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Two new species of Dasyhelea Kieffer and the immature of D. azteca Huerta & Grogan from northwestern Argentina (Diptera: Ceratopogonidae)
Zoologischer Anzeiger 272 (2018) 6–19
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Research Paper
Two new species of Dasyhelea Kieffer and the immature of D. azteca Huerta & Grogan from northwestern Argentina (Diptera: Ceratopogonidae) Florentina Díaz a,∗ , Gustavo Ricardo Spinelli b , María Marcela Ronderos a a Centro de Estudios Parasitológicos y de Vectores (CEPAVE), Facultad de Ciencias Naturales y Museo (UNLP), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Boulevard 120 S/N e/61 y 62 La Plata, Buenos Aires, Argentina b Instituto de Limnología “Dr. Raúl A. Ringuelet”(ILPLA), Facultad de Ciencias Naturales y Museo (UNLP), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Boulevard 120 S/N e/61 y 62 La Plata, Buenos Aires, Argentina
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Article history: Received 2 January 2017 Received in revised form 12 November 2017 Accepted 15 November 2017 Available online 16 November 2017 Corresponding Editor: Sven Bradler
a b s t r a c t Two new species of Dasyhelea Kieffer, D. yunga and D. allegrae, placed in the grisea and mutabilis species groups respectively, are described, illustrated and photographed based on pupae and male and female adults. In addition, the immatures of D. azteca are described for the first time. The specimens were collected in the southernmost area of the Argentinean Yungas. The pupae of D. yunga and D. allegrae were examined using a Compound Microscope (CM), while the larva and pupa of D. azteca by a Scanning Electron Microscope (SEM) and CM. © 2017 Elsevier GmbH. All rights reserved.
Keywords: Dasyhelea New species Immatures Adult Yungas Argentina
1. Introduction Dasyhelea Kieffer 1911 is a large and complex genus of Ceratopogonidae with diverse morphology and biology which occurs worldwide in a variety of small aquatic habitats (Waugh and Wirth, 1976). Taxonomically, the recognition of subgenera and/or species groups is still incipient and generally has been applied intermittently only to various regional faunas (Diaz et al., 2014). Borkent (2016) listed 77 species for the Neotropics, of which only 12 are known also from their immatures. The purpose of this paper is to describe two new species and the immatures of D. azteca Huerta & Grogan from specimens collected during entomological surveys carried out recently in the southernmost area of the Argentinean Yungas, a cloudy forest that extends on the east sloppes of the Andes and subandean chains, from Venezuela to northwestern Argentina, between 500 m and 2.500/3.500 m of altitude. It is considered by certain authors as a biogeographic province (Cabrera and Willink, 1973; Morrone,
∗ Corresponding author. E-mail address: [email protected] (F. Díaz). https://doi.org/10.1016/j.jcz.2017.11.008 0044-5231/© 2017 Elsevier GmbH. All rights reserved.
2001) or as an ecoregion by others (Brown et al., 2002, 2006), and although it is characterized by a high diversity within a great number of taxa as well by a high degree of endemisms (Cabrera and Willink, 1973; Brown et al., 2002), its ceratopogonid fauna is very poorly known. 2. Material and methods Pupae of two new species were collected in summer 2014 and D. azteca in autumn 2009 and spring 2011, 2014, using a pipette from flooded soil and with drift nets in several localities of the Yungas and in the provinces of Corrientes and Buenos Aires. Specimens were carried to the laboratory, larvae placed individually in Petri dishes and pupae isolated in a vial with a drop of water and observed daily until adult emergence. Adults were allowed to harden for 24 h before being preserved to ensure their pigmentation was complete. Larval, pupal exuviae and adults were mounted in Canada balsam following the technique described by Borkent and Spinelli (2007). For Scanning Electron Microscope (SEM), larva and pupae of D. azteca were prepared following the technique of Ronderos et al. (2000, 2008). Ink illustrations were made using an attached camera lucida. Photographs were taken with a digital camera Micrometrics
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SE Premium, through a Nikon Eclipse E200 microscope. Measurements were taken using SEM and Compound Microscope (CM). For terminology of larva see Díaz et al. (2013), Borkent (2014) for pupa and Brown et al. (2009) for adult. The studied specimens are deposited in the collection of the División Entomología, Museo de La Plata, Argentina (MLPA). 3. Results 3.1. Dasyhelea azteca Huerta & Grogan Dasyhelea azteca Huerta and Grogan 2006: 893 (male, female, Mexico); Borkent 2016: 62 (in online World catalog). 3.1.1. Diagnosis The only Neotropical species of the grisea group with the following combination of characters: adult males having the aedeagal posterolateral arms stout, with the right arm shorter than the left one, and the posterior margin of the sternite 9 with fine spicules; adult female with abdominal terga brown, hind femur with broad dark spots on midportion; scutellum with dark brown on midportion. Pupa with respiratory organ with scale-like spines and bearing 22–24 apical and 3–4 lateral pores. Larva with four pairs of sensilla campaniformia, scopae with 14–16 strong teeth, hypostoma with medial portion apparently smooth and flanked with 3 strong teeth followed by 3–4 small teeth and the epipharynx, with medial sclerite. 3.1.2. Description fourth instar larva (Figs. 1 A–G, 2 A–C) Head capsule pale brown, short, wide, tapering to apex (Fig. 1A); chaetotaxy as in Fig. 1A–E; HL 0.24–0.26 (0.25, n = 2) mm, SEM 0.252 mm; HW 0.18–0.23 (0.20, n = 2) mm, SEM 0.156 mm; HR 1.13–1.36 (1.24, n = 4), SEM 1.615; SGW 0.225 mm; SGR 1.04. Antennae cylindrical, short (Fig. 1A–D). Labrum (Figs. 1 B, D, 2 A–B) 0.80 times longer than wide; palatum (Figs. 1 B, D, 2 B) with four pairs of sensilla campaniformia (Fig. 1C–E), posterior of them two pairs of sensilla coeloconica, one simple, one serrate (Fig. 1C–E); messors well developed, stout (Fig. 1D–E); scopae well developed, with 14–16 strong teeth (Fig. 1F). Maxilla (Fig. 1B) well sclerotized; galeolacinia (Fig. 1C–F) with short, stout seta, 5–6 papillae stout and two flaplike lobe; maxillary palpus (Fig. 1C, E–F) short, buttonlike, with three small papillae; lacinial sclerite 1 stout, with short seta, lacinial sclerite 2 stout without seta (Figs. 1E–F). Mandible (Fig. 1B–F; 2A–C) strongly sclerotized, stout, with three teeth, apical tooth elongate, mesal tooth medium-sized, proximal tooth short; MDL 0.058–0.066 (0.062, n = 2) mm. Hypostoma (Fig. 1B–C, E–F) with medial portion apparently smooth, flanked by 3 strong teeth, followed by 3–4 small teeth on each side. Epipharynx massive, strongly sclerotized, lateral arms stout, short with auxiliary sclerites (Fig. 2A, C); LAW 0.10–0.12 (0.11, n = 2) mm, DCW 0.059–0.061 (0.060, n = 2) mm. Hypopharynx (Fig. 2A–C) stout, sclerotized, posterior comb with fringe. Thoracic pigmentation diffuse, pale brown. Abdominal segment whitish, with diffuse pale brown pigmentation. Caudal segment (Fig. 1G) with anterior ring of small hooks, 10–12 posterior stout hooks .
