New species and new records of Tonnoira (Diptera: Psychodidae) from Brazil

Freddy Bravo Naiara Vilarinho Maíra Xavier Araújo About the authors

Abstract

A new species of moth-fly, Tonnoira igrapiunensis sp. nov. (Psychodidae, Psychodinae), from the Brazilian Atlantic Rain Forest is described. The new species can be distinguished from other species of the genus by its eye bridge separated by 0.5 facet diameter, R5 ending slightly posterior to apex of the wing, gonostyli bifurcated with long arm shorter than length of gonocoxites, aedeagus bipartite and asymmetrical, and two asymmetrical parameres. The gonostylus is similar to that of Tonnoira andradeiSantos, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470., but in the latter species the aedeagus has only one shaft and not two as in the new species. Tonnoira bifidaBravo & Chagas, 2004Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605., Tonnoira brisolaiiSantos, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470., and Tonnoira galatiaeSantos, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470. have their distribution expanded.

Key-Words.
Psychodinae; Maruinini; Moth fly; Atlantic Rain Forest; Neotropical Region

INTRODUCTION

The American genus Tonnoira Enderlein, 1937 (Psychodinae) has been recorded from Nicaragua to Brazil (Espírito Santo State), with no records for the Caribbean islands. Twenty-five species are currently known for the genus (Bravo et al., 2008Bravo, F.; Chagas, C. & Alves, V.R. 2008. Description of two new species of Tonnoira Enderlein from caves in the Brazilian Amazon and comments about the taxonomic status of Tonnoira plumaria Quate (Diptera, Psychodidae, Psychodinae). Zootaxa, 1916: 63-68.; Chagas-Vieira, 2012Chagas-Vieira, C. 2012. New records of Tonnoira Enderlein (Diptera, Psychodidae, Psychodinae) from the Brazilian Amazon with descriptions of two new species. Zootaxa, 3318: 51-56.; Santos & Curler, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.). There are records of seventeen Tonnoira species in Brazil, which represents 70% of all known species in that genus (Bravo & Chagas, 2004Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605.; Bravo et al., 2008Bravo, F.; Chagas, C. & Alves, V.R. 2008. Description of two new species of Tonnoira Enderlein from caves in the Brazilian Amazon and comments about the taxonomic status of Tonnoira plumaria Quate (Diptera, Psychodidae, Psychodinae). Zootaxa, 1916: 63-68.; Quate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117.; Chagas-Vieira, 2012Chagas-Vieira, C. 2012. New records of Tonnoira Enderlein (Diptera, Psychodidae, Psychodinae) from the Brazilian Amazon with descriptions of two new species. Zootaxa, 3318: 51-56.; Santos & Curler, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.). We describe here a new species of Tonnoira from the Atlantic Rain Forest of Bahia State, Brazil, and expand the distributions of three previously described species.

MATERIAL AND METHODS

The specimens described here were collected by Dr. Sergio Andena and collaborators using Malaise and Pennsylvania light traps during an insect inventory in the Reserva Ecológica da Michelin (Michelin Ecological Reserve, 13°46′53″S; 38°49′44″W), which is located in the Central Corridor of the Atlantic Rain Forest, Bahia State. The Psychodidae specimens were collected in one of the most intact forest fragments in that reserve, called Vila 5, which covers 180 ha at elevations between 160 and 288 m a.s.l. (Fig. 1) (Nascimento et al., 2017Nascimento, F.E.L.; Botero, J.P.; Aragão, M. & Andena, S.R. 2017. Faunistic analysis of Cerambycidae (Insecta: Coleoptera) in an area of Atlantic Forest. Journal of Natural History, 51: 2429-2441.).

Figure 1
Atlantic Rain Forest in Reserva Ecológica Michellin, Vila 5. Photo by M. Aragão.

The specimens examined in this study were cleared with 10% potassium hydroxide (KOH), neutralized with acetic acid 20%, dehydrated in ethanol, diaphanized in clove oil and mounted in Canada balsam on slides. General morphological terminology follows Cumming & Wood (2017Cumming, J.M. & Wood, D.M. 2017. 3. Adult morphology and terminology, pp. 51-63. In: Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds.). Manual of Afrotropical Diptera. Vol. 1. Introductory chapters and keys to Diptera families. Pretoria, South African National Biodiversity Institute. 1361p.). Cercus is preferred to the apical appendages of male terminalia following Araújo & Bravo (2019Araújo, M.X. & Bravo, F. 2019. Two new species of Lepidiella Enderlein, 1937 (Diptera: Psychodidae) from the Neotropical Region with taxonomic comments about the species of the genus. Zootaxa, 4551: 487-493.). Additional information not included in the collection label was added in brackets in material examined. The type specimens and other material studied are deposited in the Entomological Collection Prof. Johann Becker of the Museu de Zoologia da Universidade Estadual de Feira de Santana, Bahia, Brazil (MZFS).

