(Diptera: Ditomyiidae) from Neotropical Mexico

Zootaxa 4072 (4): 430–440 http://www.mapress.com/j/zt/ Copyright © 2016 Magnolia Press

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http://doi.org/10.11646/zootaxa.4072.4.2 http://zoobank.org/urn:lsid:zoobank.org:pub:599A08C7-ECDC-4D6B-8A97-F41330148447

Three new species of Calliceratomyia Lane (Diptera: Ditomyiidae) from Neotropical Mexico HERON HUERTA1 & DALTON DE SOUZA AMORIM2 1

Laboratorio de Entomología, InDRE, Francisco de P. Miranda No. 177, Colonia Unidad Lomas de Plateros, México D.F., 01480, MEXICO. E-mail: [email protected] 2 Departamento de Biologia, FFCLRP, Universidade de São Paulo, Av. Bandeirantes 3900, 14.040-901 Ribeirão Preto SP BRAZIL. E-mail: [email protected]

Abstract Three new species of the ditomyiid genus Calliceratomyia Lane, so far known only from the type-species from southeastern Brazil, are described from Mexico. The new species—Calliceratomyia papaveroi sp. n., Calliceratomyia pantelhomyia sp. n. and Calliceratomyia zayolli sp. n.—are described based on adult males collected with Malaise traps in the States of Oaxaca, Chiapas and Hidalgo, Mexico. The descriptions are illustrated and the diagnosis of the genus is revised. Key words: Sciaroidea, geographic distribution, Neotropical Region, Malaise trap

Introduction The ditomyiid fungus-gnats have been treated by some authors as a subfamily of Mycetophilidae (e.g., Munroe 1974, Vockeroth 1981), but most authors now give family rank to the group (e.g., Hennig 1973, Papavero 1977, Matile 1990, Amorim & Rindal 2007, Vockeroth 2009). The family presently includes in a group the genera Symmerus Walker and Australosymmerus Freeman, very carefully reviewed by Munroe (1974). It is uncertain if the set of the remaining genera in the family constitute a monophyletic group—including Ditomyia Winnertz (exclusively Holarctic), Asioditomyia Saigusa (known from a single species from Japan), Celebesomyia Saigusa (known from a single Oriental species), Nervijuncta Marshall (with 18 species in New Zealand, one species in Chile, and one species in southern Brazil), Rhipidita Edwards (with two species described from southern Brazil) and Calliceratomyia Lane (with one species from southern Brazil). Ditomyia and Nervijuncta share a strongly modified male gonostylus and may compose a clade in the family. Both Neotropical species of Nervijuncta have been recently redescribed (Falaschi & Amorim 2009). Rhipidita and Calliceratomyia are poorly known genera in the family, with modified antennal flagellomeres and maxillary palpus, and may as well compose a small clade within the ditomyiids. Understanding the connections of Asioditomyia, Celebesomyia, these two clades, and the clade with Symmerus and Australosymmerus depends on a global study of the phylogenetic relationships within the family. Nervijuncta is typically a southern temperate genus, disjunct between New Zealand and Chile/southern Argentina. N. laffooni Lane, from southern Brazil, corresponds to a northeastern extension of this group—as happens with other groups of insects and plants (see, e.g., Oliveira & Amorim 2010). Rhipidita and Calliceratomyia, on the other hand, correspond to typically tropical elements of the family in the Neotropical fauna (Edwards 1940, Lane 1946, 1952, Papavero 1977). As commented by Falaschi & Amorim (2009), a large number of species of ditomyiids belonging to these two genera have been collected with Malaise traps in different areas of the Neotropical Region during the last 30 years, and still await description. The genus Rhipidita currently includes its type species, R. fusca, described by Edwards (1940) from Seara, Nova Teutônia district, State of Santa Catarina, southern Brazil, and R. nigra, described by Lane (1952) from Campos do Jordão, State of São Paulo, southeastern Brazil. Calliceratomyia has a single described species, C. pectinata Lane also from Campos do Jordão (Lane 1946).

430 Accepted by C. Borkent: 7 Jan. 2016; published: 3 Feb. 2016

The family Ditomyiidae is presently known from Mexico by two species of Australosymmerus (Melosymmerus), three species of Australosymmerus (Calosymmerus), and one species of Australosymmerus unassigned to any of its subgenera (Munroe 1974). This paper adds to the family three new species of Calliceratomyia from Mexico—the northern most occurrence of the genus to date—and describes in more detail the morphology of the genus.