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n = 9) mm; DAW 0.19–0.21 (0.20, n = 9) mm; DAW/DAL 1.90–2.33 (2.18, n = 9). Sensilla: three dorsolateral cephalic sclerite sensilla (Figs. 2 G, 3 A): DL-1-H medium-sized, thin seta, DL-2-H short, stout setae, DL-3-H campaniform sensillum; two clypeal/labrals (Figs. 1 H, 2 H): CL-1-H medium-sized, thin seta, CL-2-H short, thin seta; one ocular (Fig. 2H): O-2-H campaniform sensillum. Cephalothorax surface with small rounded tubercles, length 0.81–1.08 (0.96, n = 10) mm, width 0.60–0.69 (0.64, n = 10) mm. Thorax. Prothoracic extension absent; respiratory organ (Figs. 1 H–I, 2 D, G) brown, nearly straight, with scale-like spines with 22–24 apical and 3–4 lateral pores; RO length 0.16–0.20 (0.17, n = 10) mm, RO width 0.02 (n = 10) mm; without pedicel. Sensilla: two anteromedial sensilla (Figs. 2 G, 3 A): AM-1-T long, thin seta, AM-2-T short, thin seta; three anterolateral sensilla (Figs. 2 G, 3 A): AL-1-T long, thin seta, AL-2-T medium-sized, thin seta, AL-3-T campaniform sensillum; dorsals (Fig. 3B): D-1-T, D-2-T short setae, D-3-T campaniform sensillum, supraalar (SA-2-T) campaniform sensillum. Metathoracics (Fig. 3C): M-2-T, M-3-T campaniform sensilla. Abdomen. abdominal segments covered with small spinules. Sensilla: tergite 1 (Fig. 3C) with setae as follows: D-2-I peg; D-4-I, D-7-I campaniform sensilla; L-1-I long, thin seta, L-2-I, L-3-I short, thin setae. Second abdominal segment similar to the first one; segment 4 with sensillar pattern (Fig. 3D) as follows: D-2-IV short seta; D1-IV, D-4-IV, D-7-IV campaniform sensilla, D-8-IV medium-sized, thin seta, all located on flattened tubercles; L-1-IV medium-sized, thin seta, L-2-IV, L-3-IV,-4-IV short, thin setae, all located on triangular tubercles; V-1-IV campaniform sensillum, V-6-IV long, thin seta, V-7-IV short, stout seta, both on flattened tubercles. Segment 9 (Fig. 2D, I) 1.52 X longer than wide, ventral surface with many spinules; length 0.12–0.14 (0.13, n = 10) mm, width 0.13–0.17 (0.15, n = 10) mm. Terminal process (Fig. 2D, I) triangular, divergent, tip pointed, with D-5-IX campaniform sensillum, V-1-IX long, thin seta, V-2-IX short, stout seta; length 0.02–0.04 (0.03, n = 10) mm. 3.1.4. Description male pupa Similar to female with usual sexual differences. Total length 2.17–2.52 (2.28, n = 6) mm. Exuviae pale brown, except cephalotorax brown. Dorsal apotome (Fig. 2F) with DAL 0.12–0.14 (0.13, n = 8) mm; DAW 0.25–0.29 (0.27, n = 8) mm, DAW/DAL 1.96–2.32 (2.07, n = 8). Respiratory organ, RO length 0.19–0.26 (0.22, n = 8) mm, RO width 0.025–0.030 (0.027, n = 8); without pedicel. Cephalotorax: length 0.97–1.10 (1.05, n = 5) mm, width 0.62–0.72 (0.65, n = 5) mm. Segment 9 (Fig. 2J) length 0.19–0.22 (0.20, n = 8) mm, width 0.17–0.23 (0.19, n = 8) mm; genital lobe extending slightly beyond of posterior margin of segment, terminal process (Fig. 2J) length 0.040–0.050 (0.045, n = 8) mm. 3.1.5. Distribution Mexico (Morelos, Jalisco); Argentina (Tucumán, Corrientes, Buenos Aires).
3.1.3. Description female pupa (Figs. 1 H–I, D–I, 2 3 A–D) Habitus as in Fig. 2D. General coloration of exuviae brown. Total length 2.13–2.40 (2.22, n = 8) mm .
3.1.6. Type material Holotype male on microscope slide, labeled: “Holotype” and “Allotype” Dasyhelea azteca: Mexico, Morelos, Municipio Tepalcingo, Huitchila, locality “Los Sauces”, 23/28-VI-1993, forest, H. Huerta, Malaise trap.
Head. Dorsal apotome (Figs. 1 H–I, 2 E) 2.18 X broader than long, surface covered with small rounded tubercles, anterior margin straight, with 2 pairs of raised, wrinkled areas; dorsal apotome sensilla (Fig. 2E): DA-1-H short, thin seta, DA-2-H campaniform sensillum; apex of posterior margin rounded with 2 stout, rounded tubercle, one dorsomedial, one apical; DAL 0.08–0.10 (0.09,
3.1.7. Material examined Argentina: Tucumán, Departamento San Isidro de Lules, Potrero de Las Tablas, arroyo Los Tablaos y RP. 341, 26.855429, 65.431505, 690 m, 14-XI-2014, Donato-Siri, 1 male (with pupal exuvium) (MLPA), 1 female (with pupal exuvium) (MLPA). Corrientes, Estación Biológica Corrientes, arroyito, 27◦ 32 51.8 S, 58◦ 40 47.4
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Fig. 1. SEM photomicrographs of Dasyhelea azteca Huerta & Grogan (A–G) larva, (H–I) female pupa. (A) Head capsule chaetotaxy, dorsal view. (B) Head capsule chaetotaxy, ventralfrontal view. (C) Head capsule, detail of lateralfrontal view. (D) Head capsule, detail of palatum and messors. (E) Head capsule, maxilla. (F) Head capsule, detail of maxilla. (G) Caudal segment. (H) Dorsal apotome and clypeal/labrals. (I) Respiratory organ. Antennae (AN); clypeal/labral sensilla (CL); dorsal apotome (DA); galeolacinia (GL); hooks (H); hooklets (Hts); hypostoma (HY); labrum (LB); lacinial sclerite 1 (LC1); lacinial sclerite 2 (LC21); maxilla (MX); maxillary palpus (MP); messors (MS); palatum (PL); pore (p); respiratory organ (RO); sensilla campaniformia (Sca); sensilla coeloconica (Sco); scopae (SC). Head capsule and caudal segment chaetotaxy are indicated by single letters: j, collar pits; o, parahypostomal setae; p, posterior perifrontal setae; q, postfrontal setae; s, anterior perifrontal setae; t, prefrontal setae; u, mesolateral setae; w, anterolateral setae; x, paranntenal setae; z, frontal pits.
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Fig. 2. Dasyhelea azteca Huerta & Grogan, (A–C) larva, (D–E, G–I) female pupa, (F, J) male pupa. (A) Head capsule, ventral view. (B) Head capsule, ventralfrontal view. (C) Hypopharynx and epipharynx. (D) Entire pupa. (E–F) Dorsal apotome. (G) Cephalothoracic sensilla (dorsal view). (H) Clypeal/labral sensilla and ocular sensillum (ventral view). (I, J) Segment 9 (ventral view). Anterolateral sensilla (AL-1-T, AL-2-T, AL-3-T); anteromedial sensilla (AM-1-T, AM-2-T); auxiliary sclerites (AS); dorsal apotome sensilla (DA-1-H, DA-2-H); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H); dorsal comb (DC); epipharynx (epy); genital lobe (GL); hypopharynx (hyp); hypostoma (HY); labrum (LB); mandibule (MD); palatum (PL); respiratory organ (RO); segment 9 sensilla (D-5-IX, V-2-IX), terminal process (TP).
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Fig. 3. Dasyhelea azteca Huerta & Grogan, (A–D) female pupa. (A) Cephalothoracic sensilla (dorsal view). (B) Dorsal sensilla. (C) Metathorax and tergite 1. (D) Segment 4. Anterolateral sensilla (AL-1-T, AL-2-T, AL-3-T); anteromedial sensilla (AM-1-T, AM-2-T); dorsal sensilla (D-1-T, D-2-T, D-3-T); dorsolateral cephalic sclerite sensilla (DL-1H, DL-2-H, DL-3-H); metathoracic sensilla (M-2-T, M-3-T); segment 4 sensilla (D-1-IV, D-2-IV, D-3-IV, D-4-IV, D-5-IV, D-7-IV, D-8-IV, L-1-IV, L-2-IV, L-3-IV, L-4-IV,V-1-IV, V-5-IV,V-6-IV, V7-IV), supraalar campaniform sensillum (SA-2-T); tergite 1 sensilla (D-2-I, D-4-I, D-7-I, D-8-I, L-1-I, L-2-I, L-3-I).
W, 55 m, 10-XI-2011, Díaz-Marino, 3 males (with pupal exuviae) (MLPA), 2 males pupae (MLPA), 1 larval exuvium (MLPA); same data except 11-XI-2011, 5 females (with pupal exuviae) (MLPA). Buenos Aires, Isla Martin García, Casa de Bombas, 16-IV-2009, DíazMarino, 4 males (with pupal exuviae) (MLPA), 7 females (with pupal
exuviae) (MLPA), 1 exuvium larva (MLPA), 1 exuvium pupa male (MLPA). 3.1.8. Material examined with SEM Argentina: Buenos Aires, Isla Martin García, Casa de Bombas, 16-IV-2009, Díaz-Marino, 1 larva, 3 pupae.
