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Research Article
Neotropical Nilothauma Kieffer, 1921 (Diptera, Chironomidae): Key, eleven new species, re-descriptions, new combination and new records
expand article infoLuiz Carlos Pinho, Trond Andersen§
‡ Federal University of Santa Catarina, Florianópolis, Brazil
§ University of Bergen, Bergen, Norway
Open Access

Abstract

Nine new species of Nilothauma Kieffer, N. hamadae sp. nov., N. jupau sp. nov., N. karitiana sp. nov., N. leccii sp. nov., N. marianoi sp. nov., N. mateusi sp. nov., N. txukuyana sp. nov., N. werekena sp. nov. and N. yekwana sp. nov. are described and figured, based on adult males collected in Brazil and N. maya sp. nov. on an adult male from Mexico; N. terena sp. nov. is described as male, pupa and larva based on a reared specimen from Brazil. Nilothauma aleta Roback, 1960 and N. duena Roback, 1960 are re-described and recorded from Brazil. Nilothauma longissimum Mendes & Andersen, 2009 is transferred to Beardius Reiss & Sublette, 1985 and the diagnosis of Nilothauma is emended. New records of thirteen Neotropical Nilothauma species are given and a key to the males of all known species of Nilothauma is provided.

Keywords

Brazil, key, Mexico, Neotropical Region, new combination, new records, new species, taxonomy

Introduction

The genus Nilothauma was erected by Kieffer (1921), based on the Afrotropical N. pictipenne Kieffer, 1921. The Neotropical genera Neelamia Soponis, 1987 and Paranilothauma Soponis, 1987 were both placed as synonyms of Nilothauma by Mendes and Andersen (2009). Most males of Nilothauma can be recognised by having an antenna with 13 flagellomeres, low antennal ratio (except in Nilothauma longissimum Mendes & Andersen, 2009), fore-tibia with long spur on conical, apical scale, high venarum ratio and squama bare (Mendes and Andersen 2009). In addition, many males have median or lateral, sometimes strongly setose lobes or projections on tergite IX. The larvae of Nilothauma inhabit littoral and sublittoral soft sediments of lakes, streams and rivers (Epler et al. 2013).

To date, the genus comprises 52 described species that occur in all zoogeographical regions, except Antarctica (Qi et al. 2014, 2016; Niitsuma 2016; Dantas and Hamada 2017). A total of 22 species are known from the Neotropical and four from the Nearctic Regions (Andersen et al. 2016; Dantas and Hamada 2017).

After examining material collected in several localities in the Neotropical Region, mostly from the Brazilian Amazon, 24 species of Nilothauma were identified. Eleven of them proved to be new to science and are described below as adult males and for N. terena sp. nov. also as larva and pupa. The other thirteen species have their distribution range extended in the Neotropics. Nilothauma aleta Roback, 1960 and N. duena Roback, 1960, originally described from the Peruvian Amazon are re-described, based on material from Brazil and N. longissimum Mendes & Andersen, 2009 is transferred to Beardius Reiss & Sublette, 1985. A key to the males of all known species of Nilothauma is provided.

Material and methods

Alcohol-preserved specimens were dissected and slide-mounted in Euparal. Morphological terminology and abbreviations follow Sæther (1980). Measurements are taken according to Epler (1988) and given as ranges, followed by the mean when more than three specimens were measured, followed by the number of specimens measured in parenthesis.

Abbreviations used in the text as follows:

CEPA Centro de Estudos e Pesquisas Ambientais (Centre for Environmental Studies and Research);

EB Estação Biológica (Biological Station);

INPA Instituto Nacional de Pesquisas da Amazônia (Manaus, Brazil);

MZSP Museu de Zoologia da Universidade de São Paulo (São Paulo, Brazil);

PE Parque Estadual (State Park);

RPPN Reserva Particular do Patrimônio Natural (Private Natural Heritage Reserve);

UFSC Entomological Collection of the Federal University of Santa Catarina (Florianópolis, Brazil);

ZMBN University Museum of Bergen (Bergen, Norway);

ZSM Zoologische Staatsammlung München (Munich, Germany).

Type material is deposited at INPA, MZSP, UFSC, ZMBN and ZSM, as stated in each description. Vouchers are deposited at UFSC, ZMBN and ZSM.

Taxonomy

Beardius Reiss & Sublette, 1985

Beardius longissimus (Mendes & Andersen, 2009), comb. nov.

Nilothauma longissimum Mendes & Andersen, 2009: 26

Material examined

Type material, as in Mendes and Andersen (2009).

Additional material

8 males, slide-mounted: Brazil, Bahia, Camacan, RPPN Serra Bonita, Trilha Bapeba,15°20'35"S, 39°33'34"W, 4.xi.2009, light trap, A.R. Calor et al. leg. 3 males, slide-mounted, as previous, except: 15°23'32"S, 39°33'53"W, 2.xi.2009. 1 male, slide-mounted, as previous, except: 03.ii.2009. 1 male, slide-mounted, as previous, except: córrego 2, 15°23'10"S, 39°34'03"W, 819 m a.s.l., 01.viii.2008, light trap, A.R. Calor, L.S. Lecci, L.C. Pinho & R.A. Moretto leg. 1 male, slide-mounted: Brazil, São Paulo, PE Serra do Mar, Ubatuba, Picinguaba, Camburi stream, 09.ix.2006, light trap, M.R. Spies & A.E. Siegloch leg.

Remarks

Pinho et al. (2013) found that the “presence of apical thin setae on inferior volsella” (character 74, state 1), i.e. a group of two, rarely three, slender simple setae at the very tip of the inferior volsella, in addition to subapical, stouter setae, is the only synapomorphy of Beardius Reiss & Sublette, 1985 in the adult stage. In fact, the character is shared by all species of Beardius and is not found elsewhere, except in Nilothauma longissimum Mendes & Andersen, 2009, a species that was considered to be sister to all remaining Nilothauma by Mendes and Andersen (2009) in a phylogenetic analysis with Paratendipes Kieffer and Pseudochironomus Malloch as outgroups. The comparatively-high antennal ratio in N. longissimum (AR > 1.00) is the only exception in Nilothauma, which generally have very low antennal ratios (AR < 0.40). Further, the venarum ratio (VR) seems to be low in N. longissimum compared to other Nilothauma species; other characters in N. longissimum are consistent with the current diagnosis of Beardius. We therefore propose the new combination and emend the diagnosis of Nilothauma accordingly.

Distribution

The species was originally described from São Paulo State, south-eastern Brazil by Mendes and Andersen (2009); the range is now extended to Bahia State in north-eastern Brazil.

Nilothauma Kieffer, 1921

Emended diagnosis

After transferring Nilothauma longissimum Mendes & Andersen, 2009 to Beardius Reiss & Sublette, 1985 (see above), adult males of Nilothauma become more easily separated from other genera. The diagnosis given by Mendes and Andersen (2009) has to be emended as follows: “antennal ratio generally low (AR < 0.40), one species (N. longissimum sp. nov.) with AR > 1.00.” should read: “antennal ratio generally low (AR < 0.40), occasionally as high as 0.82 (N. soka Andersen, Bello González & Hagenlund, 2016).

The discovery of the pupae of N. terena sp. nov. leads to the diagnosis of the pupa given by Mendes and Andersen (2009) having to be emended as follows: “Frontal setae short, not on tubercles.” should read “Frontal setae short, occasionally long and taeniate, not on tubercles.” Further, “Sternites I–VII bare; sternite VIII with central, longitudinal field of shagreen.” should read “Sternites I–VII usually bare, sternite I occasionally with extensive shagreen; sternite VIII with central, longitudinal field of shagreen.”

Based on the larva of N. terena sp. nov., the diagnosis of the Nilothauma larvae in Epler et al. (2013) should be emended as follows: “Mandible. All teeth pale;” should read: “Mandible. All teeth pale, occasionally inner teeth with somewhat darker pigmentation;” and “Mentum. Pale;” should read: “Mentum. Pale, occasionally with somewhat darker pigmentation”.

Nilothauma aleta Roback, 1960

Figures 1A, B, 17B

Additional material

1 male, slide-mounted: Brazil, São Paulo, São Luís do Paraitinga, PE Serra do Mar, Núcleo Santa Virgínia, trilha Poço do Pito, afluente Paraibuna, 23°20'09"S, 45°08'46"W, 15.ix.2006, light trap, M.R. Spies & A.E. Siegloch leg.

Diagnostic characters

The male can be distinguished from its congeners by the combination of: tergite IX without setose dorsal lobe(s); gonostylus stout; acrostichals absent; anal point wide, covering most setae along posterior margin of tergite IX; inferior volsella slender.

Re-description

Male imago (n = 1). Total length 3.58 mm. Wing length 2.00 mm. Total length/wing length 1.79. Wing length/length of profemur 2.25.

Colouration. Head, thorax and abdomen brown; legs pale, except for ring of brown pigmentation in distal 1/2 to 2/3 of fore- and hind femora, in distal 1/3 of foretibia, in basal 1/8 of mid- and hind tibiae and in distal 1/3 of each tarsomere. Wing membrane apparently hyaline, but faint brown markings are visible when dark-field filter is applied.

Antenna. AR = 0.27. Thirteenth flagellomere 197 µm long.

Head. Temporal setae 7 in single row. Clypeus with 25 setae. Tentorium 123 µm long, maximum width 25 µm. Stipes not measurable. Palp segment lengths (in µm): 39, 34, 123, 147, 191. Third palpomere with 2 sensilla clavata subapically, longest 20 µm long. Fifth palpomere/third palpomere 1.55.

Thorax. Dorsocentrals 12 in single row, acrostichals absent, prealars 3. Scutellum with 6 setae.

Wing. VR = 1.50. Brachiolum with 1 seta, R with 13 setae, R1 with 18 setae, R4+5 with 21 setae, remaining veins bare.

Legs. Spur of fore tibia 44 µm long including 15 µm long scale. Mid-tibia with 1 spur, 15 µm long; hind tibia with 2 spurs, 25 and 29 µm long. Combs of both mid- and hind tibia 20 µm long. Width at apex of fore-tibia 39 µm, of mid-tibia 34 µm, of hind tibia 44 µm. Lengths and proportions of legs as in Table 1.

Table 1.

Lengths (in μm) and proportions of leg segments in Nilothauma aleta Roback, 1960, adult male (n = 1).

Fe Ti ta1 ta2 ta3 ta4
p1 887 601 837 522 404 256
p2 906 690 414 227 177 118
p3 1034 985 699 355 305 197
ta5 LR BV SV BR
p1 148 1.39 1.74 1.77 2.3
p2 89 0.60 3.29 2.30 2.3
p3 108 0.71 2.56 2.89 5.0

Hypopygium (Fig. 1A, B). Tergite IX without lobes, tapering to apex, with 22 short setae underneath anal point. Anal point lanceolate, 50 µm long, maximum width 37 µm. Tergite bands well developed. Laterosternite IX without setae. Phallapodeme 70 µm long; transverse sternapodeme 55 µm long. Gonocoxite 134 µm long. Inferior volsella straight, 52 µm long, 7 µm wide medially, with microtrichia and 8 simple setae apically. Superior volsella pediform, 17 µm long, 7 µm wide at base, covered with microtrichia and with 2 setae apically. Median volsella 7 µm long, with 2 simple setae, longest 12 µm. Gonostylus 95 µm long, straight. HR = 1.42. HV = 3.77.

Figure 1. 

Nilothauma aleta Roback, 1960, adult male A hypopygium, dorsal view B hypopygium with anal point and tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Female adult and immatures

Unknown.