Material and methods The holotypes and paratypes were prepared as permanent microscope slide-mountings in Euparal. Explanations of the taxonomic characters used here are found in Søli (1997) and Vockeroth (2009). The holotypes of the species described here are deposited in the Colección de Artrópodos con Importancia Médica (CAIM), México City, Distrito Federal, Mexico and paratypes of two of the species are deposited in the Diptera collection of the Museu de Zoologia da Universidade do São Paulo, Brazil (MZUSP). The wing venation terminology follows Amorim & Rindal (2007). Illustrations were made with a drawing tube attached to a compound microscope. Measurements and ratios were obtained as follows: body length, from front to tip of abdomen; wing length, from the humeral vein to the wing tip; wing width, at the widest point. Antennal length is the combined length of scape, pedicel and flagellum.

Calliceratomyia Lane Calliceratomyia Lane, 1946: 244. Type–species, Calliceratomyia pectinata Lane, 1946 (orig.des.).

Diagnosis. Flagellomeres slightly or strongly laterally compressed, at least with a short ventral extension. Maxillary palpus with three distinctive segments, segment two humpback. Fusion of medial stem to Rs almost as long as free base of M1+2. Gonocoxites fused to each other ventrally and dorsally, forming a ring around aedeagus, distal part of the gonocoxite largely projected beyond syngonocoxite.

Calliceratomyia papaveroi, sp. n. (Figs. 1, 2, 5, 10, 13, 16) Diagnosis. Flagellomeres 2–8 with a short ventral extension (Figs. 1–2). Penultimate flagellomere length about twice width, last flagellomere tiny; all flagellomeres with one ventral and one dorsal seta (Figs. 2, 5). M forking just before origin of R4 (Fig. 10). Extension of male gonocoxites distal to syngonocoxite slightly enlarged midway to apex. Gonostylus at apex almost as wide as its length. Two separate groups of spines apically on inner face of male gonostylus (Figs. 13, 16). Type-material. Holotype male, labeled: HOLOTYPE, Calliceratomyia papaveroi sp. n. México, Oaxaca, Municipio Santo Domingo Petapa, Localidad Río del Sol, Trampa Malaise, 29–30-jul-2009. Cols. Salceda, S. B., Rodríguez, A. J. & Ordoñez, A. GPS: -95° 14’ 13”N; 16° 56’ 55’’W; elevation, 480 m (CAIM). Paratype: 1 male, same data as holotype, slide-mounted (MZUSP). Description. Body length (including the antenna), 5.0 mm (Fig. 1). Head. Head wider than long. Eyes dichoptic, all facets of similar width, length of interommatidial setae close to facet width. Three ocelli present, aligned, of similar size, distance between ocelli slightly smaller than ocellus diameter. Vertex slightly elevated, with setae, transverse suture present, frons bare, face with 8 setae, length of pair of lateral setae close to first palpomere length. Clypeus more or less triangular. Mouthparts considerably reduced, labellum small; premental apodeme Y-shaped. Maxillary palpus longer than labellum, three clearly articulated brown palpomeres. First palpomere humpback, distinctively bulky distally, with 7 socketed setae, entirely covered by sensillae; second palpomere shorter than first, with some sensilla on basal half, projected beyond base of third palpomere, apically with 3 diminute spines, and one socketed setae; third palpomere more or less cylindrical, with some sensilla on basal third, 5 minute apical spines. Antenna (Figs. 1–2, 5) with 15 flagellomeres, 1.87 mm long. Scape and pedicel

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FIGURE 1. Calliceratomyia papaveroi, sp. n., male paratype, habitus. Scale bar, 0.5 mm.