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3.1.9. Remarks The reared male adults were identified as D. azteca by the peculiar aedeagus of this species, with the posterolateral arms distinctly unequal, as well as by the posteromedial margin of the sternite 9 spiculate. 3.2. Dasyhelea yunga Díaz sp. n. 3.2.1. Diagnosis The only Neotropical species in the grisea group with the following combination of characters: males with gonostylus stout and pilose, and the aedeagal posterolateral arms stout, straight, each one arm with distal portion abruptly recurved ventrad and ending in a slender filament; females with subgenital plate arrowshaped, anterior portion broad basally, narrowed at midlength, distally broad with blunt tip, posterolateral arms stout, straight and spermatheca with neck conical, long, stout. Pupa with exuviae yellowish, with dorsal apotome smooth, without clypeal/labrals and one ocular sensillum. 3.2.2. Description female pupa (Figs. 4 B, F–J, 6 F–G) General coloration of exuviae yellowish, except anterior portion of cephalotorax pale brown. Total length 2.10 mm . Head. Dorsal apotome (Fig. 4B) 2.33 X broader than long, apex rounded, surface smooth, anterior margin straight, with 2 pairs of raised, wrinkled areas; dorsal apotome sensilla (Fig. 4B): DA-1-H medium-sized, thin seta, DA-2-H campaniform sensillum, lateral to DA-1-H; posterior margin rounded with 3 stout, rounded tubercle, two dorsomedial, other apical; DAL 0.09 mm; DAW 0.21 mm; DAW/DAL 2.33. Sensilla: three dorsolateral cephalic sclerite sensilla, DL-1-H medium-sized, thin seta, DL-2-H short, stout seta, DL-3-H campaniform sensillum; without clypeal/labrals; one ocular: O-2-H campaniform sensillum (Fig. 4G). Cephalothorax surface with small rounded tubercles, length 0.96 mm, width 0.66 mm. Thorax. prothoracic extension absent; respiratory organ (Fig. 4F) pale brown, nearly straight, smooth, with 18–20 apical and 3–4 lateral pores; RO length 0.19 mm, RO width 0.03 mm; without pedicel. Sensilla: two anteromedial sensilla, AM-1-T long, thin seta, AM-2-T short, thin seta; three anterolateral sensilla, AL-1-T long, thin seta, AL-2-T medium-sized, thin seta, AL-3-T short, stout seta; dorsals (Figs. 4H, 6F): D-1-T, short, thin seta, D-2-T minute seta, D-3-T campaniform sensillum; supraalar (SA-2-T) campaniform sensillum. Metathoracics (Figs. 4 I, 6 G): M-2-T, M-3-T campaniform sensilla, M-3-T near anterior margin of metathorax. Abdomen. abdominal segments covered with small spinules. Sensilla: tergite 1 (Figs. 4 I, 6 G) with setae as follows: D-2-I peg; D-4-I, D-7-I campaniform sensilla; L-1-I long, thin seta, L-2-I, L-3-I short, thin setae. Second abdominal segment similar to the first one; segment 4 with sensillar pattern as follows: D-2-IV short seta; D-1-IV, D-4-IV, D-7-IV campaniform sensilla, D-8-IV medium-sized, thin seta, all located on flattened tubercles; L-1-IV medium-sized, thin seta, L-2-IV, L-3-IV,-4-IV short, thin setae, all located on triangular tubercles; V-1-IV campaniform sensillum, V-6-IV long, thin seta, V-7-IV short, stout seta, both on flattened tubercles. Segment 9 (Fig. 4J) 1.50 X longer than wide, length 0.14 mm, width 0.16 mm; dorsal surface with D-5-IX campaniform sensillum, D-6-IX not visible in specimen examined; ventral surface with many spinules with V-1-IX long, thin seta, V-2-IX short, stout seta. Terminal process (Fig. 4J) triangular, divergent, tip pointed, base wide two setae, one long, thin seta on small rounded base, other seta on rounded tubercle; length 0.03 mm.
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3.2.3. Description male pupa (Fig. 4A, C–E, K; 6E, H) Similar to female with usual sexual differences. Habitus as in Fig. 4A. Total length 2.45 mm. Exuviae yellowish, except cephalotorax pale brown. Dorsal apotome (Fig. 4C) with DAL 0.10–0.11 (0.105, n = 2) mm; DAW 0.25–0.27 (0.26, n = 2) mm, DAW/DAL 2.27–2.70 (2.48, n = 2). Cephalothoracic sensilla as in Fig. 4D–E; 6E. Respiratory organ, RO length 0.22–0.23 (0.225, n = 2) mm, RO width 0.02 (n = 2). Cephalotorax: length 1.11–1.23 (1.17, n = 2) mm, width 0.78 (n = 2) mm. Segment 4 as in Fig. 6H. Segment 9 (Fig. 4K) length 0.17–18 (0.175, n = 2) mm, width 0.19–12 (0.195, n = 2) mm; genital lobe moderately wide and globose extending under posterior margin of segment. Terminal process (Fig. 4K) length 0.04 (n = 2) mm. 3.2.4. Description male adult (Figs. 5 A–D, 6 A–D) Similar to female with usual sexual differences. Flagellum collapsed in the available specimens. Clypeus with 5 pairs of setae (Fig. 5A). Palpus (Fig. 5B) with third segment bearing scattered sensilla; PR 2.30–2.62 (2.46, n = 2). Scutellum with 6–8 large, 1–2 smaller setae. Wing (Fig. 5C) length 1.22–1.25 (1.23, n = 2) mm, width 0.42 (n = 2) mm, CR 0.49–0.50 (0.49, n = 2). Genitalia (Figs. 5 D, 6 A), tergite 9 truncated distally, produced beyond level of apex of gonocoxites; apicolateral process elongate, slender, with stout apical seta; cercus small with 1–2 setae; sternite 9 (Fig. 6B) 0.38 X longer than greatest width, posteromedian margin slightly concave. Gonocoxite stout, 1.80 X longer than greatest width, with pointed anteromedian process; gonostylus 0.70 X longer than gonocoxite, stout, pilose, wide base, nearly straight, blunt tip. Paramere and gonocoxal apodemes forming an asymmetrical structure (Fig. 6C); gonocoxal apodeme stout, curved; left apodeme slightly contacting paramere, right widely contacting paramere, latter stout, tapering to pointed tip. Aedeagus (Fig. 6D) 0.72 length of greatest width; anterior margin slightly concave, basal arch extending to 0.18 of total length; basal arms slender, directed posterolaterally; posterolateral arms stout, straight, slightly convergent, each arm with distal portion abruptly recurved ventrad, ending in slender filament . 3.2.5. Description female adult (Fig. 5E–J) Head dark brown (Fig. 5E). Eyes contiguous by width of 3–4 ommatidia. Antennal flagellum (Fig. 5E–F) brown; AR 0.93. Frontal sclerite (Fig. 5G) 0.54 longer than wide, broad, elliptical, with long, slender projection. Clypeus with 5 pairs of setae. Palpus pale brown, collapsed in the available specimen. PR not measurable. Thorax. Scutum brown as in Fig. 5H, scutellum paler, with 8 large, 3 smaller setae. Legs (Fig. 5H) pale brown, femorotibial joints darkish, tarsi slightly infuscated; hind tibial comb with 6 spines; prothoracic TR 2.61, mesotoracic TR 2.48, metathoracic TR 2.52. Wing (Fig. 5I), length 1.08 mm, width 0.48 mm, CR 0.53; membrane hyaline, densely covered with macrotrichia; 2nd radial cell reduced to a suture; fork of CuA1 at level of end of 1th radial cell. Halter pale brown. Abdomen. Pale brown. Subgenital plate (Fig. 5J) arrow-shaped, anterior portion broad basally, narrowed at midlength, distally broad with blunt tip; posterolateral arms stout, straight, slightly curved. Spermatheca partially collapsed, rounded (Fig. 5J), neck conical, long, stout, measuring 22.5 . 3.2.6. Distribution Argentina (Tucumán). 3.2.7. Type material Holotype male (with pupal exuvium) on microscope slide, labeled: “Holotype” Dasyhelea yunga Díaz. Argentina: Tucumán prov., Departamento San Isidro de Lules, Potrero de Las Tablas,
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Fig. 4. Dasyhelea yunga Díaz, (B, F–J) female pupa, (A, C–E, K) male pupa. (A) Entire pupa. (B-C) Dorsal apotome. (D) Dorsolateral cephalic sclerite sensilla. (E) Cephalothoracic sensilla (dorsal view). (F) Respiratory organ. (G) Clypeal/labral sensilla and ocular sensilla. (H) Dorsal sensilla. (I) Metathorax and tergite 1. (J–K) Segment 9 (ventral view). Anterolateral sensilla (AL-1-T, AL-2-T, AL-3-T); anteromedial sensilla (AM-1-T, AM-2-T); dorsal apotome sensilla (DA-1-H, DA-2-H); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H, DL-3-H); dorsal sensilla (D-1-T, D-2-T, D-3-T); genital lobe (GL); metathoracic sensilla (M-2-T, M-3-T); ocular sensillum (O); respiratory organ (RO); segment 9 sensilla (D-5-IX, V-1-IX, V-2-IX); supraalar campaniform sensillum (SA-2-T); tergite 1 sensilla (D-2-I, D-4-I, D-7-I, D-8-I, L-1-I, L-2-I, L-3-I); terminal process (TP).