Remarks

Roback (1960) described Nilothauma aleta Roback, 1960 and N. duena Roback, 1960 from the Peruvian Amazon. In their revision of Nilothauma, Adam & Sæther, (1999) regarded the two species as not belonging to Nilothauma since they lack any projections on tergite IX and stated that they appear to belong in Paratendipes Kieffer. Later, Mendes and Andersen (2009) placed Neelamia Soponis and Paranilothauma Soponis as synonyms of Nilothauma and several new Neotropical species have been described demonstrating the large morphological variation in the genus. Mendes and Andersen (2009) emended the diagnosis of Nilothauma and both N. aleta and N. duena fit well into this diagnosis.

Distribution

(Fig. 17B). The species was originally described from the Department of Huánuco, in the Peruvian Amazon by Roback (1960); the range is now extended to Serra do Mar (São Paulo State), in the Brazilian Atlantic Forest.

Nilothauma amazonense Mendes & Andersen, 2009

Figure 17A

Additional material

1 male, slide-mounted: Brazil, Santa Catarina, Grão Pará, Cachoeira do Amado, #27, 28°08'57"S, 49°21'17"W, 16.xi.2012–08.i.2013, Malaise trap, L.C. Pinho, M.C. Novaes & M.F. Haddad leg. 1 male, slide-mounted: Brazil, Santa Catarina, São Francisco do Sul, Distrito do Saí, 26°11'42"S, 48°43'53"W, 18.i–18.iii.2020, Malaise trap #150, small stream, L.C. Pinho et al. leg.

Remarks

Nilothauma amazonense Mendes & Andersen, 2009 was described, based on a single male from the Amazon. The specimens of N. amazonense from southern Brazil, however, differ slightly from the holotype. Mendes and Andersen (2009) stated that hind ta2 being shorter than ta3 (ratio of ta2/ta3 length = 0.73) is one of the diagnostic characters of the species. However, in the specimens from southern Brazil, hind ta2 and ta3 are subequal in length (ratio of ta2/ta3 length = 0.94–0.97). Body size, measured as Total Length (TL) is also larger (TL of holotype = 1.53 mm; TL of southern populations = 2.00–2.05 mm). Similar differences in body size between Amazonian and southern Atlantic Forest populations were also found in Beardius urupeatan Pinho, Mendes & Andersen, 2009 [TL Amazon = 2.32–2.51, 2.38 (6); TL southern Atlantic Forest = 2.68–3.00, 2.96 (8)]. This intraspecific variation might be due to the higher temperature in the Amazon Region when compared to the localities in the southern parts of the Atlantic Forest. Populations of chironomid species inhabiting different habitats may show variation in voltinism and more rapid growth can result in smaller body size (Tokeshi 1995; Pinho et al. 2009).

Distribution

(Fig. 17A). The species was originally described from the Amazonian Region by Mendes and Andersen (2009); the range is now extended to Santa Catarina State in southern Brazil.

Nilothauma anamariae Dantas & Hamada, 2017

Figure 17D

Additional material

1 male, slide-mounted: Brazil, Rondônia, Candeias do Jamari, Rio Preto, Ponte de Madeira, #01, 08°52'40"S, 63°38'02"W, 19–20.vii.2012, light trap, R. Boldrini & A.S. Fernandes leg. 1 male, slide mounted: Brazil, Mato Grosso, Ribeirão Cascalheira, Fazenda Campina Grande, Rio Suiá Miçu, 28–30.xi.2006, light trap, A.R. Calor, F.R. Silva & S. Mateus leg. 2 males, slide-mounted: Brazil, Mato Grosso, Ribeirão Cascalheira, Fazenda Campina Verde, Rio Suiá Miçu, 12°48.591'S, 52°06.925'W, 10.x.2007, light trap, L.C. Pinho, S. Mateus, L. Torati & F.R. Silva leg. 1 male, slide-mounted: Brazil, Pará State, Rurópolis, Rio Tambor, 29.x.2007, light trap, N. Hamada et al. leg.

Remarks

The inferior volsella can have up to 3–4 simple, curved setae apically.

Distribution

(Fig. 17D). The species was originally described from the Rio Grande do Sul State in southern Brazil by Dantas and Hamada (2017); the range is now extended to Mato Grosso, Rondônia and Pará States in central and northern Brazil.

Nilothauma aripuanense Mendes & Andersen, 2009

Figure 17B

Additional material

2 males, slide-mounted: Brazil, Rondônia, Candeias do Jamari, Rio Preto, Ponte de Madeira, #01, 08°52'40"S, 63°38'02"W, 19–20.vii.2012, light trap, R. Boldrini & A.S. Fernandes leg. 2 males, slide-mounted: Brazil, Rondônia, Teixeirópolis, Balneário com Cachoeira, 10°55'20"S, 62°22'34"W, 03.ix.2012, light #13, N. Hamada, R. Boldrini, A.S. Fernandes & J.M. Cavalcante leg. 1 male, slide-mounted: Brazil, Roraima, Boa Vista, Rio Cauamé, 02°52'06"N, 60°44'24"W, 9.iii.2009, light trap, L.M. Fusari leg. 1 male, slide-mounted: Brazil, Roraima, Boa Vista, BR-174, Igarapé Água Boa, 02°43'32"N, 60°48'43"W, 2014, N. Hamada leg. 1 male, slide-mounted: Brazil, Amazonas, Presidente Figueiredo, AM-240 Km 20, Balneário Sossego da Pantera, Igarapé da Onça, 02°02'31"S, 59°51'05"W, 02.vii.2008, light trap, C. Azevedo leg. 1 male, slide-mounted: Brazil, Amazonas, upper Rio Marauiá, downstream of Cachoeira Santo Antônio, surface float skimmed, 22.i.1963, E.J. Fittkau leg. (A485, ZSM).

Distribution

(Fig. 17B). The species was originally described from the Amazonas and Mato Grosso States by Mendes and Andersen (2009); the range is now extended to Roraima and Rondônia States in the Brazilian Amazon.

Nilothauma calori Mendes & Andersen, 2009

Figures 2A, 16C

Additional material

1 male, slide-mounted: Brazil, Amazonas, Manaus, Reserva Florestal Adolfo Ducke, Igarapé Bolívia, 02°49'15"S, 59°56'31"W, 9–12.xi.2008, Malaise trap suspensa 2, N. Hamada et al. leg. 2 males, slide-mounted: Brazil, Mato Grosso, Cuiabá, 10–11.x.1965, Brundin net, E.J. Fittkau leg. (A 580, ZSM).

Remarks

Mendes and Andersen (2009) stated that the superior volsella has a “lateral strongly sclerotized, spine-like projection”. In dorsolateral view (Fig. 2A), it can be seen that this spine-like projection originates from the base of the volsella and is equally long as the volsella proper.

Figure 2. 

Variation in superior volsella A dorsolateral view of superior volsella of Nilothauma calori Mendes & Andersen, 2009 B lateral view of superior volsella of Nilothauma complicatum Mendes & Andersen, 2009.

Distribution

(Fig. 16C). The species was originally described from Acre State by Mendes and Andersen (2009); the range is now extended to the Mato Grosso and Amazonas States.

Nilothauma complicatum Mendes & Andersen, 2009

Figures 2B, 16A

Additional material

1 male, slide-mounted: Brazil, Rondônia, Candeias do Jamari, Rio Preto, Ponte de Madeira, #01, 08°52'40"S, 63°38'02"W, 19–20.vii.2012, light trap, R. Boldrini & A.S. Fernandes leg. 3 males, slide-mounted: Brazil, Mato Grosso, Nova Xavantina, Fazenda Sr. Queté, Córrego Voadeira, 14°32.187'S, 52°30.902'W, 16.x.2007, light trap, L.C. Pinho, S. Mateus, L. Torati & F.R. Silva leg. 2 males, slide-mounted, as previous, except: Córrego Cachoeira, 14°32.817'S, 52°31.395'W. 1 male, slide-mounted, as previous, except: 14°41.577'S, 52°27.203'W, 13.x.2007. 2 males, slide-mounted, as previous, except: Estrada p/ Rancho Helena, Córrego Ponte de Pedra, 14°47.908'S, 52°37.226'W, 15.x.2007. 1 male, slide-mounted, as previous, except: Córrego Voadeira, 14°41.577'S, 52°27.203'W, 13.x.2007. 1 male, slide-mounted: Brazil, Mato Grosso, Ribeirão Cascalheira, Estrada Fazenda Manaus, 1° afluente Rio Bonito, 12°57.088'S, 51°52.480'W, 08.x.2007, light trap, L.C. Pinho, S. Mateus, L. Torati & F.R. Silva leg.

Remarks

Mendes and Andersen (2009) stated that the superior volsella has a “marginal row of flattened setae”. In lateral view (Fig. 2B), it can be seen that the volsella is quite wide medially with a row of lamellae apparently covered with weak microtrichia.

Distribution

(Fig. 16A). The species was originally described by Mendes and Andersen (2009), based on a single male from the Espírito Santo State; the range is now extended to the Mato Grosso and Rondônia States in central and northern Brazil.

Nilothauma duena Roback, 1960

Figures 3A, B, 16C

Additional material

1 male, slide-mounted: Brazil, Bahia, Camacan, Fazenda do Waldemar da farmácia, Córrego abaixo da represa de abastecimento, 15°25'16"S, 39°33'57"W, 300 m a.s.l., 05.viii.2008, light trap, A.R. Calor, L.S. Lecci, L.C. Pinho & R.A. Moretto leg. 1 male, slide-mounted: Brazil, São Paulo, Pindamonhangaba, Fazenda São Sebastião, Afluente Cedro 3, 22°50'16"S, 45°28'27"W, 18.ix.2006, light trap, M.R. Spies & A.E. Siegloch leg.

Diagnostic characters

The male can be distinguished from its congeners by the combination of: tergite IX without setose dorsal lobe(s); anal point slightly spatulate; wing unmarked; abdominal tergites I–VIII with basal half light brown, distal half pale; gonostylus and inferior volsella stout.

Figure 3. 

Nilothauma duena Roback, 1960, adult male A hypopygium, dorsal view B hypopygium with anal point and tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Re-description

Male imago (n = 2, unless otherwise stated). Total length 2.37–2.76 mm. Wing length 1.56–1.68 mm. Total length/wing length 1.58–1.66. Wing length/length of profemur 2.60–2.62.

Colouration. Head and thorax light brown; legs pale except for brown pigmentation in basal 1/3 and distal 1/5 of fore femur, basal 1/5 and distal 1/3 of fore tibia, distal 1/8 of mid- and hind femora, basal 1/8 and distal 1/8 of mid- and hind tibiae and entire tarsi. Abdominal tergites I–VIII with basal half light brown, distal half pale; hypopygium light brown. Wing membrane hyaline.

Antenna. AR = 0.17–0.18. Thirteenth flagellomere 108–118 µm long.

Head. Temporal setae 9–10 in single row. Clypeus with 13–15 setae. Tentorium 98 (1) µm long, maximum width 20 (1) µm. Stipes not measurable. Palp segment lengths (in µm): 17–25, 17–25, 32 (1), 35 (1), 62 (1). Third palpomere with 2 sensilla clavata subapically, longest 15 µm long. Fifth palpomere/third palpomere 1.92 (1).

Thorax. Dorsocentrals 16–17 in single row, acrostichals 14–16, prealars 3. Scutellum with 2–3 setae.

Wing. VR = 1.55–1.56. Brachiolum with 1 seta, R with 12–13 setae, R1 with 16–17, R4+5 with 5–22 setae, remaining veins bare.

Legs. Spur of fore tibia 54–59 µm long including 15–20 µm long scale. Mid-tibia with 1 spur, 25–29 µm long; hind tibia with 2 spurs, 49–51 and 28–31 µm long. Combs of mid-tibia 15–20 µm long, of hind tibia 18–25 µm long. Width at apex of fore tibia 48–50 µm, of mid-tibia 48–50 µm, of hind tibia 49–59 µm. Lengths and proportions of legs as in Table 2.

Table 2.

Lengths (in μm) and proportions of leg segments in Nilothauma duena Roback, 1960, adult males (n = 2).