cylindrical, about 1.2 times as long as wide. Flagellomere 1 more or less cylindrical, flagellomeres 2–8 strongly compressed laterally, with a short (but evident) ventral projection, flagellomeres 9–13 less modified, flagellomere 14 cylindrical, elongate, apical flagellomere small, cylindrical (Fig. 5); flagellomeres covered with short fine sensilla-like trichia, with one dorsal and one ventral longer seta on each flagellomere. Thorax. Scutum uniformly brown, evenly setose, no transverse suture; notopleural, supra-alar areas and postalar callus with long bristles; scutellum brown, with a pair of subapical setae. Mediotergite high, bare, dark brown; pleural sclerites brown, anepisternum, katepisternum, mesepimeron, and laterotergite bare. Postpronotum with a pair of long bristles, proepisternum with 4–5 long bristles and three small setae. Haltere brown, slightly longer than first abdominal tergite, pedicel and capitulum covered with short setae. Legs. Uniformly brown, femora light brown, covered with fine trichia, tibial spurs 1:2:2, dark brown. Fore coxa setose on anterior face, mid coxa setose anteriorly at apical

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FIGURES 2–4. Antennae of Calliceratomyia. 2. C. papaveroi, sp. n., male holotype. 3. C. pantelhomyia sp. n., male holotype. 4. C. zayolli, sp. n., male holotype. Scale bar, 0.1 mm. Flag = flagellum; Ped = pedicel, Sc = scape.



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FIGURES 5–7. Antennal flagellomeres 13–15 of Calliceratomyia. 5. C. papaveroi sp. n., male holotype. 6. C. pantelhomyia sp. n., male holotype. 7. C. zayolli sp. n., male holotype. Scale bar, 0.1 mm.

half, hind coxa with an irregular row of setae latero-posteriorly. Femora clothed with fine microtrichia and dense distal setation, forefemur slightly shorter. Foretibia length twice forefemur length. All tibiae densely covered with microtrichia and irregular rows of setae. Foretibia apical spur about 1.9 times length of tibial diameter at apex, mid and hind inner spur slightly shorter than outer spur, longer than foretibial spur. Apical comb of setae present at inner face of hind tibia. All legs with dense microtrichia and with an irregular ventral row of spines. Tarsal claws black, with basal inner teeth. Wing (Fig. 10). Membrane infuscate brown. Length, 3.8 mm; width, 1.4 mm. Membrane covered with abundant microtrichia and macrotrichia. Medial veins less sclerotized than cubital veins, all veins except A1 reaching wing margin; all veins with macrotrichia. Costa long, reaching tip of wing, barely extending beyond tip of R5. Sc short, incomplete, ending free; R1 long, slightly curved at apex, extending about 3/4 of wing length. Base of Rs oblique, slightly sinuose, originated on basal fourth of wing, Rs running considerably far from R1. R4 length approximately a third of Rs length, oblique along most of its length. M1+2 basally fused to Rs, r-m entirely obliterated; medial fork basal to origin of R4. M4 reaching wing margin beyond medial fork. CuA reaching wing margin clearly basal to medial fork. A1 less sclerotized than other posterior veins, incomplete on distal fourth. Abdomen. Dark brown, segment 1 lighter. Segments cylindrical, covered with dark fine setae, tergites and sternites 1 to 6 normally sclerotized, sternite 7 entirely membranous; sternite 8 trapezoid, tergite 8 reduced to a stripe. Male terminalia (Figs. 13, 16). Dark brown, bent downwards. Gonocoxites robust, elongate, setose, fused basally to form a syngonocoxite ring around the aedeagus, anterior margin slightly concave; portion of gonocoxites distal to syngonocoxite slightly enlarged midway to apex. Gonocoxal apodemes forming a wide posterior wall connecting gonocoxites and enclosing aedeagus. Gonostylus robust, densely setose, slight shorter than gonocoxite, as long as width at apex, ornamented apically with two well separated rows of inner spines,


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ventral row with 14–15 blunt, strongly sclerotized spines, dorsal row with 3–4 spines. Aedeagal complex consisting of a simple, straight process with a pair of short lateral branches on anterior end, an opening at apex, ventrally covered by fused parameres. Tergite 9 short, with setation along posterior margin. Cercus lobate, entirely covered with setae. Female. Unknown. Bionomics. Both specimens of the type series were collected with a Malaise trap in a tropical rainforest in the Mixteca region of the state of Oaxaca. Distribution. Mexico (Oaxaca). Etymology. This species is named after the eminent dipterist Dr. Nelson Papavero for his long contribution to dipterology and to systematics in general and as well to the history of dipterology in Mexico.