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Fig. 5. Dasyhelea yunga Díaz, (A–D) adult male, (E–J) adult female. (A) Clypeus. (B) Palpus. (C, I) Wing. (D) Gentalia. (E) Head. (F) Flagellum. (G) Frontal sclerite. (H) Thorax and legs. (J) Abdominal segment 8–9.
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Fig. 6. Dasyhelea yunga Díaz, (A–D) adult male, (E–G) female pupa, (H) male pupa. (A) Genitalia. (B) Sternite 9. (C) Paramere and gonocoxal apodemes. (D) Aedeagus. (E) Cephalothoracic sensilla (dorsal view). (F) Dorsal sensilla. (G) Metathorax and tergite 1. (H) Segment 4. Anterolateral sensilla (AL-1-T, AL-2-T, AL-3-T); anteromedial sensilla (AM-1-T, AM-2-T); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H, Dl-3-H); dorsal sensilla (D1-T, D-2-T, D-3-T); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H, DL-3-H); metathoracic sensilla (M-2-T, M-3-T); segment 4 sensilla (D-1-IV, D-2-IV, D-3-IV, D-4-IV, D-5-IV, D-7-IV, D-8-IV, L-1-IV, L-2-IV, L-3-IV, L-4-IV, V-1-IV, V-5-IV,V-6-IV, V7-IV), supraalar campaniform sensillum (SA-2-T); tergite 1 sensilla (D-2-I, D-4-I, D-7-I, D-8-I, L-1-I, L-2-I, L-3-I).
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arroyo Los Tablaos y RP. 341, 26.855429, 65.431505, 690 m, 14-XI2014, Donato-Siri. Paratype. 1 male (with pupal exuvium), 1 female (with pupal exuvium), same data as holotype. 3.2.8. Etymology The specific epithet “yunga” is a reference to the collected area. 3.3. Dasyhelea allegrae Díaz sp. n. 3.3.1. Diagnosis The only Neotropical species of the mutabilis group with the following combination of characters: males with legs entirely pale brown, apicolateral process of tergite 9 bifid at extreme tip, paramere with J-shaped distal portion, and aedeagus lacking posteromedian projection, posterolateral arms divergent, each with recurved pointed tip; females unknown. Pupa with respiratory organ paddle-shaped, with labyrinthic ducts opened by uncountable pores and basal portion dark brown, slender and annulate. 3.3.2. Description male pupa (Fig. 7A–H; 9D–G) Habitus as in Fig. 7A. General coloration of exuviae pale brown, except basal portion of respiratory organ dark brown. Total length 2.60 mm. Head. Dorsal apotome (Fig. 7B) 2.17 X broader than long, apex rounded, surface covered with small rounded tubercles, anterior margin deeply notched, with 2 pairs of raised, smooth areas; apex of posterior margin rounded, tip hyaline, with stout, dorsomedial conspicuous, elongated tubercle; DAL 0.11 mm; DAW 0.25 mm; DAW/DAL 2.17; sensilla: dorsal apotome sensilla(Fig. 7B): DA1-H medium-sized, stout seta, DA-2-H campaniform sensillum, lateral to DA-1-H. Mouthparts (Fig. 7D) with mandible, lacinia; palpus extending posterior to posterolateral margin of labium; labium divided medially by two halves slightly overlapping. Sensilla: three dorsolateral cephalic sclerite sensilla (Figs. 7 C, 9 D): DL-1-H, DL-2-H short, thin setae, DL-3-H campaniform sensillum; two clypeal/labrals (Fig. 7D): CL-1-H, CL-2-H medium-sized, thin setae; two ocular sensilla (Fig. 7D): O-1-H medium-sized, thin seta, O-2-H campaniform sensillum. Cephalothorax surface smooth or with small rounded tubercles, length 1.10 mm, width 0.67 mm. Thorax. prothoracic extension absent; respiratory organ long (Fig. 7A, E), apical portion pale brown, paddle-shaped, with labyrinthic ducts opened by uncountable pores, basal portion dark brown, slender, annulate; RO length 0.46 mm, RO width 0.10 mm; without pedicel. Sensilla: two anteromedials (Fig. 7F, 9D): AM1-T long, thin seta, AM-2-T short, stout seta; three anterolaterals (Figs. 7 F, 9 D): AL-1-T long, thin seta, AL-2-T short, stout seta, AL3-T campaniform sensillum; dorsals (Fig. 9E): D-1-T, D-2-T short, thin setae, D-3-T campaniform sensillum, supraalar (SA-2-T) campaniform sensillum. Metathoracics (Figs. 7 G, 9 F): M-2-T, M-3-T campaniform sensilla, M-3-T near anterior margin of metathorax.