Fe Ti ta1 ta2 ta3 ta4
p1 601–640 453–502 660–690 374–376 276–278 207–217
p2 621–670 453–473 296–305 148–158 108–110 69–79
p3 739–778 670–699 404–443 217–236 207–210 148–150
ta5 LR BV SV BR
p1 105–108 1.37–1.46 1.72–1.74 1.60–1.66 1.8–2.7
p2 49–59 0.65–0.66 3.48–3.49 3.56–3.74 1.8–2.7
p3 79–99 0.60–0.63 2.78–2.79 3.33–3.49 3.6–4.7

Hypopygium (Fig. 3A, B). Tergite IX without dorsal lobes, with triangular posterior margin with 13–15 weak setae along posterior margin to each side of base of anal point. Anal point spatulate, 22–30 µm long, maximum width 10–12 µm. Tergite bands well developed. Laterosternite IX with 1–2 setae. Phallapodeme 47–60 µm long; transverse sternapodeme 55–62 µm long. Gonocoxite 112–125 µm long. Inferior volsella straight, 40–45 µm long, 15–20 µm wide medially, with microtrichia and 10–11 strong, simple setae apically. Superior volsella tapering to apex, 20–37 µm long, 12–14 µm wide at base, covered with microtrichia and apparently bare at tip. Median volsella 7–10 µm long, with 3–4 setae, longest 20–22 µm long. Gonostylus 87–90 µm long, straight. HR = 1.29–1.43. HV = 2.72–3.17.

Female adult and immatures

Unknown.

Remarks

See remarks for N. aleta Roback, 1960.

Distribution

(Fig. 16C). The species was originally described by Roback (1960) from the Department of Huánuco, in the Peruvian Amazon; the range is now extended to Serra Bonita (Bahia State) and Serra do Mar (São Paulo State), in the Brazilian Atlantic Forest.

Nilothauma fittkaui (Soponis, 1987)

Figure 17A

Additional material

2 males, slide-mounted: Brazil, São Paulo, Campos do Jordão, PE Campos do Jordão, Córrego Canhambora, 1538 m a.s.l., 22°41'44"S, 45°29'30"W, 13.i.2006, light trap, M.R. Spies leg. 1 male, slide-mounted, Costa Rica, La Selva, 03.iv.1993, Malaise trap, O.A. Sæther leg.

Distribution

(Fig. 17A). The species was described by Soponis (1987) from Amazonas and later recorded from Acre, Espírito Santo and Para States in Brazil and from Ecuador by Mendes and Andersen (2009). The range is now extended south to São Paulo State in Brazil and north to Costa Rica in Central America.

Nilothauma hamadae sp. nov.

Figures 4A, B, 16A

Type material

Holotype male, slide-mounted: Brazil, Amazonas, Barcelos, Rio Aracá, Foz do Igarapé Cuieiras, 00°19'15"N, 63°16'15"W, 35 m a.s.l., 30.vii–01.viii.2009, light trap #11, N. Hamada et al. leg. (UFSC).

Etymology

The specific epithet is a noun in the genitive case which honours Neusa Hamada for her great contribution to the knowledge of Amazonian Chironomidae.

Figure 4. 

Nilothauma hamadae sp. nov. adult male A hypopygium, dorsal view B hypopygium with anal point and tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Diagnostic characters

The male can be distinguished from its congeners by the combination of: tergite IX with single, median setose dorsal lobe, consisting of a low, but wide protuberance with about 25 strong setae; anal point very broad (about half the width of tergite IX) and lanceolate; median volsella separated from superior volsella.

Description

Male imago (n = 1). Total length 1.94 mm. Wing length 1.13 mm. Total length/wing length 1.71. Wing length/length of profemur 2.30.

Colouration. Head, thorax and abdomen medium brown. Legs light brown. Wing membrane hyaline.

Antenna. AR = 0.40. Thirteenth flagellomere 217 µm long.

Head. Temporal setae 7 in single row. Clypeus with 15 setae. Tentorium 75 µm long, maximum width 12 µm. Stipes 92 µm long. Palp segment lengths (in µm): 25, 25, 75, 127, 144. Third palpomere with 2 sensilla clavata subapically, longest 20 µm long. Fifth palpomere/third palpomere 1.93.

Thorax. Dorsocentrals 7 in single row, acrostichals 10, prealars 2. Scutellum with 4 setae.

Wing. VR = 1.37. Brachiolum with 1 seta, R with 13 setae, R1 with 7 setae, R4+5 with 14 setae, remaining veins bare.

Legs. Spur of fore tibia 34 µm long including 12 µm long scale. Mid-tibia with 1 spur, 15 µm long; hind tibia with 2 spurs, 20 and 25 µm long. Combs of mid-tibia 12 µm long, of hind tibia 15 µm long. Width at apex of fore tibia 34 µm, of mid-tibia 34 µm, of hind tibia 39 µm. Lengths and proportions of legs as in Table 3.

Table 3.

Lengths (in μm) and proportions of leg segments in Nilothauma hamadae sp. nov., adult male (n = 1).

Fe Ti ta1 ta2 ta3 ta4
p1 493 364
p2 473 364 217 108 79 49
p3 532 542 315 167 158 99
ta5 LR BV SV BR
p1
p2 49 0.59 3.62 3.86 3.3
p3 69 5.0

Hypopygium (Fig. 4A, B). Tergite IX narrow, tapering to apex, with central rounded lobe bearing 25 simple, strong setae; with 9 simple setae along posterior margin underneath the anal point. Anal point lanceolate, 42 µm long, 27 µm wide. Tergite bands lacking. Laterosternite IX with 1 seta. Phallapodeme 40 µm long; transverse sternapodeme 15 µm long. Gonocoxite 75 µm long. Inferior volsella strongly curved, 37 µm long, 7 µm wide medially, with microtrichia in basal half, with 4 simple setae apically and 1 simple seta subapically. Superior volsella 30 µm long, 4 µm wide at base, covered with microtrichia and fringed at apex. Median volsella 7 µm long, with 2 simple setae, longest 10 µm. Gonostylus 112 µm long, with basal half strongly curved and distal half straight. HR = 0.67. HV = 1.73.

Female adult and immatures

Unknown.

Distribution

(Fig. 16A). Known from Barcelos (Amazonas State), in the Brazilian Amazon.

Nilothauma jaraguaense Mendes & Andersen, 2009

Figure 17D

Additional material

1 male, slide-mounted: Brazil, São Paulo, Salesópolis, EB Boraceia, Rio Claro, Poço Verde, 18.ix.2002, light trap, A.S. Melo, C.G. Froehlich, R. Mariano, A. Prather & R. Blahnik leg. 1 male, slide-mounted: Brazil, São Paulo, Jundiaí, PE Serra do Japi, 23.ix.2008, light trap, R. Mariano & L.S. Lecci leg.

Distribution

(Fig. 17D). The species was described by Mendes and Andersen (2009), based on a single male from Parque Estadual do Jaraguá in São Paulo State, Brazil.

Nilothauma jupau sp. nov.

Figures 5A–C, 16A

Type material

Holotype male, slide-mounted: Brazil, Rondônia, Teixeirópolis, Balneário com Cachoeira, 10°55'20"S, 62°22'34"W, 03.ix.2012, light trap #13, N. Hamada, R. Boldrini, A.S. Fernandes & J.M. Cavalcante leg. (UFSC). Paratype: 1 male, slide-mounted, same data as holotype (INPA).

Etymology

The specific epithet honours the Jupaú, indigenous people from Rondônia State, Brazilian Amazon. The name is to be regarded as a noun in apposition.

Diagnostic characters

The male can be distinguished from its congeners by the combination of: tergite IX with thorn and without setose dorsal lobe(s); anal point spatulate; wing with conspicuous dark markings; abdominal tergites II, III, VI, VII and VIII dark brown.

Figure 5. 

Nilothauma jupau sp. nov. adult male A wing B hypopygium, dorsal view C hypopygium with anal point and tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Description

Male imago (n = 2, unless otherwise stated). Total length 2.04–2.14 mm. Wing length 1.07–1.14 mm. Total length/wing length 1.78–2.01. Wing length/length of profemur 2.03–2.27.

Colouration. Head and thorax brown; legs pale, except for entire fore femur, distal half of mid- and hind femora and distal 1/3 of fore- and hind tibiae with brown pigmentation; abdomen pale, except for brown pigmentation in segments II, III, VI, VII and VIII. Wing membrane with extensive dark markings.

Antenna. AR = 0.28–0.32. Thirteenth flagellomere 115–134 µm long.

Head. Temporal setae 7–8 in single row. Clypeus with 20–24 setae. Tentorium 57–72 µm long, maximum width 15–17 µm. Stipes 95–100 µm long. Palp segment lengths (in µm): 13–14, 30–32, 85–95, 105–117, 115–134. Third palpomere with 2–3 sensilla clavata subapically, longest 14–15 µm. Fifth palpomere/third palpomere 1.21–1.69.

Thorax. Dorsocentrals 6–7 in single row, acrostichals 8–14, prealars 2. Scutellum with 2 setae.

Wing (Fig. 5A). VR = 1.63–1.64. Brachiolum with 1 seta, R with 8–10 setae, R1 with 6 setae, R4+5 with 11–15 setae, remaining veins bare.

Legs. Spur of fore tibia 39–49 µm long including 20–25 µm long scale. Mid-tibia with 1 spur, 20–25 µm long; hind tibia with 2 spurs, 20–25 and 23–28 µm long. Combs of mid-tibia 15–20 µm long, of hind tibia 15–20 µm long. Width at apex of fore tibia 44 µm, of mid-tibia 39–44 µm, of hind tibia 47–51 µm. Lengths and proportions of legs as in Table 4.

Table 4.

Lengths (in μm) and proportions of leg segments in Nilothauma jupau sp. nov., adult males (n = 2).

Fe Ti ta1 ta2 ta3 ta4
p1 502–522 404–424
p2 483–522 335–345 180–187 87–89 57–59 37–39
p3 542–571 512–522 270–276 138–148 138–148 85–89
ta5 LR BV SV BR
p1
p2 28–30 0.54–0.56 3.22–3.31 4.22–4.56 1.8–2.8
p3 59–69 0.53–0.54 2.93–3.23 3.82–3.96 1.5–3.2

Hypopygium (Fig. 5B, C). Tergite IX without dorsal lobes, with rounded posterior margin, with 4–5 clustered setae anteriorly to base of anal point and 8–12 weaker setae to each side of anal point. Anal point spatulate, 40–42 µm long, maximum width 7–10 µm. Tergite bands well developed. Laterosternite IX with 2–3 setae, with thorn. Phallapodeme 42–52 µm long; transverse sternapodeme 20–22 µm long. Gonocoxite 70–75 µm long, with longer microtrichia dorsomedially. Inferior volsella slightly curved, 30–32 µm long, 15–18 µm wide medially, with microtrichia and 5–6 simple setae in apical one third. Superior volsella slender, 12–20 µm long, 4–5 µm wide at base, covered with microtrichia and with 2 setae at apex, longest 8–13 µm. Median volsella consisting of small tubercle situated underneath superior volsella, 6–7 µm long, with 2 setae at apex, longest 6–7 µm long. Gonostylus 70–92 µm long, straight. HR = 0.81–1.00. HV = 2.21–3.05.

Female adult and immatures

Unknown.

Distribution

(Fig. 16A). Known from Rondônia State, Brazilian Amazon.

Nilothauma karitiana sp. nov.

Figures 6A–C, 16C

Type material

Holotype male, slide-mounted: Brazil, Rondônia, Candeias do Jamari, Rio Preto, Ponte de Madeira, 08°52'40"S, 63°38'02"W, 19–20.vii.2012, light trap #01, R. Boldrini & A.S. Fernandes leg. (UFSC). Paratype: 1 male adult, slide-mounted: BRAZIL, Amazonas, Barcelos, Rio Aracá, #9, 69 m a.s.l., 00°24'39"N, 63°23'12"W, 28.vii–06.viii.2009, light trap #3, N. Hamada et al. leg. (INPA).