Calliceratomyia pantelhomyia, sp. n. (Figs. 3, 6, 8, 9, 11, 14, 17, 19) Diagnosis. Flagellomeres 1–12 with a short ventral extension (Fig. 3). Last flagellomere almost as long as penultimate; a single ventral longer seta on flagellomeres 1–4, flagellomere 14 without longer ventral seta (Fig. 6). M forking just opposite to origin of R4 (Fig. 11). Extension of male gonocoxites distal to syngonocoxite more or less cylindrical. Gonostyle slightly longer than width at apex. Two separate groups of spines apically on inner face of male gonostylus, the dorsal row with 3–4 spines (Figs. 14, 17, 19).

FIGURES 8–9. Calliceratomyia pantelhomyia, sp. n., male holotype. 8. Frontolateral view of head. 9. Maxillary palpus. Scale bar, 0.1 mm.

Type material. Holotype male, labeled: HOLOTYPE, Calliceratomyia pantelhomyia sp. n. México, Chiapas, Municipio Pantelho, 6 km al sur, rumbo al poblado de Pantelho, Trampa Malaise, 29–30-jul-2009. Cols. Salceda, S. B. & Rodríguez, A. J. GPS: -92° ’29 55’’N; 16° 59’47”W; elevation, 460 m. (CAIM). Paratypes: 3 males, same data of holotype (1 in CAIM, 2 in MZUSP). Description. Body length (including antenna), 6.0 mm. Head. Head wider than long. Eyes dichoptic, all facets of similar width, length of interommatidial setae close to facet width. Three ocelli present, aligned, of similar size. Vertex slightly elevated, with setae; transverse suture incomplete; frons bare; face with 8–14 setae, length of lateral pair of setae as long as first palpomere. Clypeus more or less triangular. Mouthparts considerably reduced, labellum small; pre-mental apodeme Y-shaped. Maxillary palpus (Figs. 8–9) longer than labellum, with three brown, clearly articulated palpomeres. First palpomere humpback, distinctively bulky distally, with 7–8 socketed setae, entirely NEW MEXICAN SPECIES OF DITOMYIID GENUS CALLICERATOMYIA


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FIGURES 10–12. Wings of Calliceratomyia. 10. C. papaveroi sp. n., male holotype. 11. C. pantelhomyia sp. n., male holotype. 12. C. zayolli sp. n., male holotype. Scale bar, 0.5 mm.

covered by sensillae; second palpomere shorter than first, with sensilla on basal half, 5–6 minute apical spines, and 2–5 socketed setae; third palpomere more or less cylindrical, with some sensilla on basal third and 8 minute apical spines. Antenna (Figs. 3, 6) length, 2.4 mm, with 15 flagellomeres. Scape and pedicel cylindrical, approximately 1.2 times as long as wide. Flagelomeres 1–12 more or less laterally compressed, flagellomeres 14–15 cylindrical, last flagellomere about as long as flagellomere 14 (Fig. 6); flagellomeres covered with sensilla and short fine setae, flagellomeres 1–13 with a dorsal longer seta, only flagellomere 1–4 with a ventral longer seta. Thorax. Scutum with a mesal darker stripe and a pair of additional lateral darker stripes on a lighter background, evenly setose, no


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transverse suture; notopleural, supra-alar area and postalar callus with long bristles; scutellum brown, with three subapical setae. Mediotergite high, bare, dark brown; remaining pleural sclerites brown, anepisternum, katepisternum, mesepimeron, and laterotergite bare. Postpronotum with a pair of long bristles, proepisternum with 2–3 bristles and 4–5 small setae. Haltere brown, slightly longer than first abdominal tergite, pedicel and capitulum covered with short setae. Legs. As in previous species. Wing. (Fig. 11) Membrane infuscated brown. Length, 4.0– 4.3 mm; width, 1.7–1.8 mm (n=2). M1+2 basally fused to Rs, fork slightly basal to level of origin of R4. Otherwise, as in previous species. Abdomen. Dark brown, basal and distal margins of segments 2–6 lighter. Segments cylindrical, covered with dark fine setae, segments 1 to 6 sclerotized, sternite 7 entirely membranous; sternite 8 trapezoid, tergite 8 very short. Male terminalia (Figs. 14, 17, 19). Dark brown, bent downward. Gonocoxites elongate, more or less cylindrical, setose, fused basally to form a syngonocoxite ring around the aedeagus. Gonostylus robust, densely setose, slight shorter than gonocoxite, longer than width at apex, ornamented with two well separated apical rows of spines, ventral row with 16–18 blunt, strongly sclerotized spines, the dorsal row with 3–4 spines. Aedeagal complex consisting of a simple, straight process with a pair of short lateral branches on anterior end and an opening at apex, ventrally covered by the fused parameres. Tergite 9 short, with setation along posterior margin. Cerci lobate, entirely covered with setae.