TAXONOMY

Genus Tonnoira Enderlein, 1937

Type species:Tonnoira pelliticornis Enderlein (orig. des.), known only by female.

Comments:Santos & Curler (2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.) provide an accurate diagnosis of Tonnoira. In some species of Tonnoira the R5 ends at pointed apex but other species this vein ends slightly beyond the apex, thus, these character states should be included in the diagnosis.

Tonnoira igrapiunensis sp. nov.

Figs. 2-12

Figures 2-12
Tonnoira igrapiunensis sp. nov., adult male. (2) Head in frontal view (paratype 56227). (3) scape, pedicel and first flagellomeres (paratype 56227). (4) Flagellomeres 4, 5 and 6 (paratype 56227). (5) Apical flagellomeres and apiculus (paratype 56226). (6) pedicel with the marginal sensory setae (paratype 56227). (7) Palpus (paratype 56227). (8) wing. (9) Hypandrium, gonocoxites, gonostylus, ejaculatory apodeme, aedeagus and parameres (holotype). (10) gonostylus, lateral view (paratype 56227). (11) gonocoxites, gonocoxal apodemes fused, parameres and ejaculatory apodeme (holotype 56225). (12) Epandrium, cerci and hypoproct (paratype 56227). Abbreviations: am = apical membrane; lsae = long shaft of aedeagus; pr = paramere; ssae = small shaft of aedeagus.

Type material: Holotype: male, BRAZIL, Bahia, Igrapiúna, Reserva Ecológica Michelin, Vila 5, 28.v-12.vii.2012, Malaise [trap]. Aragão, M.; Menezes, E.; Mota, E.; Andena, S, leg. (MZFS #56225). Paratypes: 6 males, same data as holotype (MZFS #56227, #56229, #56230, #56231, #56232, #56233); 2 males, same locality as holotype, 27-28.x. 2012, light [trap], Aragão, M., Menezes, E., Mota, E. leg. (MZFS #56226, #56234); 1 male, same locality as holotype, 28.x-18.xi.2012, Malaise [trap], Aragão, M., Menezes, E. leg. (MZFS #56228).

Description: Male: Head: slightly wider than long; short vertex; eye bridge with four facet rows; eyes separated by almost one facet diameter; interocular suture inverted Y-shaped (Fig. 2). Antenna: scape wider apically than at base, 2.0 times the length of the pedicel (Fig. 3); pedicel spherical, with two groups of three small sensorial setae both in front of and behind the proximal margin (Fig. 6); flagellum with 14 flagellomeres, all cylindrical (Figs. 3-5); ascoids not observed, probably lost; 14th flagellomere with long apiculus, 0.3 times the length of the flagellomere (Fig. 5). Wing (Fig. 8): Sc short not reaching C; R2+3 not connected to R4; R5 ending slightly beyond the wing apex; radial fork basal to medial fork. Terminalia: hypandrium bent backwards, narrower laterally (Fig. 9); gonocoxites 1.2 times the length of gonostyli, with setae only dorsally (Figs. 9, 11); gonostyli bifurcate; lateral branch of gonostylus longer than mesal branch (Figs. 9, 10), divergent in lateral view (Fig. 10); aedeagus bipartite, asymmetrical, short shaft 0.8 times the length of the long shaft; short shaft with two sclerites fused basally, dorsal sclerite plate-like, shorter than the narrow ventral sclerite, both covered by an apical membrane (Fig. 9); parameres asymmetrical, long shaft with curved apex and short shaft plate-like with rounded apex (Figs. 9, 11); gonocoxal apodemes fused medially and articulated with parameres (Fig. 11); epandrium wider than long, bare ventrally, with one basal aperture (Fig. 12); hypoproct longer than epandrium, with apex tapered; epiproct shorter than hypoproct, with rounded apex; hypoproct and epiproct both with apical micropilosity (Fig. 12); cerci with two tenacula (Fig. 12).

Female: Unknown.