Etymology

The specific epithet honours the Karitiana, indigenous people from the Rio Jamari Basin in the Rondônia State (Brazil). The name is to be regarded as a noun in apposition.

Figure 6. 

Nilothauma karitiana sp. nov. adult male A hypopygium, dorsal view B superior volsella, dorsal view C hypopygium with anal point and tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Diagnostic characters

The male can be distinguished from its congeners by the combination of: tergite IX without setose dorsal lobe(s); anal point spatulate; wing unmarked; superior volsella fused to median volsella; gonostylus very long, narrow basally and apically, swollen medially.

Description

Male imago (n = 2, unless otherwise stated). Total length 3.32–3.78 mm. Wing length 1.71–1.83 mm. Total length/wing length 1.94–2.07. Wing length/length of profemur 2.02–2.06.

Colouration. Head, thorax, legs and abdomen uniformly light brown. Wing membrane hyaline.

Antenna. AR = 0.22 (1). Thirteenth flagellomere 217 µm long.

Head. Temporal setae 9 (1) in single row. Clypeus with 10–11 setae. Tentorium 105 (1) µm long, maximum width 17 (1) µm. Stipes 122 (1) µm long. Palp segment lengths (in µm): 25–35, 27–37, 60 (1), 80 (1), 154 (1). Third palpomere with 3 (1) sensilla clavata subapically, longest 20 (1) µm long. Fifth palpomere/third palpomere 2.57

Thorax. Dorsocentrals 9–13 in single row, acrostichals 12, prealars 2–3. Scutellum with 2 setae.

Wing. VR = 1.33–1.46. Brachiolum with 1 seta, R with 11–14 setae, R1 with 10–12 setae, R4+5 with 3–4 setae at apex, remaining veins bare.

Legs. Spur of fore tibia 59–64 µm long including 17–20 µm long scale. Mid-tibia with 1 spur, 23–25 µm long; hind tibia with 2 spurs, 23–25 and 29–33 µm long. Combs of mid-tibia 15–20 µm long, of hind tibia 20–23 µm long. Width at apex of fore tibia 59 µm, of mid-tibia 59 µm, of hind tibia 64 µm. Lengths and proportions of legs as in Table 5.

Table 5.

Lengths (in μm) and proportions of leg segments in Nilothauma karitiana sp. nov., adult males (n = 1–2).

Fe Ti ta1 ta2 ta3 ta4
p1 847–887 729–808 1054 542 424 335
p2 739–798 532–561 374 158 108 69
p3 896–965 867–926 493–522 246–256 240–246 144–148
ta5 LR BV SV BR
p1 138 1.30 1.91 1.61 2.2
p2 49 0.67 4.51 3.63 3.2
p3 95–99 0.56–0.57 3.01–3.27 3.58–3.62 4.0–5.0

Hypopygium (Fig. 6A, B). Tergite IX without dorsal lobes, tapering to apex, with 2–3 median and 8–9 setae to each side of anal point. Anal point spatulate, 55–57 µm long, 17–20 µm wide. Tergite bands well developed. Laterosternite IX with 1–2 setae. Phallapodeme 82–90 µm long; transverse sternapodeme 42–55 µm long. Gonocoxite 138–142 µm long. Inferior volsella slightly curved, 97–107 µm long, 17–20 µm wide medially, with microtrichia and 6–7 simple setae subapically. Superior volsella digitiform, 55–65 µm long, 17–20 µm wide at base, covered with microtrichia and with 4 setae apically. Median volsella fused to superior volsella, consisting of 2–4 small tubercles each bearing single, simple seta, longest 22–25 µm. Gonostylus 204–232 µm long, straight, narrow basally and apically, swollen medially. HR = 0.62–0.70. HV = 1.62–1.63.

Female adult and immatures

Unknown.

Distribution

(Fig. 16C). Known from Rondônia and Amazonas States, in the Brazilian Amazon.

Nilothauma leccii sp. nov.

Figures 7A, B, 16B

Type material

Holotype male, slide-mounted: Brazil, São Paulo, São Sebastião, Rio das Pedras, 23°44'27"S, 45°37'12"W, 28.x.2005, light trap, A.R. Calor et al. leg. (UFSC).

Etymology

The specific epithet is a noun in the genitive case honouring Lucas Silveira Lecci, for his friendship and prolific fieldwork.

Diagnostic characters

The male can be separated from its congeners by its large size combined with unmarked wing; spatulate anal point; superior volsella leaf-shapped; inferior volsella with strong, split setae and digitiform and strongly setose gonostylus.

Figure 7. 

Nilothauma leccii sp. nov. adult male A hypopygium, dorsal view B hypopygium with anal point and tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Description

Male imago (n = 1). Total length 4.35 mm. Wing length 1.96 mm. Total length/wing length 2.22. Wing length/length of profemur 2.45.

Colouration. Thorax and legs brown, abdomen light brown. Wing membrane without dark markings.

Antenna. AR = 0.16. Thirteenth flagellomere 124 µm long.

Head. Temporal setae 6 in single row. Clypeus with 15 setae. Tentorium 113 µm long, maximum width 25 µm. Stipes not measurable. Palp segment lengths (in µm): 37, 33, 74, 107, 138. Third palpomere with 2 sensilla clavata subapically, longest about 25 µm long. Fifth palpomere/third palpomere 1.86.

Thorax. Antepronotum with 4 setae. Dorsocentrals 17 partly biserial posterior, acrostichals 14, prealars 6. Scutellum with 13 setae.

Wing. VR = 1.53. Brachiolum with 2 setae, R with 15 setae, R1 with 24 setae, R4+5 with 31 setae, remaining veins bare.

Legs. Spur of fore tibia 65 µm long including 41 µm long scale. Mid-tibia with 1 spur, 47 µm long; hind tibia with 2 spurs, 43 and 65 µm long. Combs of mid-tibia 29 µm long, of hind tibia 47 µm long. Width at apex of fore tibia 69 µm, of mid-tibia 73 µm, of hind tibia 89 µm. Lengths and proportions of legs as in Table 6.

Table 6.

Lengths (in μm) and proportions of leg segments in Nilothauma leccii sp. nov., adult male (n = 1).

Fe Ti ta1 ta2 ta3 ta4
p1 801 670 972 490 384 286
p2 874 605 400 180 139 90
p3 964 989 596 302 261 155
ta5 LR BV SV BR
p1 139 1.45 1.88 1.51 1.7
p2 65 0.66 3.97 3.69 4.3
p3 98 0.60 3.09 3.27 4.8

Hypopygium (Fig. 7A, B). Tergite IX without dorsal lobes, posterior margin subtriangular with 7 setae to each side of the anal point. Anal point spatulate, 59 µm long, 10 µm wide basally, 17 µm wide medially. Tergite bands not continuous. Laterosternite IX with 4 setae. Phallapodeme 61 µm long; transverse sternapodeme 35 µm long. Gonocoxite 171 µm long. Inferior volsella weakly curved, 113 µm long, 15 µm wide subapically, with microtrichia and 7 strong, apically split setae. Superior volsella leaf-shaped, 69 µm long, 10 µm wide at base, 23 µm wide medially, covered with microtrichia and with few weak setae ventrally and along inner margin. Median volsella narrow, 48 µm long, covered with microtrichia and with 4 setae on tubercles at apex, setae about 23 µm long. Gonostylus digitiform, strongly setose, 163 µm long, 37 µm wide medially. HR = 1.05. HV = 2.67.

Female adult and immatures

Unknown.

Distribution

(Fig. 16B). Only known from São Paulo State in Brazil.

Nilothauma marianoi sp. nov.

Figures 8A, B, 16B

Type material

Holotype male, slide-mounted: Brazil, Bahia, Barreiras, Rio de Janeiro, cachoeira Acaba Vidas, 11°53'40"S, 45°36'57"W, 722 m a.s.l., 14.x.2008, light trap, A.R. Calor, R. Mariano & S. Mateus leg. (UFSC).

Etymology

The specific epithet is a noun in the genitive case honouring Rodolfo Mariano, for his friendship and prolific fieldwork.

Diagnostic characters

The male can be distinguished from its congeners by the combination of: wing without dark markings; tergite IX without setose dorsal lobe(s) or thorns, with single, strong median seta, with narrowly triangular posterior margin and small, apical, parallel-sided anal point.

Figure 8. 

Nilothauma marianoi sp. nov. adult male A hypopygium, dorsal view B hypopygium with anal point and tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Description

Male imago (n = 1). Total length 3.51 mm. Wing length 1.45 mm. Total length/wing length 2.43. Wing length/length of profemur 2.27.

Colouration. Thorax and legs brown, abdomen light brown. Wing membrane without dark markings.

Antenna. AR = 0.19. Thirteenth flagellomere 152 µm long.

Head. Temporal setae 6 in partly double row. Clypeus with 9 setae. Tentorium 98 µm long, maximum width 18 µm. Stipes not measurable. Palp segment lengths (in µm): 17, 18, 44, 99, 117. Sensilla clavata on third palpomere not discernable. Fifth palpomere/third palpomere 2.66.

Thorax. Dorsocentrals 6 in single row, acrostichals 4, prealars 2. Scutellum with 2 setae.

Wing. VR = 1.39. Brachiolum with 1 seta, R with 8 setae, R4+5 with 1 apical seta, remaining veins bare.

Legs. Spur of fore tibia 68 µm long including 28 µm long scale. Mid-tibia with 1 spur, 47 µm long; hind tibia with 2 spurs, 43 and 61 µm long. Combs of mid-tibia 21 µm long, of hind tibia 26 µm long. Width at apex of fore tibia 47 µm, of mid-tibia 48 µm, of hind tibia 52 µm. Lengths and proportions of legs as in Table 7.

Table 7.

Lengths (in μm) and proportions of leg segments in Nilothauma marianoi sp. nov., adult male (n = 1).

Fe Ti ta1 ta2 ta3 ta4
p1 637 458 645 401 310 221
p2 621 425 261 131 90 65
p3 694 686 384 180 180 114
ta5 LR BV SV BR
p1 106 1.41 1.68 1.70 2.6
p2 49 0.62 3.90 4.05 5.0
p3 74 0.55 3.22 3.60 7.1

Hypopygium (Fig. 8A, B). Tergite IX without dorsal lobes, with single, median, strong setae, posterior margin narrowly subtriangular with 7 setae to each side. Anal point situated apically, small, parallel-sided with rounded apex, 14 µm long, 4 µm wide basally, 3 µm wide medially. Tergite bands lacking. Laterosternite IX with 2 setae. Phallapodeme 51 µm long; transverse sternapodeme 11 µm long. Gonocoxite 104 µm long. Inferior volsella weakly curved, 103 µm long, 11 µm wide subapically, with microtrichia and 18 setae in apical one-half. Superior volsella subquadrangular, 48 µm long, 17 µm wide medially, covered with microtrichia. Median volsella consisting of 14 µm long tubercle, covered with microtrichia and with 1 strong apical seta, setae about 19 µm long. Gonostylus nearly straight, 104 µm long, 10 µm wide medially, 17 µm wide subapically. HR = 0.72. HV = 2.44.

Female adult and immatures

Unknown.

Distribution

(Fig. 16B). Only known from Bahia State in Brazil.

Nilothauma mateusi sp. nov.

Figures 9A, B; 16B

Type material

Holotype male, slide-mounted: Brazil, Mato Grosso, Nova Xavantina, Fazenda Sr. Queté, Córrego Cachoeira, 14°32.817'S, 52°31.395'W, 16.x.2007, light trap, L.C. Pinho, S. Mateus, L. Torati & F.R. Silva leg. (UFSC).