FIGURES 13–15. Ventral view of male terminalia of Calliceratomyia. 13. C. papaveroi sp. n., holotype. 14. C. pantelhomyia sp. n., holotype. 15. C. zayolli sp. n., male holotype. Scale bar, 0.1 mm.



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FIGURES 16–19. Ventral view of male terminalia of Calliceratomyia. 16. C. papaveroi sp. n., holotype. 17. C. pantelhomyia sp. n., holotype. 18. C. zayolli sp. n., male holotype. 19. C. pantelhomyia sp. n., holotype, inner view of distal end of gonostylus. Scale bar, 16–18: 0.064 mm.

Female. Unknown. Bionomics. Four specimens were collected with Malaise trap in a rainforest in the mountainous area of Chiapas. Distribution. Mexico (Chiapas). Etymology. The specific epithet of this species refers to the name of the type-locality, Pantelho, and the Greek name myia, standing for fly.

Calliceratomyia zayolli, sp. n. (Figs. 4, 7, 12, 15, 18) Diagnosis. Flagellomeres 1–12 with a short ventral extension, last flagellomere slightly shorter than penultimate flagellomere; flagellomeres 1–3 with one longer ventral seta, flagellomere 14 with two longer setae (Figs. 4, 7). M forking considerably basal to origin of R4 (Fig. 12). Extension of male gonocoxites distal to syngonocoxite enlarged midway to apex, an obvious fold along the inner basal margin of gonocoxites. Gonostylus length almost width at apex. Two separate groups of spines apically on inner face of male gonostylus (Figs. 15, 18).


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Type-material. Holotype male, labeled: HOLOTYPE Calliceratomyia zayolli sp. n. Mexico, Hidalgo, Localidad Tlanchinol, km 3.5 carretera Tlanchinol-Apantlazol, GPS: 20’59’16”N, 98’39’36”W, elevation, 1,540 m, in cloud forest (CAIM). Description. Body length (including the antenna), 5.0 mm. Head. Head wider than long. Eyes dichoptic all facets of similar width, length of interommatidial setae close to facet width. Three ocelli present, aligned, of similar size, distance between ocelli slightly smaller than their diameter. Vertex slightly elevated, with setae, transverse suture present, frons bare, face with 14 setae, lateral pair of setae as long as first palpomere. Clypeus more or less triangular. Mouthparts considerably reduced, labellum small; premental apodeme Y-shaped. Maxillary palpus longer than labellum, with three brown, clearly articulated palpomeres. First palpomere humpback, distinctively bulky distally, with 8 socketed setae, entirely covered by sensillae; second palpomere shorter than first, with some sensilla at basal half, projected beyond base of third palpomere, apically with 4 diminute spines and one socketed seta; third palpomere more or less cylindrical, with some sensilla on basal third and 6 minute apical spines. Antenna (Figs. 4, 7) with 15 flagellomeres, 2.2 mm length. Scape and pedicel cylindrical, about 1.2 times as long as wide. Flagellomere 1 more or less cylindrical, flagellomeres 2–8 strongly compressed laterally, with a short but evident ventral projection, flagellomeres 9–13 less modified, flagellomere 14 cylindrical, elongate; apical flagellomere small, cylindrical, 0.7 times length of previous flagellomere (Fig. 7); flagellomeres covered with short fine sensilla-like trichia, one dorsal and one ventral longer seta on flagellomeres 1–3; flagellomeres 13–14 with two setae; all flagellomeres with longer dorsal setae. Thorax. Scutum uniformly brown, evenly setose, without transverse suture; notopleural, supra-alar areas and postalar callus with long bristles; scutellum brown, with a pair of subapical setae. Mediotergite high, bare, dark brown; pleural sclerites brown, anepisternum, katepisternum, mesepimeron, and laterotergite bare. Postpronotum with a pair of long bristles, proepisternum with three long bristles and three small setae. Haltere brown, slightly longer than the first abdominal tergite, pedicel and capitulum covered with short setae. Legs. Similar to previous species. Wing. (Fig. 12) Membrane infuscate brown. Length, 4.7 mm; width, 1.8 mm. M1+2 basally fused to Rs, medial fork more basal than base of R4. Otherwise similar to previous species. Abdomen. Similar to previous species. Male terminalia (Figs. 15, 18). Dark brown, bent downward. Gonocoxites robust, elongate, enlarged midway to apex, setose, fused basally forming a syngonocoxite ring around aedeagus, anterior margin wide, slightly concave. Inner basal margin of gonocoxites with an obvious fold. Gonostylus clearly longer than width at apex, densely setose, slight shorter than gonocoxite, ornamented with two well separated rows of spines apically, ventral row with 12 blunt, strongly sclerotized spines, dorsal row with 7–8 spines. Aedeagal complex consisting of a simple, straight process, with a pair of short lateral branches on anterior end, an opening at apex, ventrally covered by fused parameres. Tergite 9 short, with setation along posterior margin. Cercus lobate, entirely covered with setae. Female. Unknown. Bionomics. The only known specimen was collected with a Malaise trap in a cloud forest in the north part of the state of Hidalgo, Mexico. Distribution. Mexico (Hidalgo). Etymology. The specific epithet refers to zayolli-, which means “fly” in Nahuatl language. The term is used in apposition.