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spinules with V-1-IX, V-2-IX long, thin setae, V-1-IX longer than V2-IX; length 0.21 mm, width 0.14 mm; genital lobe moderately long and globose extending under posterior margin of segment. Terminal processes (Fig. 7A, H) triangular, divergent, tip pointed length 0.11 mm. 3.3.3. Description male adult (Figs. 8 A–E, 9 A–C) Head brown (Fig. 8A). Eyes contiguous by width of 2–3 ommatidia. Antenna with flagellum (Fig. 8A) brown; AR 1.11. Clypeus (Fig. 8B) with 4 pairs of setae. Palpus (Fig. 8C) brown, collapsed in the available specimen; segments 2–5 in proportion of 20-58-2126, PR not measurable. Thorax brown; scutellum paler, with 7–8 large setae, 2–3 thinner ones. Legs entirely pale brown; apex of hind tibial with 8 spines; prothoracic tarsal ratio 1.66, mesothoracic tarsal ratio 1.77, metathoracic tarsal ratio 1.59. Wing (Fig. 8D) length 1.23 mm, width 0.40 mm, costal ratio 0.43; membrane hyaline, densely covered with macrotrichia; 2nd radial cell with lumen; fork of CuA1 at level of end of 2nd radial cell. Halter missing in the available specimen. Abdomen brown. Genitalia (Figs. 8 E, 9 A) tergite 9 slightly tapering, distal margin rounded; extending to apex of gonocoxites; apicolateral process stout, short, bifid at extreme tip, outer tooth larger than inner one, each one with thin seta; cercus not visible in the available specimen; sternite 9 not discernible due mounting process. Gonocoxite stout, 1.5x longer than greatest width with small, well sclerotized, pointed posteromedian process; gonostylus 0.9 length of gonocoxite, base moderately broad, slightly curved, tapering distally, tip pointed. Paramere and gonocoxal apodemes forming an asymmetrical structure (Fig. 9B); left apodeme stouter, broadly connected to paramere, right slender, slightly connected to paramere; latter stout, J-shaped, tip pointed. Aedeagus (Fig. 9C) as long as of greatest width; W/L ratio 0.10, basal arch stout, heavily sclerotized, concave; basal arms stout, recurved; without posteromedian projection; posterolateral arms broad basally, tapering distally, divergent, with recurved pointed tip . 3.3.4. Female Unknown. 3.3.5. Distribution Argentina (Catamarca). 3.3.6. Type material Holotype male (with pupal exuvium) on microscope slide, labeled: “Holotype” Dasyhelea allegrae Díaz. Argentina: Catamarca: Dpto Andalgalá, Río Las Quintas y RP. 341, 27,3684296, 65,9812444, 1557 m, 11-XI-2014, Donato-Siri (MLPA). 3.3.7. Etymology This species is named after Allegra Haded, goddaughter of Florentina Díaz. 4. Taxonomic discussion
Abdomen. Abdominal segments covered with small spinules. Sensilla: tergite 1(Figs. 7 G, 9 F): with setae as follows: D-2-I peg; D-4-I, D-7-I campaniform sensilla; L-1-I long, thin seta, L-2-I, L3-I short, thin setae. Second abdominal segment similar to the first one; segment 4 with sensillar pattern (Fig. 9G) as follows: D-2-IV medium-sized, stout seta on notorious triangular tubercle, D-I-IV, D-4-IV, D-7-IV campaniform sensilla, D-8-IV medium-sized, thin seta, all located on triangular tubercles; L-1-IV long, thin seta, L-2-IV, L-3-IV, L-4-IV short, stout setae, all located on triangular tubercles; V-6-IV medium-sized, thin seta, V-7-IV short, stout seta, both on flattened tubercles. Segment 9 (Fig. 7A, H) 1.50 X longer than width, dorsal surface with D-5-IX campaniform sensillum, D6-IX not visible in specimen examined; ventral surface with many
Two of the described species, Dasyhelea azteca and Dasyhelea yunga n. sp., belong to the grisea group as defined by Waugh and Wirth (1976), and this group is equivalent to the subgenus Dasyhelea s. str. (Dominiak, 2012). The larva of D. azteca is very similar to its Neotropical congeners D. eloyi Díaz & Ronderos and D. necrophila Spinelli & Rodríguez by virtue of the pale brown head capsule, whitish body and caudal segment bearing a ring of short spines anteriorly; however, D. eloyi differs by the palatum bearing 3 pairs of sensilla campaniformia and 3 pairs of sensilla coeloconica, the scopae armed with 4–6 teeth, the mandibule with 4 stout teeth, the hypostoma crenulated with 11–12 strong teeth and the epipahrynx lacking auxiliary sclerites; D. necrophila differs by the palatum with
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Fig. 7. Dasyhelea allegrae Díaz, (A–H) male pupa. (A) Entire pupa. (B) Dorsal apotome. (C) Dorsolateral cephalic sclerite sensilla. (D) Clypeal/labral sensilla and ocular sensilla. (E) Respiratory organ. (F) Cephalothoracic sensilla (dorsal view). (G) Metathorax and tergite 1. (H) Segment 9 (ventral view). Anterolateral sensilla (AL-1-T, AL-2-T, AL-3-T); anteromedial sensilla (AM-1-T, AM-2-T); dorsal apotome sensilla (DA-1-H, DA-2-H); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H, DL-3-H); genital lobe (GL); metathoracic sensilla (M-2-T, M-3-T); respiratory organ (RO); segment 9 sensilla (D-5-IX, V-1-IX, V-2-IX); tergite 1 sensilla (D-2-I, D-4-I, D-7-I, D-8-I, L-1-I, L-2-I, L-3-I); terminal process (TP).
2 pairs of sensilla campaniformia and 2 pairs of sensilla coeloconica, the mandibule with 4 stout teeth, the hypostoma crenulated bearing 3 medial and 8 lateral strong teeth. The pupa of D. azteca is very similar to D. yunga by virtue of the single ocular sensillum and the tergite of the first abdominal segment with the L-1-I represented by a long and thin seta; however, D. yunga differs by the yellowish general coloration, the surface of the dorsal apotome smooth and bearing 3 tubercles, the absence of clypeal/labral sensilla and the respiratory organ with 18–20 apical and 3–4 lateral pores. The pupae of D. eloyi and D. necrophila differs from the two species herein described by the longer and thinner clypeal labrals, the number of pores in the respiratory organ (16–18 apical and 6–8 lateral pores in the D. eloyi, 30 apical and 4 lateral in D. necrophila),
the shorter lateral sensilla of the first abdominal segment in D. eloyi, and by the anterolateral sensilla AL-3-T represented by a peg in D. necrophila. The pupa of another Neotropical species, D. correntina Ronderos & Díaz, differs from D. azteca and D. yunga by the larger size (total length 3.48 mm), the exuviae dark brown, the respiratory organ with 10–12 apical and 6–7 lateral pores and the shorter L-1-I sensillum of the first abdominal tergite. The adults of D. yunga are very similar to D. eloyi by virtue of the fork of CuA1 at level of end of 1th radial cell and the distal portion of the aeadeagal posterolateral arms abruptly recurved ventrad and ending in a slender filament. However, D. eloyi can be distinguished from D. yunga by the frontal sclerite trilobed, the clypeus with 8 setae, the halter dark brown, the posterolateral arms of the female subgenital plate curved, the neck
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Fig. 8. Dasyhelea allegrae Díaz, (A–E) adut male. (A) Head. (B) Clypeus. (C) Palpus. (D) Wing. (E) Genitalia.
of the spermatheca long, stout and oblique, the pointed gonostili and the male sternite 9 rounded distally. The third species herein described, Dasyhelea allegrae n. sp., belongs to the mutabilis species group. Its adult male is very similar to the Patagonian species D. ingrami Díaz & Spinelli by virtue of the tergite 9 rounded distally, the paramere and gonocoxal apodemes forming an asymmetrical structure and the aedeagus lacking a posteromedian projection and with the posterolateral arms single, divergent and with recurved pointed tips; however, the male of D. ingrami differs by the apicolateral process triangular, the gonostili with wide base and tapering abruptly at midlength,
its tip hook-like and the stouter paramere with blunt tip. Apart from the peculiar respiratory organ, the pupa of this new species is very similar to D. pseudolacustris Díaz & Spinelli and D. serrana Díaz & Spinelli, both recently described by Diaz et al. (2014) from Argentinean Patagonia; however, D. pseudolacustris differs by the rounded tubercle of the posterior margin of the dorsal apotome with 6–7 strong spicules, the clypeal/labrals short and hyaline, the smooth surface of the cephalotorax and the respiratory organ slender with 42–45 apical and 6–8 lateral pores; a detailed revision of the pupa of D. pseudolacustris reveals that it has two dorsal apotome sensilla and three dorsolateral sensilla instead of one and two
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Fig. 9. Dasyhelea allegrae Díaz, (A–C) adult male, (D–G) male pupa. (A) Genitalia. (B) Paramere and gonocoxal apodemes. (C) Aedeagus. (D) Cephalothoracic sensilla (dorsal view). (E) Dorsal sensilla. (F) Metathorax and tergite 1. (G) Segment 4. Anterolateral sensilla (AL-1-T, AL-2-T, AL-3-T); anteromedial sensilla (AM-1-T, AM-2-T); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H, Dl-3-H); dorsal sensilla (D1-T, D-2-T, D-3-T); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H, DL-3-H); metathoracic sensilla (M-2-T, M-3-T); segment 4 sensilla (D-1-IV, D-2-IV, D-3-IV, D-4-IV, D-5-IV, D-7-IV, D-8-IV, L-1-IV, L-2-IV, L-3-IV, L-4-IV, V-5-IV,V-6-IV, V7-IV), supraalar campaniform sensillum (SA-2-T); tergite 1 sensilla (D-2-I, D-4-I, D-7-I, D-8-I, L-1-I, L-2-I, L-3-I).