Etymology

The specific epithet is a noun in the genitive case honouring Sidnei Mateus, for his friendship and prolific fieldwork.

Diagnostic characters

The male can be distinguished from its congeners by the combination of: wing without markings; tergite IX with pair of rounded lobes submedially with about 14 long setae; anal point parallel-sided; superior volsella small, subtriangular, projecting medially, with 2 setae on tubercles apically.

Figure 9. 

Nilothauma mateusi sp. nov. adult male A hypopygium, dorsal view B hypopygium with anal point and tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Description

Male imago (n = 1). Total length 2.09 mm. Wing length 0.87 mm. Total length/wing length 2.40. Wing length/length of profemur 2.41.

Colouration. Thorax and legs brown, abdomen light brown. Wing membrane without dark markings.

Antenna. AR = 0.19. Thirteenth flagellomere 82 µm long.

Head. Temporal setae 7 in single row. Clypeus with 9 setae. Tentorium 55 µm long, maximum width 12 µm. Stipes not measurable. Palp segment I–III lengths (in µm): 21, 19, 55; remaining palp segments lost. Third palpomere with 2 sensilla clavata subapically, longest about 10 µm.

Thorax. Dorsocentrals 8 in single row, acrostichals 6, prealars 2. Scutellum with 2 setae.

Wing. VR = 1.64. Brachiolum with 1 seta, R with 6 setae, R1 with 2 setae, R4+5 with 4 setae, remaining veins bare.

Legs. Spur of fore tibia 37 µm long including 18 µm long scale. Mid-tibia with 1 spur, 23 µm long; hind tibia with 2 spurs, 19 and 28 µm long. Combs of mid-tibia 14 µm long, of hind tibia 17 µm long. Width at apex of fore tibia 29 µm, of mid-tibia 30 µm, of hind tibia 33 µm. Lengths and proportions of legs as in Table 8.

Table 8.

Lengths (in μm) and proportions of leg segments in Nilothauma mateusi sp. nov., adult male (n = 1).

Fe Ti ta1 ta2 ta3 ta4
p1 360 272 340 136 112 68
p2 352 248 140 64 52 32
p3 416 392 208 108 112 72
ta5 LR BV SV BR
p1 40 1.25 2.73 1.86 2.0
p2 24 0.56 4.30 4.29 2.8
p3 40 0.53 3.06 3.88 4.6

Hypopygium (Fig. 9A, B). Tergite IX with 4 weak setae to each side of the anal point and pair of rounded lobes submedially, each with about 14 long setae, longest about 50 µm long. Anal point parallel-sided with rounded apex, 23 µm long, 12 µm wide basally, 8 µm wide medially. Tergite bands not continuous. Laterosternite IX with 1 seta. Phallapodeme 35 µm long; transverse sternapodeme 19 µm long. Gonocoxite 73 µm long. Inferior volsella straight, 43 µm long, 7 µm wide subapically, with microtrichia and 4 strong apical setae. Superior volsella projecting medially, subtriangular with two apical tubercles, 14 µm long, 7 µm wide at base, 4 µm wide subapically, covered with microtrichia and with 2 apical setae, longest 9 µm long. Median volsella consisting of single strong tubercle, about 12 µm long, with single 10 µm long setae at apex. Gonostylus curved, 101 µm long, 10 µm wide medially, 14 µm wide subapically. HR = 0.72. HV = 2.07.

Female adult and immatures

Unknown.

Distribution

(Fig. 16B). Only known from Mato Grosso State in Brazil.

Nilothauma maya sp. nov.

Figures 10A, B, 16B

Type material

Holotype male, slide-mounted: Mexico, Campeche, Calacmul, Ejido Nuevo Becan, El Chorro, 18°35'26"N, 89°15'29"W, 130 m a.s.l., 30.iv.1997, light trap, A. Contreras-Ramos et al. leg. (ZMBN).

Etymology

Named after the Maya people, who used to live in the area. The name is to be regarded as a noun in apposition.

Diagnostic characters

The male can be distinguished from its congeners by the combination of: pale brown species; wing without markings; tergite IX without setose dorsal lobe(s) or spine; anal point spatulate; superior volsella slender, curved, tapering; gonostylus curved, with strong setae on protruberance on inner margin in basal one-third.

Figure 10. 

Nilothauma maya sp. nov. adult male A hypopygium, dorsal view B hypopygium with anal point and tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Description

Male imago (n = 1). Total length 2.43 mm. Wing length 1.12 mm. Total length/wing length 2.17. Wing length/length of profemur 2.15.

Colouration. Pale brown. Wing membrane without dark markings.

Antenna. AR = 0.18. Thirteenth flagellomere 112 µm long.

Head. Temporal setae 4 in single row. Clypeus with 13 setae. Tentorium and stipes not measurable. Palp segment lengths (in µm): 21, 28, 65, 117, 144. Third palpomere with 2 sensilla clavata subapically, longest 19 µm long. Fifth palpomere/third palpomere 2.22.

Thorax. Dorsocentrals 8 in single row, acrostichals 8, prealars 2. Scutellum with 2 setae.

Wing. VR = 1.44. Brachiolum with 1 seta, R with 10 setae, R1 with 12 setae, R4+5 with 15 setae, remaining veins bare.

Legs. Spur of fore tibia 48 µm long including 21 µm long scale. Mid-tibia with 1 spur, 26 µm long; hind tibia with 2 spurs, 25 and 33 µm long. Combs of mid-tibia 12 µm long, of hind tibia 17 µm long. Width at apex of fore tibia 36 µm, of mid-tibia 40 µm, of hind tibia 44 µm. Lengths and proportions of legs as in Table 9.

Table 9.

Lengths (in μm) and proportions of leg segments in Nilothauma maya sp. nov., adult male (n = 1).

Fe Ti ta1 ta2 ta3 ta4
p1 520 372
p2 484 308 208 76 56 32
p3 504 484 232 132 140 88
ta5 LR BV SV BR
p1
p2 28 0.68 5.21 3.81 3.4
p3 48 0.48 2.99 4.26 4.7

Hypopygium (Fig. 10A, B). Tergite IX without lobes, with 8 setae above anal point and 5 somewhat weaker setae to each side of anal point. Anal point spatulate, 36 µm long, maximum width 11 µm. Tergite bands continuous. Laterosternite IX with 2 setae. Phallapodeme 48 µm long; transverse sternapodeme 27 µm long. Gonocoxite 104 µm long. Inferior volsella curved, 55 µm long, 14 µm wide subapically, with microtrichia and 4 strong apical setae. Superior volsella slender, curved, tapering, 54 µm long, 11 µm wide at base, 3 µm wide subapically, covered with microtrichia and with 2 weak apical setae. Median volsella consisting of two small tubercles, about 4 µm long, each with single setae at apex, longest 15 µm long. Gonostylus 131 µm long, curved, with single, strong setae on protuberance on inner margin at 27 µm from base, setae 21 mm long. HR = 0.79. HV = 1.85.

Female adult and immatures

Unknown.

Remarks

The slide is distorted and the drawings are composites of left and right side.

Distribution

(Fig. 16B). Only known from Campeche State in Mexico.

Nilothauma reissi (Soponis, 1987)

Figure 17C

Additional material

1 male, slide-mounted: Brazil, Santa Catarina, São Francisco do Sul, Distrito do Saí, 26°13'40"S, 48°40'50"W, CEPA Vila da Glória, 11–15.xi.2019, #143, light trap, L.C. Pinho et al. leg. 1 male, slide-mounted: Brazil, São Paulo, Santa de Rosa Viterbo, bridge at Tio Zito, 27.ix. 2000, light trap, H.F. Mendes & T. Andersen leg. 1 male, slide-mounted: Brazil, Mato Grosso, Ribeirão Cascalheira, Estrada Fazenda Manaus, 1° af. Rio Bonito, 12°57.088'S, 51°52.480'W, 08.x.2007, light trap, L.C. Pinho, S. Mateus, L. Torati & F.R. Silva leg.

Distribution

(Fig. 17C). The species was described from the Amazonas by Soponis (1987) and was later recorded from Minas Gerais and São Paulo States in northern and south-eastern Brazil by Mendes and Andersen (2009). The range is now extended to Mato Grosso and Santa Catarina States.

Nilothauma soka Andersen, Bello-González & Hagenlund, 2016

Figure 17C

Additional material

2 males, slide-mounted: Brazil, Rondônia, Candeias do Jamari, Rio Preto, Ponte de Madeira, #01, 08°52'40"S, 63°38'02"W, 19–20.vii.2012, light trap, R. Boldrini & A.S. Fernandes leg. 2 males, slide-mounted: Brazil, Roraima, Boa Vista, Rio Cauamé, 02°52'06"N, 60°44'24"W, 9.iii.2009, light trap, L.M. Fusari leg. 2 males, slide-mounted: Brazil, Amazonas, Barcelos, Rio Aracá, #9, 69 m a.s.l., 00°24'39"N, 63°23'12"W, 28.vii–06.viii.2009, light trap #3, N. Hamada et al. leg. 3 males, slide-mouted: Brazil, Amazonas, Barcelos, Rio Aracá, Foz do Igarapé Cuieiras, 00°19'15"N, 63°16'15"W, 35 m a.s.l., 30.vii–01.viii.2009, light trap #11, N. Hamada et al. leg.

Distribution

(Fig. 17C). The species was originally described from the Amazonas State by Andersen et al. (2016); the range is now extended to the Rondônia and Roraima States in the Brazilian Amazon.

Nilothauma strebulosum (Adam & Sæther, 2000)

Figure 16D

Additional material

1 male, slide-mounted: Brazil, Mato Grosso, Nova Xavantina, Fazenda Sr. Queté, Córrego Voadeira, 14°32.187'S, 52°30.902'W, 16.x.2007, light trap, L.C. Pinho, S. Mateus, L. Torati & F.R. Silva leg.

Distribution

(Fig. 16D). The species was originally described from Costa Rica by Adam and Sæther (2000); the range is now extended to Mato Grosso State, central Brazil.

Nilothauma terena sp. nov.

Figures 11A, B, 12A–H, 16A

Type material

Holotype male with larval and pupal exuvia, slide-mounted: Brazil, São Paulo, São Carlos, Campus UFSCar, Córrego do Fazzari, 21°59'S, 47°54'W, 11.ix.2008, L.C. Pinho & F.L. Silva leg. (UFSC).

Etymology

The specific epithet honours the Terena indigenous people from São Paulo State (Brazil). The name is to be regarded as a noun in apposition.

Diagnostic characters

The male can be distinguished from its congeners by having tergite IX with broadly rounded posterior margin without anal point, with anterolateral thorns, with dorsolateral lobes with few, strong setae and posteriolateral, narrowly subtriangular projection. The pupa can be recognised by having long, taeniate frontal setae and sternite I with extensive shagreen. The larva can be recognised by apparently having antenna with five segments only and by having mentum and inner teeth of mandible somewhat darker pigmented.

Figure 11. 

Nilothauma terena sp. nov. adult male A hypopygium, dorsal view B hypopygium with d tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Description

Male imago (n = 1). Total length 2.67 mm. Wing length 1.21 mm. Total length/wing length 2.21. Wing length/length of profemur 2.11.

Colouration. Thorax and legs brown, abdomen light brown. Wing membrane without dark markings.

Antenna. AR = 0.21. Thirteenth flagellomere 120 µm long.

Head. Temporal setae 5 in single row. Clypeus with 13 setae. Tentorium 62 µm long, maximum width 15 µm. Stipes not measurable. Palp segment lengths (in µm): 20, 23, 57, 92, 106. Third palpomere with 2 sensilla clavata subapically, longest about 25 µm. Fifth palpomere/third palpomere 1.87.

Thorax. Dorsocentrals 5 in single row, acrostichals 4, prealars 2. Scutellum with 2 setae.

Wing. VR = 1.46. Brachiolum with 1 seta, R with 6 setae, R1 with 1 seta, R4+5 with 1 apical seta, remaining veins bare.