Remarks: The new species can be distinguished from other species of Tonnoira by the eye bridge with four facet rows, separated by 0.5 facet diameter; R5 ending slightly posterior to apex of wing; gonostyli bifurcated, dorsal arm shorter than ventral arm; long arm of gonostyli shorter than length of gonocoxites; cerci with two tenacula; aedeagus bipartite, asymmetrical, short shaft 0.8 times the length of the long shaft; parameres asymmetrical, longer shaft with apex curved. The gonostylus is similar to that of T. andradeiSantos, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470., but differs, mainly, by the bifid aedeagus in the new species and with only one shaft in T. andradei. The ejaculatory apodeme and hypoproct in the new species is longer than the same structures in T. andradei.

Etymology: The species name is a reference to the municipality of Igrapiúna where the specimens of the new species were collected.

Distribution: Known only from the type locality.

Tonnoira bifida Bravo & Chagas, 2004Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605.

Tonnoira bifidaBravo & Chagas, 2004Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605.: 601, figs. 1-9.

Material examined: 1 male, BRAZIL, Bahia, Varzedo, Sr. Getúlio, x.2016, Dias & Campos, leg. (MZFS).

Distribution: Brazil: Bahia, Serra da Jibóia (type locality: 12°52′S, 39°28′W, 720 m.a.s.l.); Bahia, Varzedo (new record, 12°57′45.1″S, 39°27′12.1″W, 280 m.a.s.l.).

Tonnoira brisolaii Santos, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.

Tonnoira brisolaiiSantos, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470. (in Santos & Curler, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.): 464, figs. 1-5.

Material examined: 1 male, BRAZIL, Bahia, Serra da Jibóia, Sede Gambá, 10.v.2017, Ligth [trap], Silva-Neto, Mendes & Moura, leg. (MZFS).

Distribuição: Brazil: Espírito Santo, Santa Teresa, (type locality); Bahia, Serra da Jibóia (new record, 12°52′S, 39°28′W, 720 m.a.s.l.).

Tonnoira galatiae Santos, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.

Tonnoira galatiaeSantos, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470. (in Santos & Curler, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.): 468, figs. 14-17.

Material examined: 3 males, BRAZIL, Bahia, Igrapiuna, Reserva Ecológica Michelin, Vila 5, 22.ix-28.x.2012, Malaise [trap]. Aragão, M.; Menezes, E. leg. (MZFS); 1 male same data as the previous one, except 16.xii.2012-20.i.2013 (MZFS); 2 males, same data as the previous one, except 20.i-24.ii.2013 (MZFS); 2 males, same data as the previous one, except 24.ii-31.iii.2013 (MZFS); 1 male, same data as the previous one, except 31.iii-28.iv.2013 (MZFS).

Distribution: Brazil: Espírito Santo, Santa Teresa (type locality: 19°54′S, 40°33′W, 820 m.a.s.l.); Bahia, Igrapiúna (new record: 13°50′S; 39°10′W, 160 m.a.s.l.).