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respectively, as they were originally described (Diaz et al., 2014); the pupa of D. serrana can be distinguished by the yellowish exuviae, the dorsal surface of the dorsal apotome covered with strong spicules and bearing one dorsal apotome sensillum, the two short clypeal/labrals, the respiratory organ with base and apex smooth and with 6–7 apical and 5–6 lateral pores and the seta D-8-IV of the fourth abdominal segment short and thin. 5. Bionomics Larvae and pupae of D. azteca were collected in March 2009 and spring 2011, 2014 from small temporary ponds near of margin of Rio de La Plata River and stream in Corrientes. Larvae lasted three days to reach to pupal stage in the laboratory and the pupae that reached the adult stage took 3–4 days to emerge. The pupae of D. yunga and D. allegrae were collected in November 2015 from arroyo Los Tablaos and Río Las Quintas respectively and lasted 2–3 days to reach the adult stage. Pupae exibit the typical semi-circular slow adbominal movement typical of other ceratopogonid pupae. Acknowledgments We are grateful to Nélida Caligaris for technical assistance and Mónica Caviglia for the English proofreading. This work was supported by Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Argentina. PIP 03051 and PIP 0568. References Borkent, A., Spinelli, G.R., 2007. Neotropical Ceratopogonidae (Diptera: Insecta). In: Adis, J., Arias, J.R., Rueda-Delgado, G., Wantzen, K.M. (Eds.), Aquatic Biodiversity in Latin America (ABLA), vol. 4. Pensoft, Sofia-Moscow, pp. 198. Borkent, A., 2014. The pupae of the biting midges of the world (Diptera: ceratopogonidae), with a generic key and analysis of the phylogenetic relationships between genera. Zootaxa 3879, 1–327.
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Borkent, A., 2016. World Species of Biting Midges (Diptera: Ceratopogonidae), http://www.inhs.uiuc.edu/research/FLYTREE/CeratopogonidaeCatalog.pdf. Last update May 16 2016. 245 pp. Brown, A.D., Grau, A., Lomáscolo, T., Gasparri, N.I., 2002. Una estrategia de ˜ (Yungas) de Argentina. conservación para las selvas subtropicales de montana Ecotrópicos 15, 147–159. Brown, A.D., Pacheco, S., Lomáscolo, T., Malizia, L., 2006. Situación ambiental de los ˜ bosques Andino Yunguenos. In: Brown, A.D., Martinez Ortiz, U., Acerbi, M., Corcuera, J. (Eds.), La Situación Ambiental Argentina 2005. Fundación Vida Silvestre Argentina, Buenos Aires, pp. 53–56. Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E., Zumbado, M.A., 2009. Manual of Central American Diptera, vol. 1. NRC Research Press, Ottawa, Ontario, Canada, 714 pp. Cabrera, A.L., Willink, A., 1973. Biogeografía de América Latina Washington D.C. OEA. Monografía n◦ 13, Serie Biología., pp. 109. Díaz, F., Ronderos, M.M., Spinelli, G.R., Ferreira-Keppler, R.L., Torreias, S.R., 2013. A new species of Dasyhelea Kieffer (Diptera: Ceratopogonidae) from brazilian amazonia. Zootaxa 3686 (1), 085–093. Diaz, F., Spinelli, G.R., Donato, M., Ronderos, M.M., 2014. A phylogenetic analysis of the mutabilis species group of Dasyhelea Kieffer, and a taxonomic revision of the Patagonian species (Diptera: ceratopogonidae). J. Nat. Hist. 48 (35–36), 2117–2175. Dominiak, P., 2012. Biting midges of the genus Dasyhelea Kieffer (Diptera: Ceratopogonidae) in Poland. Pol. J. Entomol. 81, 211–304. Huerta, H., Grogan, W.L.Jr., 2006. A new species and new record of biting midges of the genus Dasyhelea Kieffer (Diptera: Cetatopogonidae) from Morelos and Jalisco, Mexico. Proceedings of the Entomological Society of Washington 108 (4), 892–898. Kieffer, J.J., 1911. Nouvelles descriptions de chironomides obtenus d’éclosion. Bulletin de la Société de Moselle (Metz) 27, 1–60. Morrone, J.J., 2001. Biogeografía de América Latina y el Caribe. Manuales y Tesis SEA 3 Zaragoza (Spain)., pp. 148. Ronderos, M.M., Spinelli, G.R., Sarmiento, P., 2000. Preparation and mounting of biting midges of the genus Culicoides Latreille (Diptera: ceratopogonidae) to be observed with a scanning electron microscope. Trans. Am. Entomol. Soc. 126 (1), 125–132. Ronderos, M.M., Díaz, F., Sarmiento, P., 2008. A new method using acid to clean and a technique for preparation of eggs of biting midges (Diptera: Ceratopogonidae) for scanning electron microscope. Trans. Am. Entomol. Soc. 134 (3+4), 471–476. Waugh, W.T., Wirth, W.W., 1976. A revision of the genus Dasyhelea Kieffer of the eastern United States north of Florida (Diptera: Ceratopogonidae). Ann. Entomol. Soc. Am. 69 (2), 219–247.
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Zoologischer Anzeiger journal homepage: www.elsevier.com/locate/jcz
Research Paper
Two new species of Dasyhelea Kieffer and the immature of D. azteca Huerta & Grogan from northwestern Argentina (Diptera: Ceratopogonidae) Florentina Díaz a,∗ , Gustavo Ricardo Spinelli b , María Marcela Ronderos a a Centro de Estudios Parasitológicos y de Vectores (CEPAVE), Facultad de Ciencias Naturales y Museo (UNLP), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Boulevard 120 S/N e/61 y 62 La Plata, Buenos Aires, Argentina b Instituto de Limnología “Dr. Raúl A. Ringuelet”(ILPLA), Facultad de Ciencias Naturales y Museo (UNLP), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Boulevard 120 S/N e/61 y 62 La Plata, Buenos Aires, Argentina
a r t i c l e
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Article history: Received 2 January 2017 Received in revised form 12 November 2017 Accepted 15 November 2017 Available online 16 November 2017 Corresponding Editor: Sven Bradler
a b s t r a c t Two new species of Dasyhelea Kieffer, D. yunga and D. allegrae, placed in the grisea and mutabilis species groups respectively, are described, illustrated and photographed based on pupae and male and female adults. In addition, the immatures of D. azteca are described for the first time. The specimens were collected in the southernmost area of the Argentinean Yungas. The pupae of D. yunga and D. allegrae were examined using a Compound Microscope (CM), while the larva and pupa of D. azteca by a Scanning Electron Microscope (SEM) and CM. © 2017 Elsevier GmbH. All rights reserved.
Keywords: Dasyhelea New species Immatures Adult Yungas Argentina
1. Introduction Dasyhelea Kieffer 1911 is a large and complex genus of Ceratopogonidae with diverse morphology and biology which occurs worldwide in a variety of small aquatic habitats (Waugh and Wirth, 1976). Taxonomically, the recognition of subgenera and/or species groups is still incipient and generally has been applied intermittently only to various regional faunas (Diaz et al., 2014). Borkent (2016) listed 77 species for the Neotropics, of which only 12 are known also from their immatures. The purpose of this paper is to describe two new species and the immatures of D. azteca Huerta & Grogan from specimens collected during entomological surveys carried out recently in the southernmost area of the Argentinean Yungas, a cloudy forest that extends on the east sloppes of the Andes and subandean chains, from Venezuela to northwestern Argentina, between 500 m and 2.500/3.500 m of altitude. It is considered by certain authors as a biogeographic province (Cabrera and Willink, 1973; Morrone,
∗ Corresponding author. E-mail address: [email protected] (F. Díaz). https://doi.org/10.1016/j.jcz.2017.11.008 0044-5231/© 2017 Elsevier GmbH. All rights reserved.