Legs. Spur of fore tibia 62 µm long including 32 µm long scale. Mid-leg missing; hind tibia with 2 spurs, 28 and 46 µm long. Combs of hind tibia 18 µm long. Width at apex of fore tibia 37 µm, of hind tibia 39 µm. Lengths and proportions of legs as in Table 10.

Table 10.

Lengths (in μm) and proportions of leg segments in Nilothauma terena sp. nov., adult male (n = 1).

Fe Ti ta1 ta2 ta3 ta4
p1 572 359
p2
p3 563 574 310 155 147 90
ta5 LR BV SV BR
p1
p2
p3 49 0.57 3.22 3.58 7.1

Hypopygium (Fig. 11A, B). Tergite IX with rounded posterior margin with altogether 6 marginal setae in two posteriolateral groups; with dorsolateral lobes with 5 strong setae, longest setae 65 µm long; with posteriolateral, narrowly subtriangular projections, 14 µm long, 6 µm wide at base. Tergite band not apparent. Laterosternite IX without setae; with small anteriolateral thorn. Phallapodeme 33 µm long; transverse sternapodeme 11 µm long. Gonocoxite 83 µm long. Inferior volsella digitiform, curved, 55 µm long, 11 µm wide medially, with microtrichia and 12 short setae subapically. Superior volsella 29 µm long, 12 µm wide at base, 13 µm wide medially, covered with microtrichia and with marginal setae. Median volsella subtriangular with 2 apical setae on small tubercles, setae about 6 µm long. Gonostylus weakly curved, 97 µm long, 14 µm wide medially. HR = 0.85. HV = 2.75.

Female adult

Unknown.

Pupa

(n = 1). Total length 3.59 mm. Exuviae pale brown.

Cephalothorax (Fig. 12A, B). Frontal apotome (Fig. 12A) with few wrinkles, frontal setae taeniate, 154 μm long. Thoracic horn not discernible; basal ring oval, 13 μm in diameter. Scutum with field of few weak tubercles. Antepronotals 2; precorneals 2; dorsocentrals 4, Dc1 39 μm in front of Dc2, Dc2 96 μm in front of Dc3, Dc3 23 μm in front of Dc4.

Figure 12. 

Nilothauma terena sp. nov. pupa (A–E) and larva (F–H) A frontal apotome B thorax C abdomen, dorsal view D sternite I E paratergite VIII F antenna G mandible H mentum and ventromental plates.

Abdomen (Fig. 12C–E). Tergite I bare; tergites II–VI with transverse anterior band of somewhat stronger spinules, merging with median field of finer shagreen; anterior band of shagreen on tergite VI separated from posterior shagreen patch; tergite VII with anterior and posterior shagreen patches; tergite VIII with anterior shagreen patch connected with narrow posterior field of finer shagreen; tergite IX bare. Sternite I (Fig. 12D) with extensive shagreen; sternite II–VII bare; sternite VIII with narrow, longitudinal field of fine shagreen. Tergite II with 159 μm long row of 36 hooks, each hook about 8 μm long. Conjunctives III/IV and IV/V with spinules extending on to preceding segment. Pedes spurii B weakly developed on segment II. Anal comb 51 μm long, consisting of 3 spurs.

Abdominal setation. Lateral setae on segments I–VIII as: 0, 3, 3, 3, 4, 4, 4, 4; posterior lateral seta on tergite IV and all lateral setae on tergites V–VIII taeniate, remaining setae hair-like. All tergites with 1 pair of O setae.

Anal lobe. As long as broad, with 1 taeniate dorsal setae and complete fringe of 19 taeniae on each side, longest 170 μm. Male genital sac over-reaches anal lobe by 119 μm.

Fourth instar larva

(n = 1). Head capsule 228 μm long. Postmentum 145 μm long.

Head. Antenna (Fig. 12F) apparently with five segments only, length of antennal segments (in μm): 19, 10, 4, 11, 6. AR = 0.61. Basal antennal segment 10 μm wide; blade 32 μm long; accessory blade about 5 μm long. Premandible not measurable, teeth not discernible. Mandible (Fig. 12G) 69 μm long, seta subdentalis 19 μm long, inner teeth somewhat darker pigmented. Mentum (Fig. 12H) somewhat darker pigmented, 41 μm wide; middle part 10 μm wide with 2 minute inner teeth and pair of slightly larger lateral teeth; with 6 pairs of pointed, medially curved lateral teeth. Ventromental plates 98 μm wide, medially separated by 10 μm. Seta submenti 46 μm long.

Abdomen. Lost.

Distribution

(Fig. 16A). Known from São Paulo State, south-eastern Brazil.

Nilothauma txukuyana sp. nov.

Figures 13A, B, 16B

Type material

Holotype male, slide-mounted: Brazil, Pará, Rio Paru do Oeste, Malloca Apicó, 20.iv.1962, at light, E.J. Fittkau leg. (A 366-1, ZSM). Paratypes: 15 males, same data as holotype (ZSM, ZMBN, UFSC).

Etymology

The specific epithet honours the Txukuyana, indigenous people from Amazonas and Pará States in Brazil and from Suriname. The name is to be regarded as a noun in apposition.

Diagnostic characters

The male can be distinguished from its congeners by having tergite IX with broadly-rounded posterior margin without anal point, with dorsolateral lobes with few, strong setae and posteriolateral, strongly setose, subtriangular projection.

Figure 13. 

Nilothauma txukuyana sp. nov. adult male A hypopygium, dorsal view B hypopygium with tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Description

Male imago (n = 5–8). Total length 2.17–2.44, 2.25 mm. Wing length 1.00–1.09, 1.05 mm. Total length/wing length 2.02–2.33, 2.16. Wing length/length of profemur 2.16–2.26, 2.21.

Colouration. Head, thorax and legs brown; abdomen light brown. Wing membrane without dark markings.

Antenna. AR = 0.26–0.29, 0.28. Thirteenth flagellomere 132–156, 145 µm long.

Head. Temporal setae 6–10, 7 in single row. Clypeus with 13–16, 15 setae. Tentorium 69–83, 77 µm long, maximum width 14–19, 19 µm. Stipes not measurable. Palp segment lengths (in µm): 18–25, 21; 23–28, 26; 56–60, 57; 76–81, 79; 102–115, 107. Third palpomere with 2 sensilla clavata subapically, longest about 15 µm long. Fifth palpomere/third palpomere 1.72–1.98, 1.85.

Thorax. Dorsocentrals 4–7, 6 in single row, acrostichals apparently 4–6, 5 anterior, prealars 1–2, 2. Scutellum with 2 setae.

Wing. VR = 1.44–1.54, 1.50. Brachiolum with 1 seta, R with 7–9, 8 setae, R1 with 5–8, 7 setae, R4+5 with 9–13, 11 setae apically, remaining veins bare.

Legs. Spur of fore tibia 48–54, 52 µm long including 20–25, 22 µm long scale. Mid-tibia with 1 spur, 29–35, 32 µm long; hind tibia with 2 spurs, 22–28, 25 and 33–39, 36 µm long. Combs of mid-tibia 17–18, 18 µm long, of hind tibia 19–22, 21 µm long. Width at apex of fore tibia 33–37, 35 µm, of mid-tibia 33–38, 36 µm, of hind tibia 39–41, 40 µm. Lengths and proportions of legs as in Table 11.

Table 11.

Length (in µm) and proportions of legs of Nilothauma txukuyana sp. nov., adult males (n = 5–7).

Fe Ti ta1 ta2
p1 457–523, 482 319–359, 338 474–547, 515 194–221, 203
p2 449–474, 462 286–310, 301 147–179, 162 65–74, 69
p3 507–556, 529 458–482, 467 245–278, 263 114–139, 127
ta3 ta4 ta5 LR
p1 147–163, 157 106–123, 114 57–65, 59 1.49–1.59, 1.52
p2 41–49, 46 25–33, 29 24–32, 26 0.50–0.58, 0.54
p3 114–147, 129 73–90, 78 41–49, 47 0.54–0.58, 0.56
BV SV BR
p1 2.47–2.54, 2.51 1.54–1.64, 1.59 2.43–3.46, 2.71
p2 5.13–5.73, 5.45 4.36–5.06, 4.73 2.77–3.76, 3.38
p3 3.20–3.57, 3.30 3.70–4.00, 3.79 4.33–5.00, 4.60

Hypopygium (Fig. 13A, B). Tergite IX with rounded posterior margin with 4–7, 5 marginal setae; with dorsolateral lobes with 3–4, 4 strong setae, longest setae 44–55, 50 µm long; with posteriolateral subtriangular, strongly setose projection, 35–41, 38 µm long, 22–25, 24 µm wide at base. Tergite band not apparent. Laterosternite IX with single setae. Phallapodeme 47–55, 49 µm long; transverse sternapodeme 17–19, 18 µm long. Gonocoxite 83–89, 87 µm long. Inferior volsella digitiform, curved, 62–72, 67 µm long, 11–14, 12 µm wide medially, with microtrichia and 13–17, 15 setae subapically. Superior volsella 25–29, 27 µm long, 8–11, 10 µm wide at base, 7–10, 9 µm wide medially, covered with microtrichia and with marginal setae. Median volsella small, broadly triangular, apparently without setae. Gonostylus nearly straight, 126–135, 130 µm long, 17–19, 18 µm wide medially. HR = 0.65–0.68, 0.66. HV = 1.78–1.87, 1.82.

Female imago and immatures

Unknown.

Distribution

(Fig. 16B). Known from Pará State, Brazil.

Nilothauma werekena sp. nov.

Figures 14A–C, 16D

Type material

Holotype male, slide-mounted: Brazil, Amazonas, Barcelos, Rio Aracá, Foz do Igarapé Cuieiras, #11, 00°19'15"N, 63°16'15"W, 35 m a.s.l., 30.vii–01.viii.2009, light trap, N. Hamada et al. leg. (UFSC). Paratypes: 4 males, slide-mounted, same data as holotype (INPA). 2 males, slide-mounted, same data as previous, except: #9, 00°24'39"N, 63°23'12"W, 69 m a.s.l., 28.vii–06.viii.2009, light trap #3, N. Hamada et al. leg (MZSP).

Figure 14. 

Nilothauma werekena sp. nov. adult male A hypopygium, dorsal view B superior volsella, dorsal view C hypopygium with anal point and tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Etymology

The specific epithet honours the Werekena indigenous people from the Rio Negro Basin in the Amazon. The name is to be regarded as a noun in apposition.

Diagnostic characters

The male can be distinguished from its congeners by the combination of: tergite IX with one low, but wide median dorsal protruberance with about 30 strong setae; anal point spatulate; superior volsella covered with microtrichia, fused to median volsella; laterosternite IX with thorn.

Description

Male imago (n = 5–7, unless otherwise stated). Total length 1.89–2.26, 2.11 mm. Wing length 0.98–1.11, 1.05 mm. Total length/wing length 1.90–2.15, 1.99. Wing length/length of profemur 2.21–2.53, 2.37.

Colouration. Head, thorax, legs and abdomen uniformly brown. Wing membrane without dark markings.

Antenna. AR = 0.32–0.39, 0.35. Thirteenth flagellomere 230–274, 260 µm long.

Head. Temporal setae 7–8, 7 in single row. Clypeus with 10–14, 12 setae. Tentorium 47–75, 65 µm long, maximum width 12–20, 17 µm. Stipes 80–117, 100 (4) µm long. Palp segment lengths (in µm): 20–32, 27; 22–27, 25; 65–85, 72; 95–125, 110; 87–132, 115. Third palpomere with 2–5, 4 sensilla clavata subapically, longest 12–15, 14 µm long. Fifth palpomere/third palpomere 1.21–1.75, 1.54.

Thorax. Dorsocentrals 6–8, 7 in single row, acrostichals 10–14, 12, prealars 2. Scutellum with 2 setae.