Species included in Tonnoira

Twenty-six species are recognized in Tonnoira: T. andradeiSantos, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470. (Brazil: Espírito Santo); T. bifidaBravo & Chagas, 2004Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605. (Brazil: Bahia); T. bifurcataQuate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117. (Brazil: Rondônia); T. bitaleaQuate, 1999Quate, L. W. 1999. Taxonomy of neotropical Psychodidae (Diptera 3. Psychodines of Barro Colorado Island and San Blas, Panama. Memoirs on Entomology International, 14: 409-441. (Panama, Brazil: Amazonas); T. bitenaculaQuate, 1996Quate, L.W. 1996. Preliminary taxonomy of Costa Rican Psychodidae (Diptera), exclusive of Phlebotominae. Revista de Biología Tropical, 44(Suppl. 1): 1-81. (Costa Rica); T. brisolaiiSantos, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470. (Brazil: Espírito Santo); T. castaneaQuate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117. (Brazil: Amazonas, Suriname); T. cavernicolaQuate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117. (Bolivia, Guiana Francesa); T. didymaQuate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117. (French Guiana, Suriname); T. distinctaBravo, Alves & Chagas, 2008Bravo, F.; Chagas, C. & Alves, V.R. 2008. Description of two new species of Tonnoira Enderlein from caves in the Brazilian Amazon and comments about the taxonomic status of Tonnoira plumaria Quate (Diptera, Psychodidae, Psychodinae). Zootaxa, 1916: 63-68. (Brazil: Amazonas); T. ferreiraiSantos, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470. (Brazil: Espirito Santo); T. fusiformisQuate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117. (Costa Rica, Ecuador); T. galatiaeSantos, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470. (Brazil: Espírito Santo); T. igrapiunensis sp. nov. (Brazil: Bahia); T. longipennisBravo & Chagas, 2004Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605. (Brazil: Bahia); T. magnaBravo & Chagas, 2004Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605. (Brazil: Bahia); T. mirabilisWagner, 1981Wagner, R. 1981. Two new moth-flies (Diptera, Psychodidae) from South America. Studies on Neotropical Fauna and Environment, 16: 217-220. (Brazil: Amazonas, Suriname); T. pelliticornis (Peru); T. protuberataQuate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117. (Venezuela); T. psacadopteraQuate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117. (Venezuela); T. rapiformisQuate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117. (Brazil, Suriname); T. rectilataQuate, 1999Quate, L. W. 1999. Taxonomy of neotropical Psychodidae (Diptera 3. Psychodines of Barro Colorado Island and San Blas, Panama. Memoirs on Entomology International, 14: 409-441. (Nicaragua, Panama, Suriname, Brazil: Amazonas, Rondônia); T. robustaBravo, Alves & Chagas, 2008Bravo, F.; Chagas, C. & Alves, V.R. 2008. Description of two new species of Tonnoira Enderlein from caves in the Brazilian Amazon and comments about the taxonomic status of Tonnoira plumaria Quate (Diptera, Psychodidae, Psychodinae). Zootaxa, 1916: 63-68. (Brazil: Amazonas); T. sicilisQuate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117. (French Guiana); T. spinaChagas-Vieira, 2012Chagas-Vieira, C. 2012. New records of Tonnoira Enderlein (Diptera, Psychodidae, Psychodinae) from the Brazilian Amazon with descriptions of two new species. Zootaxa, 3318: 51-56. (Brazil: Amazonas); T. tripenisChagas-Vieira, 2012Chagas-Vieira, C. 2012. New records of Tonnoira Enderlein (Diptera, Psychodidae, Psychodinae) from the Brazilian Amazon with descriptions of two new species. Zootaxa, 3318: 51-56. (Brazil: Amazonas).

In Brazil, eighteen species of Tonnoira are recorded only from the Amazon and Atlantic Rain forests, two regions with high annual rainfall levels. Ten species occur in the Brazilian Amazon (states of Amazonas, Para, and Rondônia), and eight in the Atlantic Rain Forest (states of Espírito Santo and Bahia).

Key to males of Tonnoira

The male of the type species of the genus, T. pelliticornis, is unknown.

1. Gonostylus bifurcate 2

- Gonostylus simple, not bifurcate 9

2. Gonostylus with lateral and mesal branches of different length; branches subparallel or divergent 3

- Gonostylus with lateral and mesal branches of same length, crossed (Santos & Curler, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.: Fig. 4) T. brisolai

3. Lateral branch of gonostylus longer than mesal branch; mesal branch of gonostylus shorter than the length of gonocoxite (e.g., Santos & Curler, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.: fig. 12) 4

- Lateral branch of gonostylus shorter than mesal branch; mesal branch of gonostylus longer than the length of gonocoxite (e.g., Bravo & Chagas, 2004Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605.: fig. 6) 6

4. Eye bridge with 5 facet rows; cercus with one tenaculum T. protuberata

- Eye bridge with 4 facet rows; cercus with two tenacula 5

5. Eyes separated by 0.5 facet diameter; parameres symmetrical (Santos & Curler, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.: fig. 12) T. andradei

- Eyes separated by almost 1 facet diameter; parameres asymmetrical (fig. 9) T. igrapiunensis sp. nov.

6. Lateral branch of gonostylus very short, 0.2 times the length of mesal branch (Santos & Curler, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.: fig. 8) T. ferrerai

- Lateral branch of gonostylus 0.4-0.6 times the length of mesal branch (e.g., Bravo & Chagas, 2004Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605.: fig. 6) 7

7. Aedeagus with one branch apically U-shaped with one of the branchs longer than other (Quate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117.: fig. 69a) T. bifurcata

- Aedeagus with two branches, none of them apically U-shaped 8

8. Lateral branch of gonostylus with the same sclerotinization of mesal branch; hypandrium narrower than width of gonocoxite, strip-like (Quate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117.: fig. 70) T. didyma

- Lateral branch strongly sclerotinized; hypandrium wider than width of gonostylus (Bravo & Chagas, 2004Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605.: fig. 6) T. bifida