2001) or as an ecoregion by others (Brown et al., 2002, 2006), and although it is characterized by a high diversity within a great number of taxa as well by a high degree of endemisms (Cabrera and Willink, 1973; Brown et al., 2002), its ceratopogonid fauna is very poorly known. 2. Material and methods Pupae of two new species were collected in summer 2014 and D. azteca in autumn 2009 and spring 2011, 2014, using a pipette from flooded soil and with drift nets in several localities of the Yungas and in the provinces of Corrientes and Buenos Aires. Specimens were carried to the laboratory, larvae placed individually in Petri dishes and pupae isolated in a vial with a drop of water and observed daily until adult emergence. Adults were allowed to harden for 24 h before being preserved to ensure their pigmentation was complete. Larval, pupal exuviae and adults were mounted in Canada balsam following the technique described by Borkent and Spinelli (2007). For Scanning Electron Microscope (SEM), larva and pupae of D. azteca were prepared following the technique of Ronderos et al. (2000, 2008). Ink illustrations were made using an attached camera lucida. Photographs were taken with a digital camera Micrometrics
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SE Premium, through a Nikon Eclipse E200 microscope. Measurements were taken using SEM and Compound Microscope (CM). For terminology of larva see Díaz et al. (2013), Borkent (2014) for pupa and Brown et al. (2009) for adult. The studied specimens are deposited in the collection of the División Entomología, Museo de La Plata, Argentina (MLPA). 3. Results 3.1. Dasyhelea azteca Huerta & Grogan Dasyhelea azteca Huerta and Grogan 2006: 893 (male, female, Mexico); Borkent 2016: 62 (in online World catalog). 3.1.1. Diagnosis The only Neotropical species of the grisea group with the following combination of characters: adult males having the aedeagal posterolateral arms stout, with the right arm shorter than the left one, and the posterior margin of the sternite 9 with fine spicules; adult female with abdominal terga brown, hind femur with broad dark spots on midportion; scutellum with dark brown on midportion. Pupa with respiratory organ with scale-like spines and bearing 22–24 apical and 3–4 lateral pores. Larva with four pairs of sensilla campaniformia, scopae with 14–16 strong teeth, hypostoma with medial portion apparently smooth and flanked with 3 strong teeth followed by 3–4 small teeth and the epipharynx, with medial sclerite. 3.1.2. Description fourth instar larva (Figs. 1 A–G, 2 A–C) Head capsule pale brown, short, wide, tapering to apex (Fig. 1A); chaetotaxy as in Fig. 1A–E; HL 0.24–0.26 (0.25, n = 2) mm, SEM 0.252 mm; HW 0.18–0.23 (0.20, n = 2) mm, SEM 0.156 mm; HR 1.13–1.36 (1.24, n = 4), SEM 1.615; SGW 0.225 mm; SGR 1.04. Antennae cylindrical, short (Fig. 1A–D). Labrum (Figs. 1 B, D, 2 A–B) 0.80 times longer than wide; palatum (Figs. 1 B, D, 2 B) with four pairs of sensilla campaniformia (Fig. 1C–E), posterior of them two pairs of sensilla coeloconica, one simple, one serrate (Fig. 1C–E); messors well developed, stout (Fig. 1D–E); scopae well developed, with 14–16 strong teeth (Fig. 1F). Maxilla (Fig. 1B) well sclerotized; galeolacinia (Fig. 1C–F) with short, stout seta, 5–6 papillae stout and two flaplike lobe; maxillary palpus (Fig. 1C, E–F) short, buttonlike, with three small papillae; lacinial sclerite 1 stout, with short seta, lacinial sclerite 2 stout without seta (Figs. 1E–F). Mandible (Fig. 1B–F; 2A–C) strongly sclerotized, stout, with three teeth, apical tooth elongate, mesal tooth medium-sized, proximal tooth short; MDL 0.058–0.066 (0.062, n = 2) mm. Hypostoma (Fig. 1B–C, E–F) with medial portion apparently smooth, flanked by 3 strong teeth, followed by 3–4 small teeth on each side. Epipharynx massive, strongly sclerotized, lateral arms stout, short with auxiliary sclerites (Fig. 2A, C); LAW 0.10–0.12 (0.11, n = 2) mm, DCW 0.059–0.061 (0.060, n = 2) mm. Hypopharynx (Fig. 2A–C) stout, sclerotized, posterior comb with fringe. Thoracic pigmentation diffuse, pale brown. Abdominal segment whitish, with diffuse pale brown pigmentation. Caudal segment (Fig. 1G) with anterior ring of small hooks, 10–12 posterior stout hooks .
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n = 9) mm; DAW 0.19–0.21 (0.20, n = 9) mm; DAW/DAL 1.90–2.33 (2.18, n = 9). Sensilla: three dorsolateral cephalic sclerite sensilla (Figs. 2 G, 3 A): DL-1-H medium-sized, thin seta, DL-2-H short, stout setae, DL-3-H campaniform sensillum; two clypeal/labrals (Figs. 1 H, 2 H): CL-1-H medium-sized, thin seta, CL-2-H short, thin seta; one ocular (Fig. 2H): O-2-H campaniform sensillum. Cephalothorax surface with small rounded tubercles, length 0.81–1.08 (0.96, n = 10) mm, width 0.60–0.69 (0.64, n = 10) mm. Thorax. Prothoracic extension absent; respiratory organ (Figs. 1 H–I, 2 D, G) brown, nearly straight, with scale-like spines with 22–24 apical and 3–4 lateral pores; RO length 0.16–0.20 (0.17, n = 10) mm, RO width 0.02 (n = 10) mm; without pedicel. Sensilla: two anteromedial sensilla (Figs. 2 G, 3 A): AM-1-T long, thin seta, AM-2-T short, thin seta; three anterolateral sensilla (Figs. 2 G, 3 A): AL-1-T long, thin seta, AL-2-T medium-sized, thin seta, AL-3-T campaniform sensillum; dorsals (Fig. 3B): D-1-T, D-2-T short setae, D-3-T campaniform sensillum, supraalar (SA-2-T) campaniform sensillum. Metathoracics (Fig. 3C): M-2-T, M-3-T campaniform sensilla. Abdomen. abdominal segments covered with small spinules. Sensilla: tergite 1 (Fig. 3C) with setae as follows: D-2-I peg; D-4-I, D-7-I campaniform sensilla; L-1-I long, thin seta, L-2-I, L-3-I short, thin setae. Second abdominal segment similar to the first one; segment 4 with sensillar pattern (Fig. 3D) as follows: D-2-IV short seta; D1-IV, D-4-IV, D-7-IV campaniform sensilla, D-8-IV medium-sized, thin seta, all located on flattened tubercles; L-1-IV medium-sized, thin seta, L-2-IV, L-3-IV,-4-IV short, thin setae, all located on triangular tubercles; V-1-IV campaniform sensillum, V-6-IV long, thin seta, V-7-IV short, stout seta, both on flattened tubercles. Segment 9 (Fig. 2D, I) 1.52 X longer than wide, ventral surface with many spinules; length 0.12–0.14 (0.13, n = 10) mm, width 0.13–0.17 (0.15, n = 10) mm. Terminal process (Fig. 2D, I) triangular, divergent, tip pointed, with D-5-IX campaniform sensillum, V-1-IX long, thin seta, V-2-IX short, stout seta; length 0.02–0.04 (0.03, n = 10) mm. 3.1.4. Description male pupa Similar to female with usual sexual differences. Total length 2.17–2.52 (2.28, n = 6) mm. Exuviae pale brown, except cephalotorax brown. Dorsal apotome (Fig. 2F) with DAL 0.12–0.14 (0.13, n = 8) mm; DAW 0.25–0.29 (0.27, n = 8) mm, DAW/DAL 1.96–2.32 (2.07, n = 8). Respiratory organ, RO length 0.19–0.26 (0.22, n = 8) mm, RO width 0.025–0.030 (0.027, n = 8); without pedicel. Cephalotorax: length 0.97–1.10 (1.05, n = 5) mm, width 0.62–0.72 (0.65, n = 5) mm. Segment 9 (Fig. 2J) length 0.19–0.22 (0.20, n = 8) mm, width 0.17–0.23 (0.19, n = 8) mm; genital lobe extending slightly beyond of posterior margin of segment, terminal process (Fig. 2J) length 0.040–0.050 (0.045, n = 8) mm. 3.1.5. Distribution Mexico (Morelos, Jalisco); Argentina (Tucumán, Corrientes, Buenos Aires).
3.1.3. Description female pupa (Figs. 1 H–I, D–I, 2 3 A–D) Habitus as in Fig. 2D. General coloration of exuviae brown. Total length 2.13–2.40 (2.22, n = 8) mm .
3.1.6. Type material Holotype male on microscope slide, labeled: “Holotype” and “Allotype” Dasyhelea azteca: Mexico, Morelos, Municipio Tepalcingo, Huitchila, locality “Los Sauces”, 23/28-VI-1993, forest, H. Huerta, Malaise trap.