Wing. VR = 1.21–1.32, 1.27. Brachiolum with 1 seta, R with 11–12, 11 setae, R1 with 5–8, 7 setae, R4+5 with 11–17, 15 setae, remaining veins bare.

Legs. Spur of fore tibia 39–44, 42 µm long including 15–20, 17 µm long scale. Mid-tibia with 1 spur, 20–25, 22 µm long; hind tibia with 2 spurs, 20–25, 23 and 28–31, 30 µm long. Combs of mid-tibia 17–19, 18 µm long, of hind tibia 18–21, 19 µm long. Width at apex of fore tibia 29–39, 34 µm, of mid-tibia 34–39, 37 µm, of hind tibia 34–44, 39 µm. Lengths and proportions of legs as in Table 12.

Table 12.

Lengths (in μm) and proportions of leg segments in Nilothauma werekena sp. nov., adult males (n = 5–7, unless otherwise stated).

Fe Ti ta1 ta2
p1 374–501, 433 315–394, 345 443–522, 473 246–286, 266
p2 345–463, 424 296–345, 325 177–207, 197 89–99, 94
p3 443–532, 493 463–522, 502 266–305, 286 138–158, 148
ta3 ta4 ta5 LR
p1 187–217, 207 118–148, 138 (4) 69–89, 79 (4) 1.36–1.47, 1.40
p2 69–79, 74 39–49, 44 30–39, 35 0.56–0.63, 0.60
p3 128–158, 148 (4) 89–99, 94 (4) 59–69, 55 (4) 0.57–0.61, 0.59
BV SV BR
p1 1.84–2.20, 1.95 1.55–1.66, 1.60 2.0–3.3, 2.5
p2 3.50–4.25, 3.81 3.42–4.05, 3.71 2.3–4.7, 3.3
p3 2.70–2.89, 2.81 (4) 3.39–3.45, 3.41 5.0–7.0, 5.9

Hypopygium (Fig. 14A–C). Tergite IX without dorsal lobe(s), with low, but wide median dorsal protruberance with 29–32, 31 strong, clustered median setae; posterior margin rounded to subrectangular, with 8–11, 9 weak setae to each side of base of anal point. Anal point spatulate, 37–47, 40 µm long, maximum width 5–10, 7 µm. Tergite bands not continuous. Laterosternite IX with 1–2, 1 seta; with anterolateral thorn. Phallapodeme 40–50, 45 µm long; transverse sternapodeme 15–25, 20 µm long. Gonocoxite 62–82, 72 µm long. Inferior volsella strongly curved, 22–30, 27 µm long, 5–8, 6 µm wide medially, with microtrichia and 3 strong, simple setae apically. Superior volsella tongue-shaped to slightly pediform, 32–40, 37 µm long, 15–25, 20 µm wide at base, densely covered with microtrichia. Median volsella fused to superior volsella, consisting of 2–3, 2 small tubercles, each bearing single, long seta. Gonostylus 70–100, 90 µm long, basal half curved, distal half straight. HR = 0.63–0.96, 0.83. HV = 2.26–2.70, 2.34.

Female adult and immatures

Unknown.

Distribution

(Fig. 16D). Known from Barcelos (Amazonas State), in the Brazilian Amazon.

Nilothauma yekwana sp. nov.

Figures 15A, B, 16A

Type material

Holotype male, slide-mounted: Brazil, Roraima, Boa Vista, BR-174, Igarapé Água Boa, 02°43'32"N, 60°48'43"W, 2014, N. Hamada leg (UFSC).

Etymology

The specific epithet honours the Ye’kwana, indigenous people from the Roraima State, Brazil. The name is to be regarded as a noun in apposition.

Figure 15. 

Nilothauma yekwana sp. nov. adult male A hypopygium, dorsal view B hypopygium with tergite IX removed, dorsal aspect to the left and ventral aspect to the right.

Diagnostic characters

The male can be distinguished from its congeners by the combination of: tergite IX with two setose dorsolateral lobes; anal point absent; posterior margin of tergite IX subrectangular; inferior volsella straight, tapering to apex; superior volsella curved, projecting posteriolaterally; median volsella broad, triangular, bearing 7 strong setae.

Description

Male imago (n = 1). Total length 1.77 mm. Wing length 0.91 mm. Total length/wing length 1.95. Wing length/length of profemur 2.19.

Colouration. Head, thorax, legs and abdomen uniformly light brown. Wing membrane without dark markings.

Antenna. AR = 0.22. Thirteenth flagellomere 137 µm long.

Head. Temporal setae 7 in single row. Clypeus with 15 setae. Tentorium 65 µm long, maximum width 12 µm. Stipes 80 µm long. Palp segment I–III lengths (in µm): 22, 17, 50; segment IV and V lost. Third palpomere with 2 sensilla clavata subapically, longest 15 µm.

Thorax. Dorsocentrals 5 in single row, acrostichals 10, prealars 2. Scutellum with 2 setae.

Wing. VR = 1.48. Brachiolum with 1 seta, R with 6 setae, R4+5 with 2 setae at apex, remaining veins bare.

Legs. Spur of fore tibia 44 µm long including 15 µm long scale. Mid-tibia with 1 spur, 20 µm long; hind tibia with 2 spurs, 20 and 25 µm long. Combs of mid-tibia 15 µm long, of hind tibia 18 µm long. Width at apex of fore tibia 39 µm, of mid-tibia 39 µm, of hind tibia 44 µm. Lengths and proportions of legs as in Table 13.

Table 13.

Lengths (in μm) and proportions of leg segments in Nilothauma yekwana sp. nov., adult male (n = 1).

Fe Ti ta1 ta2 ta3 ta4
p1 414 296
p2 394 256 177 59 39 30
p3 443 414 217 99 99 79
ta5 LR BV SV BR
p1
p2 30 0.57 5.25 3.67 2.0
p3 49 0.52 3.60 3.95 4.4

Hypopygium (Fig. 15A, B). Tergite IX with 2 dorsolateral, densely setose lobes, setae about 15 µm long; with 2 strong setae anterolaterally, 2 medially and 4 close to posterior margin; posterior margin subquadrangular, anal point absent. Tergite bands lacking. Laterosternite IX with 1 seta. Phallapodeme 37 µm long; transverse sternapodeme 10 µm long. Gonocoxite 65 µm long. Inferior volsella straight, tapering to apex, 40 µm long, 7 µm wide medially, with microtrichia and 9 simple setae subapically. Superior volsella curved, projecting posteriolaterally, 22 µm long, 5 µm wide at base, covered with microtrichia and fringed at apex. Median volsella broad, triangular, 15 µm long, with 7 strong setae (one of them bifid), longest 20 µm. Gonostylus 75 µm long, straight. HR = 0.87. HV = 2.36.

Female adult and immatures

Unknown.

Distribution

(Fig. 16A). Known from Roraima State, Brazilian Amazon.

Nilothauma zitoi Mendes & Andersen, 2009

Figure 16D

Additional material

1 male, slide-mounted: Brazil, Mato Grosso, Ribeirão Cascalheira, Fazenda Campina Verde, Rio Suiá Miçu, 12°48.591'S, 52°06.925'W, 10.x.2007, light trap, L.C. Pinho, S. Mateus, L. Torati & F.R. Silva leg.

Figure 16. 

Distribution maps of Neotropical Nilothauma species A N. hamadae sp. nov., N. jupau sp. nov., N. terena sp. nov., N. yekwana sp. nov., N. complicatum Mendes & Andersen, 2009 B N. leccii sp. nov., N. marianoi sp. nov., N. mateusi sp. nov., N. txukuyana sp. nov., N. maya sp. nov. C N. karitiana sp. nov., N. duena Roback, 1960, N. calori Mendes & Andersen, 2009 D N. werekena sp. nov., N. zitoi Mendes & Andersen, 2009, N. strebulosum (Adam & Sæther, 2000).

Distribution

(Fig. 16D). The species was originally described by Mendes and Andersen (2009), based on a single male from São Paulo State; the range is now extended to the Mato Grosso State.

Figure 17. 

Distribution maps of Neotropical Nilothauma species A N. fittkaui (Soponis, 1987), N. amazonense Mendes & Andersen, 2009 B N. aripuanense Mendes & Andersen, 2009, N aleta Roback, 1960 C N. reissi (Soponis, 1987), N. soka Andersen, Bello-González & Hagenlund, 2016 D N. anamariae Dantas & Hamada, 2017, N. jaraguaense Mendes & Andersen, 2009.

Key to the males of Nilothauma Kieffer of the world

Modified from Qi et al. (2014), Niitsuma (2016) and Andersen et al. (2016), with the inclusion of sixteen species.