9. Cercus with 1 tenaculum 10

- Cercus with 2 or 3 tenacula 13

10. Base of R3 with cluster of black “granules” (Quate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117.: fig. 63) T. psacadoptera

- Base of R3 normal, withou cluster of black “granules” 11

11. Hypandrium wide, of the same length of width of gonostylus; gonocoxite 0.8 times the length of gonostylus (Chagas-Vieira, 2012Chagas-Vieira, C. 2012. New records of Tonnoira Enderlein (Diptera, Psychodidae, Psychodinae) from the Brazilian Amazon with descriptions of two new species. Zootaxa, 3318: 51-56.: fig. 16) T. tripenis

- Hypandrium stripe-like; gonocoxite about as long as gonostylus 12

12. Aedeagus “tripartite with short spur, a sickle shaped lateral shaft, and broad, paddle shaped central part twisted” (Quate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117.: p. 29, fig. 68) T. sicilis

- Aedeagus “bipartite, larger shaft with basal three-quarters broad, asymmetrically narrowing at distal one-quarter, smaller shaft small, blunt, about one-half as long as other shaft” (Quate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117.: p. 27, fig. 67a) T. castanea

13. Cercus with 3 tenacula 14

- Cercus with 2 tenacula 15

14. Aedeagus ending at the same level of apex of gonostylus; ejaculatory apodeme shorter than gonostylus (Wagner, 1981Wagner, R. 1981. Two new moth-flies (Diptera, Psychodidae) from South America. Studies on Neotropical Fauna and Environment, 16: 217-220.: fig. 3) T. mirabilis

- Aedeagus ending above of apex of gonostylus; ejaculatory apodeme longer than gonostylus (Chagas-Vieira, 2012Chagas-Vieira, C. 2012. New records of Tonnoira Enderlein (Diptera, Psychodidae, Psychodinae) from the Brazilian Amazon with descriptions of two new species. Zootaxa, 3318: 51-56.: fig. 12) T. spina

15. Aedeagus symmetrical, base broad, apex sagittate; parameres absent (Quate, 1996Quate, L.W. 1996. Preliminary taxonomy of Costa Rican Psychodidae (Diptera), exclusive of Phlebotominae. Revista de Biología Tropical, 44(Suppl. 1): 1-81.: fig. 13b; Quate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117.: fig. 71) 16

- Aedeagus symmetrical or asymmetrical, long, subcylindrical, pointed or rounded at apex; parameres present 17

16. Hypandrium present, strip-like; gonocoxite as wide as long (Quate, 1996Quate, L.W. 1996. Preliminary taxonomy of Costa Rican Psychodidae (Diptera), exclusive of Phlebotominae. Revista de Biología Tropical, 44(Suppl. 1): 1-81.: fig. 13b) T. bitenacula

- Hypandrium vestigial, broken in the middle; gonocoxite longer than wide (Quate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117.: fig. 71) T. rapiformis

17. Aedeagus ending far beyond the apex of gonostylus (Bravo & Chagas, 2004Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605.: fig. 1) T. longipennis

- Aedeagus ending near or just beyond the apex of gonocoxites, never beyond the apex of gonostylus 18

18. Aedeagus sickle shaped; ejaculatory apodeme triangular, short (Quate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117.: fig. 77) T. rectilata

- Aedegus straight or little curved; ejaculatory apodeme sub-oval, longer than wide 19

19. First flagellomere fusiform 20

- First flagellomere cylindrical 23

20. “Gonocoxal apodeme with pair of slender, acute processes flanking aedeagus and extending nearly to the tip of aedeagus” (Quate, 1999Quate, L. W. 1999. Taxonomy of neotropical Psychodidae (Diptera 3. Psychodines of Barro Colorado Island and San Blas, Panama. Memoirs on Entomology International, 14: 409-441.: p. 429, fig. 5K) T. bitalea

- Gonocoxal apodeme not expanded posteriorly 21

21. Hypandrium, wide, posteriorly bilobed (Bravo & Chagas, 2004Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605.: fig. 23) T. magna

- Hypandrium, narrow, without lobes 22

22. Eyes separated by 0.5 facet diameters; gonostylus shorter than gonocoxites; ejaculatory apodeme shorter than gonocoxite (Bravo et al., 2008Bravo, F.; Chagas, C. & Alves, V.R. 2008. Description of two new species of Tonnoira Enderlein from caves in the Brazilian Amazon and comments about the taxonomic status of Tonnoira plumaria Quate (Diptera, Psychodidae, Psychodinae). Zootaxa, 1916: 63-68.: figs. 11, 16) T. robusta

