in T. cavernicola).

Description
Male.Head.Wider than long; eye bridge of four rows of facets; eyes separation resembling a sand-watch, with the upper part separated by less than one facet diameter and almost joined at the waisted part; interocular suture as an inverted 'Y,' short, length about 1.5 facet diameters; the frontal patch of alveoli trapezoidal slightly concave in upper and lower margins; palpus incomplete in the material examined, first segment of the palpus almost spherical; labium without strong sclerite; labella bulbous with two rows of setae on apical margin.Antennal scape 1.6 times the length of the pedicel, almost cylindrical, apical margin wider than basal margin; pedicel spherical, smaller than scape; flagellomeres cylindrical, about three times the length of the pedicel, with scattered setae on the surface, apical flagellomeres absent in examined material; ascoids missing in reviewed material.
Wing.Length two times its width; wing membrane hyaline; junction of R 2+3 basal to M 1+2 , 'stem' of R 2+3 very short; R 5 ending at wing apex; CuA 2 with an almost 90° bend at the apex.
Terminalia (Figures 2C-E, 3).Gonocoxites are 1.2 times the length of gonostylus.Gonostylus covered with scattered alveoli, conical with rounded apex.Aedeagus with basiphallus longer than gonocoxite.Epandrium about width 1.4 times its length.Hypoproct triangular, not very prominent, covered in small setulae.Surstyli tapered from base to apex, curved dorsally, each with two tenacula, one inserted dorsoapically and one inserted dorsolaterally, separated by about the length of one tenaculum; apical tenacula with crow-like apex; dorsal tenacula spoonlike apex.

Female. Unknown.
Etymology.From the combination of Latin 'cōnus' (meaning cone) and surstylus, referencing the conical shape of the surstylus.Species name to be treated as a name in apposition.
Distribution.Only known from the type locality in Costa Rica.
Remarks.Tonnoira conistylus sp.nov.can be placed in the key provided by Bravo et al. (2020) by modifying it as follows: 24. Eyes separated by one facet diameter; aedeagus extending to half of gonostylus (see Bravo et al. 2008: fig. 6)  Remarks.One of the males has the paramere bent toward the coverglass, giving it the appearance of not having a paramere and difficulting the visualization.The position of the aedeagus makes it look slightly different from the other male examined but, nonetheless, based on the locality and other characters compared favorably to the other specimen and the illustrations by Bravo et al. (2020); I have no doubts that both belong to the same species and that they can be positively identified as T. distincta.

Discussion
To date, only three species of Tonnoira have been collected in caves, namely: T. cavernicola, T. distincta, and T. robusta.The new report of T. distincta for Ecuador was also collected in a cave.Although the association of Tonnoira species with caves is poorly studied this new record collected in a similar environment as the type series in a different country leads me to believe that there is still a high amount of understudied interactions inside the Psychodinae subfamily in their natural environments.

Figure 1 .
Figure 1.Updated distribution map of known Tonnoira species.