1 Tergite IX without setose dorsal lobe(s) or projection(s) 2
Tergite IX with one to four setose dorsal lobes or projection(s) (e.g. Figs 16, 20) 19
2 Anal point present 3
Anal point absent 15
3 Tergite IX with median cluster of about 30 strong setae 4
Tergite IX with few, clustered setae, if numerous they are scattered (as in N. aripuanense) 5
4 Superior volsella slender, without microtrichia and with lateral spine. Brazil Nilothauma jaquei Dantas & Hamada, 2017
Superior volsella pediform to lingulate, covered with microtrichia and without lateral spine (Fig. 14). Brazil Nilothauma werekena sp. nov.
5 Wing with conspicuous dark markings (Fig. 5); abdominal tergites II, III, and VI–VIII dark brown. Brazil Nilothauma jupau sp. nov.
Wing unmarked, at most with faint colour (as in Nilothauma aleta, best seen in dark-field filter); abdominal tergites uniformly pale to brown 6
6 Gonostylus stout or swollen (Figs 1, 3, 6, 7) 7
Gonostylus slender (Fig. 10) 10
7 Gonostylus very long, narrow basally and apically, swollen at mid-length (Fig. 6). Brazil Nilothauma karitiana sp. nov.
Gonostylus stout, not distinctly swollen at mid-length (Figs 1, 3) 8
8 Acrostichals absent; anal point wide, covering most setae along posterior margin of tergite IX (Fig. 1). Peru, Brazil Nilothauma aleta Roback, 1960
Acrostichals present; anal point comparatively narrow, nearly parallel-sided, with most setae placed lateral to base of anal point 9
9 Superior volsella tapering to apex; inferior volsella short, stout, with short, simple setae (Fig. 3). Peru, Brazil Nilothauma duena Roback, 1960
Superior volsella wider at mid-length; inferior volsella long and slender, with long simple or apically split setae (Fig. 7). Brazil Nilothauma leccii sp. nov.
10 Superior volsella narrow, straight, curved or weakly sinuous, projecting posterio-medially, with one to six apical setae 11
Superior volsella wider in distal half, projecting posterio-medially or posterio-laterally, with microtrichia only 13
11 Tergite IX with numerous scattered setae; anal point broadly lanceolate, about 20 µm wide. Brazil Nilothauma aripuanense Mendes & Andersen, 2009
Tergite IX with one to four median setae and about 12 setae along posterior margin; anal point comparatively narrow 12
12 Tergite IX with single seta anterior to anal point; gonostylus with two to four sub-basal dorsal setae not arising from protuberances. Brazil Nilothauma paucisetis Dantas & Hamada, 2017
Tergite IX with four aligned setae anterior to anal point; gonostylus with single sub-basal seta arising from distinct inner protuberance (Fig. 10). Neotropical Mexico Nilothauma maya sp. nov.
13 Anal point parallel-sided, about 4 µm wide; inferior volsella with about 18 slender, simple setae (Fig. 8). Brazil Nilothauma marianoi sp. nov.
Anal point spatulate, 15–23 µm wide; inferior volsella with few (about 3) simple, slender setae or numerous (about 12) stout, split setae apically 14
14 Superior volsella boot-shaped, projecting posterior-laterally; inferior volsella narrow, with few simple, slender setae apically. Brazil Nilothauma soka Andersen, Bello-González & Hagenlund, 2016
Superior volsella straight, projecting posterior-medially; inferior volsella wide, with numerous stout, split setae apically. Brazil Nilothauma anamariae Dantas & Hamada, 2017
15 Inferior volsella branched subapically. Brazil Nilothauma complicatum Mendes & Andersen, 2009
Inferior volsella simple 16
16 Superior volsella pediform, without ventral transverse fold, with setae and microtrichia 17
Superior volsella diamond-shaped, with ventral transverse fold, with microtrichia only 18
17 Wing vein R1 with setae; gonostylus nearly parallel-sided in apical half. Brazil, Ecuador Nilothauma fittkaui (Soponis, 1987)
Wing vein R1 bare; gonostylus widest in apical one-third. Brazil Nilothauma reissi (Soponis, 1987)
18 Apex of superior volsella projecting caudad. Brazil Nilothauma sooretamense Mendes & Andersen, 2009
Apex of superior volsella projecting mesad. Brazil Nilothauma involucrum Mendes & Andersen, 2009
19 Anal point lacking or rudimentary, completely covered by microtrichia 20
Anal point present, with microtrichia at most in basal half 22
20 Tergite IX with four lobes or projections, one anterior pair with strong, long setae and one posterio-lateral, triangular pair with weaker setae 21
Tergite IX with one or two dorsal lobes 27
21 Tergite IX with posterio-lateral pair of projections short, subequal in length to anterior pair (Fig. 11); laterosternite IX with thorn. Brazil N. terena sp. nov.
Tergite IX with posterio-lateral pair of projections long, more than three times longer than anterior pair (Fig. 13); laterosternite IX without thorn. Brazil N. txukuyana sp. nov.
22 Dorsal projections of tergite IX differ in shape. Ghana Nilothauma insolitum Adam & Sæther, 1999
Dorsal projections of tergite IX of the same shape 23
23 Median volsella fused to superior volsella; superior volsella broadly pediform. Brazil Nilothauma fazzariense Mendes & Andersen, 2009
Median volsella distinct and separated from superior volsella; superior volsella digitate, curved, with or without lateral spine 24
24 Dorso-lateral projections of tergite IX overreaching posterior margin of tergite. Brazil Nilothauma roquei Mendes & Andersen, 2009
Dorso-lateral projections of tergite IX not extended beyond posterior margin of tergite 25
25 Superior volsella with lateral spine; laterosternite IX with thorn; posterior margin of tergite IX broadly rounded. Brazil Nilothauma calori Mendes & Andersen, 2009
Superior volsella without lateral spine; laterosternite IX without thorn; posterior margin of tergite IX subrectangular 26
26 Median volsella consisting of single, small tubercle bearing one seta; inferior volsella curved, not tapering to apex. Brazil, Costa Rica Nilothauma strebulosum (Adam & Sæther, 2000)
Median volsella broad, triangular, bearing 7 strong setae; inferior volsella straight, tapering to apex (Fig. 15). Brazil Nilothauma yekwana sp. nov.
27 Tergite IX with single, median setose dorsal lobe 28
Tergite IX with two setose dorsal lobes 37
28 Superior volsella without microtrichia 29
Superior volsella covered with microtrichia 32
29 Dorsal projection weakly developed and undivided; situated posteriorly on tergite IX, close to base of anal point. China, Japan, Thailand Nilothauma japonicum Niitsuma, 1985
Dorsal projection well developed, divided or undivided, situated anteriorly on tergite IX, at some distance from base of anal point 30
30 Dorsal projection three-pronged at apex, without setae. Ghana Nilothauma fuscina Adam & Sæther, 1999
Dorsal projection bell-shaped, with numerous setae 31
31 Superior volsella with single apical seta; anal point narrow. Brazil Nilothauma matogrossense Mendes & Andersen, 2009
Superior volsella with one dorsolateral and one apical spine-like seta; anal point broad. Ghana Nilothauma duminola Adam & Sæther, 1999
32 Anal point very broad, about half as wide as tergite IX and lanceolate (Fig. 4). Brazil Nilothauma hamadae sp. nov.
Anal point narrow, spatulate to tapering, but not lanceolate 33
33 Anal point tapering or only slightly widened medially 34
Anal point distinctly spatulate 35
34 Wing with distinct dark areas at RM, FCu, along apical half of An and in cells r4+5 and m1+2. Cuba Nilothauma granma Andersen, Bello-González & Hagenlund, 2016
Wing without dark areas. Canada, USA Nilothauma babiyi (Rempel, 1937)
35 Dorsal projection large, covering most of tergite IX. Australia Nilothauma adunatum Adam & Sæther, 1999
Dorsal projection of tergite IX small, with setae at apex only 36
36 Wing length > 2.4 mm; AR = 0.28; median volsella consisting of two tubercles, each with single, strong apical seta. Canada Nilothauma verrucum Adam & Sæther, 1999
Wing length < 1.3 mm; AR = 0.13; median volsella consisting of three tubercles, each with strong, apical seta. Venezuela Nilothauma canaima Andersen, Bello-González & Hagenlund, 2016
37 Dorsal projections of tergite IX of the same shape 38
Dorsal projections of tergite IX of different shapes 42
38 Anal point short, digitiform, with microtrichia in basal half; gonostylus distinctly widened in apical one-third. Brazil Nilothauma zitoi Mendes & Andersen, 2009
Anal point well developed, lanceolate or parallel-sided; gonostylus nearly parallel-sided in apical half 39
39 Superior volsella short, subtriangular, with two apical tubercles bearing setae; inferior volsella with long, strong simple setae (Fig. 9). Brazil Nilothauma mateusi sp. nov.
Superior volsella long, pediform to tongue-shaped; inferior volsella with slender, simple apically split setae 40
40 Anal point parallel-sided; laterosternite IX with thorn. Chile Nilothauma spiesi Mendes & Andersen, 2009
Anal point lanceolate; laterosternite IX without thorn 41
41 Inferior volsella and gonostylus with apically split setae; median volsella curved, tapering, with microtrichia and setae. Brazil Nilothauma jaraguaense Mendes & Andersen, 2009
Inferior volsella and gonostylus with simple setae only; median volsella short, parallel-sided, with two apical setae, without microtrichia. Brazil Nilothauma amazonense Mendes & Andersen, 2009
42 Wing with dark areas or bands; anterior projection on tergite IX long and deeply divided 43
Wing without dark markings; anterior projection on tergite IX variably developed 50
43 Anterior projection on tergite IX with setae not restricted to apex. South Africa Nilothauma harrisoni Adam & Sæther, 1999
Anterior projection on tergite IX with apical setae only 44
44 Posterior projection on tergite IX deeply divided into antero-dorsal and postero-ventral parts 45
Posterior projection on tergite IX not as above 47
45 Anterior projection on tergite IX with simple, separated, apical setae; anterior part of posterior projection apically pointed; wing with postero-median spot extending on both sides of Cu1. Afrotropical Nilothauma pictipenne Kieffer, 1921
Anterior projection on tergite IX with setae forming fan-like structure; anterior part of posterior projection with blunt apex; wing with postero-median spot exclusively proximal of Cu1 46
46 Wing with four dark areas; setae on anterior projection on tergite IX branched apically. Ghana Nilothauma flabellatum Adam & Sæther, 1999
Wing with three dark areas; apical setae on anterior projection on tergite IX lamellate, not branched. Ghana Nilothauma kakumense Adam & Sæther, 1999
47 Posterior projection on tergite IX either with a disto-dorsal lobe or subapical constriction 48
Posterior projection on tergite IX without distal lobe or constriction 49
48 Posterior projection on tergite IX with long antero-lateral arms and disto-dorsal lobe. Tanzania Nilothauma anderseni Adam & Sæther, 1999
Posterior projection on tergite IX without antero-lateral arms, with apical constriction and five apical setae. Zimbabwe Nilothauma latocaudatum Adam & Sæther, 1999
49 Posterior projection on tergite IX without microtrichia, except on long, antero-lateral arms; superior volsella without antero-median extension; median volsella present. China, Japan Nilothauma nojirimaculatum Sasa, 1991
Posterior projection on tergite IX without antero-lateral arms, covered with microtrichia; superior volsella with antero-median extension; median volsella apparently absent. Japan Nilothauma hibaraquartum Sasa, 1993
50 Anterior projection on tergite IX with apically plumose setae 51
Apical setae on anterior projection on tergite IX not plumose 58
51 Anal point with microtrichia along median ridge and apical margin. Oriental China Nilothauma pandum Qi, Lin, Wang & Shao, 2014
Anal point without microtrichia 52
52 Superior volsella pad-like, with extensive microtrichia. Palaearctic Japan Nilothauma hibaratertium Sasa, 1993
Superior volsella slender, microtrichia absent or limited within small area when present 53
53 Superior volsella without lateral spur. Thailand Nilothauma mergae Adam & Sæther, 1999
Superior volsella with lateral spur 54
54 Superior volsella without setal brush or fringe, with only one to three apical setae 55
Superior volsella with apical setal brush or fringe 56
55 Anal point with microtrichia along median ridge; superior volsella relatively long when compared to median volsella (length ratio, Svo/Mvo > 4.0); inferior volsella with simple setae only. Oriental China Nilothauma aristatum Qi, Tang & Wang, 2016
Anal point bare; length ratio Svo/Mvo about 2.0; inferior volsella with apically split setae. Oriental China Nilothauma acre Adam & Sæther, 1999
56 Superior volsella four to five times as long as median volsella. Palaearctic Japan Nilothauma niidaensis Niitsuma, 2016
Superior volsella two to three times as long as median volsella 57
57 Anterior projection on tergite IX two to four times as long as broad. Canada, USA Nilothauma bicorne (Townes, 1945)
Anterior projection on tergite IX broader than long. USA Nilothauma mirabile (Townes, 1945)
58 Anal point trifid; anterior projection on tergite IX very long, tapering to parallel-sided apex, with apical setae only; posterior projection on tergite IX distally very slender, with five apical setae. D. R. Congo, Ghana Nilothauma burmeisteri Adam & Sæther, 1999
Anal point simple; anterior projection on tergite IX wart-like, with setae not only at apex; posterior projection on tergite IX triangular or apically rounded 59
59 Posterior projection on tergite IX apically rounded; superior volsella with two to four lobes 60
Posterior projection on tergite IX triangular; superior volsella without lobes 61
60 Anterior projection on tergite IX partially divided apically; superior volsella with two lobes. Oriental China Nilothauma bilobatum Qi, Tang & Wang, 2016
Anterior projection on tergite IX undivided apically; superior volsella with four lobes. Oriental China Nilothauma quatuorlobum Yan, Tang & Wang, 2005
61 Anal point parallel-sided; anterior projection on tergite IX with setae thickened at apices. Ghana Nilothauma ankasense Adam & Sæther, 1999
Anal point spatulate; anterior projection on tergite IX with setae not thickened at apices 62
62 Superior volsella tapering, widest near base. Europe Nilothauma brayi (Goetghebuer, 1921)
Superior volsella widest about one-third from apex. Australia Nilothauma infissum Adam & Sæther, 1999

Acknowledgements

We are indebted to Dr. Neusa Hamada and all her team at INPA for providing rich material from the Amazon and to Dr. Martin Spies for the loan of material from the Zoologische Staatssammlung München, Germany. Thanks are also due to Dr. Sidnei Mateus, Dr. Marcia R. Spies, Dr. Adolfo R. Calor and the staff at the Laboratory of Systematics of Diptera (UFSC) for all help and for providing us with freshly collected material. Fieldwork in Santa Catarina was funded by the Santa Catarina Research Foundation (FAPESC) – project “Insetos aquáticos do Parque Estadual da Serra Furada, SC” (FAPESC 11323/2012–9) and by the City Hall of São Francisco do Sul – project “Nascentes do Saí”.

References

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