- Eyes separated by 0.2 facet diameters; gonostylus longer than gonocoxites; ejaculatory apodeme longer than gonocoxite (Santos & Curler, 2014Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.: fig. 16) T. galatiae

23. Aedeagus with one shaft, “abruptly narrowing to acute apex” (Quate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117.: p. 32, fig. 74) T. fusiformis

- Aedeagus with one shaft evenly narrowing in acute apex 24

24. Eyes separated by less than 1 facet diameter; “aedeagus extending little beyond tip of gonocoxite” (Quate & Brown, 2004Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117.: p. 31, fig. 72) T. cavernicola

- Eyes separated by 1 facet diameter; aedeagus extending to half of gonostylus (Bravo et al., 2008Bravo, F.; Chagas, C. & Alves, V.R. 2008. Description of two new species of Tonnoira Enderlein from caves in the Brazilian Amazon and comments about the taxonomic status of Tonnoira plumaria Quate (Diptera, Psychodidae, Psychodinae). Zootaxa, 1916: 63-68.: fig. 6) T. distincta

ACKNOWLEDGEMENTS

This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001. Freddy Bravo received a research grant from Conselho Nacional de Desenvolvimento Científico e Tecnológico (№ 306441/2015-2). Maíra Xavier Araújo also has a grant from CNPq (№ 150571/2018-6). We would like to thank the reviewers for their comments in the manuscript.

REFERENCES

  • Araújo, M.X. & Bravo, F. 2019. Two new species of Lepidiella Enderlein, 1937 (Diptera: Psychodidae) from the Neotropical Region with taxonomic comments about the species of the genus. Zootaxa, 4551: 487-493.
  • Bravo F. & Chagas, C. 2004. Espécies Novas de Tonnoira Enderlein (Diptera: Psychodidae) do Nordeste brasileiro. Neotropical Entomology, 33: 601-605.
  • Bravo, F.; Chagas, C. & Alves, V.R. 2008. Description of two new species of Tonnoira Enderlein from caves in the Brazilian Amazon and comments about the taxonomic status of Tonnoira plumaria Quate (Diptera, Psychodidae, Psychodinae). Zootaxa, 1916: 63-68.
  • Chagas-Vieira, C. 2012. New records of Tonnoira Enderlein (Diptera, Psychodidae, Psychodinae) from the Brazilian Amazon with descriptions of two new species. Zootaxa, 3318: 51-56.
  • Cumming, J.M. & Wood, D.M. 2017. 3. Adult morphology and terminology, pp. 51-63. In: Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds.). Manual of Afrotropical Diptera. Vol. 1. Introductory chapters and keys to Diptera families. Pretoria, South African National Biodiversity Institute. 1361p.
  • Nascimento, F.E.L.; Botero, J.P.; Aragão, M. & Andena, S.R. 2017. Faunistic analysis of Cerambycidae (Insecta: Coleoptera) in an area of Atlantic Forest. Journal of Natural History, 51: 2429-2441.
  • Quate, L.W. 1996. Preliminary taxonomy of Costa Rican Psychodidae (Diptera), exclusive of Phlebotominae. Revista de Biología Tropical, 44(Suppl. 1): 1-81.
  • Quate, L. W. 1999. Taxonomy of neotropical Psychodidae (Diptera 3. Psychodines of Barro Colorado Island and San Blas, Panama. Memoirs on Entomology International, 14: 409-441.
  • Quate, L.W. & Brown, B.V. 2004. Revision of Neotropical Setomimini (Diptera: Psychodidae: Psychodinae). Contribution in Science, Natural History Museum of Los Angeles, 500: 1-117.
  • Santos, C.B. & Curler, G.R. 2014. Four new species of Tonnoira Enderlein (Diptera: Psychodidae, Psychodinae) from the Brazilian Atlantic forest. Zootaxa, 3760: 463-470.
  • Wagner, R. 1981. Two new moth-flies (Diptera, Psychodidae) from South America. Studies on Neotropical Fauna and Environment, 16: 217-220.

  • Published with the financial support of the "Programa de Apoio às Publicações Científicas Periódicas da USP"

Publication Dates

  • Publication in this collection
    07 Sept 2020
  • Date of issue
    2020

History

  • Received
    10 Dec 2019
  • Accepted
    20 Feb 2020
  • Published
    08 June 2020
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