Discussion Vockeroth (2009) was the first to mention this group of ditomyiids for Central America. His chapter has beautiful illustrations of the wing and the antenna of an undescribed species from Costa Rica, but the genus was unfortunately misidentified as Rhipidita (see Vockeroth, 2009: p. 254, figs. 11.2–3). The species is certainly a Calliceratomyia, with strongly pectinate flagellomeres 2–13. That species is still undescribed, as the shape of the final flagellomere indicates, and is obviously closely related to C. pectinata, the type-species of the genus. There is an additional undescribed pectinate species of Calliceratomyia from Peru belonging to The Canadian National Collection of Insects, Arachnids and Nematodes (CNC), Ottawa, Canada. The three species described here, despite not having ventrally elongated flagellomeres, fit in the genus Calliceratomyia. This is indicated by the long gonocoxites basally fused together projecting beyond the level of the aedeagus. As well, the maxillary palpus is 3-segmented. The flagellomeres of the three species from Mexico



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described here have ventral extensions, even though much shorter than in C. pectinata and in the undescribed species illustrated by Vockeroth (2009). Both known species of Rhipidita, on the other hand, have flagellomeres with short pectination, palpomeres 2 and 3 are fused together, and the shape of the gonocoxites and the gonostyli differs from C. pectinata and the species described here. There is no question about these new species belonging to Calliceratomyia Lane. The new described species are considerably similar to each other, but they have consistent differences in the antennae, wings, and male terminalia to allow their delimitation as separate species. They differ from C. pectinata by the much shorter ventral pectination of the flagellomeres and by the presence of separate groups of spines apically on the gonostylus. C. papaveroi, sp. n. has a very short last antennal flagellomere, the ventral extension of flagellomeres 2–8 is wider distally, and all flagellomeres have a distinctive ventral seta. C. pantelhomyia, sp. n. and C. zayolli, sp. n. have the last antennal flagellomere almost as long as the penultimate, the ventral extension of flagellomeres 2–10 is slender (giving a rather triangular shape to the flagellomeres), and a ventral distinctive seta is present only on flagellomeres 1–4. C. pantelhomyia, sp. n. has the medial fork just opposite to the origin of R4, which is much more basal in C. zayolli, sp. n. In C. pantelhomyia, sp. n. the ventral mesal margin is almost straight, while the gonostylus in C. zayolli have 7–8 spines at the dorsal row and the anterior margin of the syngonocoxite has a medial furrow.

Acknowledgments The first author (HH) expresses his sincere thanks to Beatriz Salceda and Rodriguez A. A. for collecting of the type material. The second author has a CNPq Research Fellowship (Proc. 309.240/2014). The authors are very thankful for the kind and useful reviews from Neal Evenhuis and anonymous colleague reviewer.

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