Head. Dorsal apotome (Figs. 1 H–I, 2 E) 2.18 X broader than long, surface covered with small rounded tubercles, anterior margin straight, with 2 pairs of raised, wrinkled areas; dorsal apotome sensilla (Fig. 2E): DA-1-H short, thin seta, DA-2-H campaniform sensillum; apex of posterior margin rounded with 2 stout, rounded tubercle, one dorsomedial, one apical; DAL 0.08–0.10 (0.09,
3.1.7. Material examined Argentina: Tucumán, Departamento San Isidro de Lules, Potrero de Las Tablas, arroyo Los Tablaos y RP. 341, 26.855429, 65.431505, 690 m, 14-XI-2014, Donato-Siri, 1 male (with pupal exuvium) (MLPA), 1 female (with pupal exuvium) (MLPA). Corrientes, Estación Biológica Corrientes, arroyito, 27◦ 32 51.8 S, 58◦ 40 47.4
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Fig. 1. SEM photomicrographs of Dasyhelea azteca Huerta & Grogan (A–G) larva, (H–I) female pupa. (A) Head capsule chaetotaxy, dorsal view. (B) Head capsule chaetotaxy, ventralfrontal view. (C) Head capsule, detail of lateralfrontal view. (D) Head capsule, detail of palatum and messors. (E) Head capsule, maxilla. (F) Head capsule, detail of maxilla. (G) Caudal segment. (H) Dorsal apotome and clypeal/labrals. (I) Respiratory organ. Antennae (AN); clypeal/labral sensilla (CL); dorsal apotome (DA); galeolacinia (GL); hooks (H); hooklets (Hts); hypostoma (HY); labrum (LB); lacinial sclerite 1 (LC1); lacinial sclerite 2 (LC21); maxilla (MX); maxillary palpus (MP); messors (MS); palatum (PL); pore (p); respiratory organ (RO); sensilla campaniformia (Sca); sensilla coeloconica (Sco); scopae (SC). Head capsule and caudal segment chaetotaxy are indicated by single letters: j, collar pits; o, parahypostomal setae; p, posterior perifrontal setae; q, postfrontal setae; s, anterior perifrontal setae; t, prefrontal setae; u, mesolateral setae; w, anterolateral setae; x, paranntenal setae; z, frontal pits.
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Fig. 2. Dasyhelea azteca Huerta & Grogan, (A–C) larva, (D–E, G–I) female pupa, (F, J) male pupa. (A) Head capsule, ventral view. (B) Head capsule, ventralfrontal view. (C) Hypopharynx and epipharynx. (D) Entire pupa. (E–F) Dorsal apotome. (G) Cephalothoracic sensilla (dorsal view). (H) Clypeal/labral sensilla and ocular sensillum (ventral view). (I, J) Segment 9 (ventral view). Anterolateral sensilla (AL-1-T, AL-2-T, AL-3-T); anteromedial sensilla (AM-1-T, AM-2-T); auxiliary sclerites (AS); dorsal apotome sensilla (DA-1-H, DA-2-H); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H); dorsal comb (DC); epipharynx (epy); genital lobe (GL); hypopharynx (hyp); hypostoma (HY); labrum (LB); mandibule (MD); palatum (PL); respiratory organ (RO); segment 9 sensilla (D-5-IX, V-2-IX), terminal process (TP).
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Fig. 3. Dasyhelea azteca Huerta & Grogan, (A–D) female pupa. (A) Cephalothoracic sensilla (dorsal view). (B) Dorsal sensilla. (C) Metathorax and tergite 1. (D) Segment 4. Anterolateral sensilla (AL-1-T, AL-2-T, AL-3-T); anteromedial sensilla (AM-1-T, AM-2-T); dorsal sensilla (D-1-T, D-2-T, D-3-T); dorsolateral cephalic sclerite sensilla (DL-1H, DL-2-H, DL-3-H); metathoracic sensilla (M-2-T, M-3-T); segment 4 sensilla (D-1-IV, D-2-IV, D-3-IV, D-4-IV, D-5-IV, D-7-IV, D-8-IV, L-1-IV, L-2-IV, L-3-IV, L-4-IV,V-1-IV, V-5-IV,V-6-IV, V7-IV), supraalar campaniform sensillum (SA-2-T); tergite 1 sensilla (D-2-I, D-4-I, D-7-I, D-8-I, L-1-I, L-2-I, L-3-I).
W, 55 m, 10-XI-2011, Díaz-Marino, 3 males (with pupal exuviae) (MLPA), 2 males pupae (MLPA), 1 larval exuvium (MLPA); same data except 11-XI-2011, 5 females (with pupal exuviae) (MLPA). Buenos Aires, Isla Martin García, Casa de Bombas, 16-IV-2009, DíazMarino, 4 males (with pupal exuviae) (MLPA), 7 females (with pupal
exuviae) (MLPA), 1 exuvium larva (MLPA), 1 exuvium pupa male (MLPA). 3.1.8. Material examined with SEM Argentina: Buenos Aires, Isla Martin García, Casa de Bombas, 16-IV-2009, Díaz-Marino, 1 larva, 3 pupae.
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3.1.9. Remarks The reared male adults were identified as D. azteca by the peculiar aedeagus of this species, with the posterolateral arms distinctly unequal, as well as by the posteromedial margin of the sternite 9 spiculate. 3.2. Dasyhelea yunga Díaz sp. n. 3.2.1. Diagnosis The only Neotropical species in the grisea group with the following combination of characters: males with gonostylus stout and pilose, and the aedeagal posterolateral arms stout, straight, each one arm with distal portion abruptly recurved ventrad and ending in a slender filament; females with subgenital plate arrowshaped, anterior portion broad basally, narrowed at midlength, distally broad with blunt tip, posterolateral arms stout, straight and spermatheca with neck conical, long, stout. Pupa with exuviae yellowish, with dorsal apotome smooth, without clypeal/labrals and one ocular sensillum. 3.2.2. Description female pupa (Figs. 4 B, F–J, 6 F–G) General coloration of exuviae yellowish, except anterior portion of cephalotorax pale brown. Total length 2.10 mm . Head. Dorsal apotome (Fig. 4B) 2.33 X broader than long, apex rounded, surface smooth, anterior margin straight, with 2 pairs of raised, wrinkled areas; dorsal apotome sensilla (Fig. 4B): DA-1-H medium-sized, thin seta, DA-2-H campaniform sensillum, lateral to DA-1-H; posterior margin rounded with 3 stout, rounded tubercle, two dorsomedial, other apical; DAL 0.09 mm; DAW 0.21 mm; DAW/DAL 2.33. Sensilla: three dorsolateral cephalic sclerite sensilla, DL-1-H medium-sized, thin seta, DL-2-H short, stout seta, DL-3-H campaniform sensillum; without clypeal/labrals; one ocular: O-2-H campaniform sensillum (Fig. 4G). Cephalothorax surface with small rounded tubercles, length 0.96 mm, width 0.66 mm. Thorax. prothoracic extension absent; respiratory organ (Fig. 4F) pale brown, nearly straight, smooth, with 18–20 apical and 3–4 lateral pores; RO length 0.19 mm, RO width 0.03 mm; without pedicel. Sensilla: two anteromedial sensilla, AM-1-T long, thin seta, AM-2-T short, thin seta; three anterolateral sensilla, AL-1-T long, thin seta, AL-2-T medium-sized, thin seta, AL-3-T short, stout seta; dorsals (Figs. 4H, 6F): D-1-T, short, thin seta, D-2-T minute seta, D-3-T campaniform sensillum; supraalar (SA-2-T) campaniform sensillum. Metathoracics (Figs. 4 I, 6 G): M-2-T, M-3-T campaniform sensilla, M-3-T near anterior margin of metathorax. Abdomen. abdominal segments covered with small spinules. Sensilla: tergite 1 (Figs. 4 I, 6 G) with setae as follows: D-2-I peg; D-4-I, D-7-I campaniform sensilla; L-1-I long, thin seta, L-2-I, L-3-I short, thin setae. Second abdominal segment similar to the first one; segment 4 with sensillar pattern as follows: D-2-IV short seta; D-1-IV, D-4-IV, D-7-IV campaniform sensilla, D-8-IV medium-sized, thin seta, all located on flattened tubercles; L-1-IV medium-sized, thin seta, L-2-IV, L-3-IV,-4-IV short, thin setae, all located on triangular tubercles; V-1-IV campaniform sensillum, V-6-IV long, thin seta, V-7-IV short, stout seta, both on flattened tubercles. Segment 9 (Fig. 4J) 1.50 X longer than wide, length 0.14 mm, width 0.16 mm; dorsal surface with D-5-IX campaniform sensillum, D-6-IX not visible in specimen examined; ventral surface with many spinules with V-1-IX long, thin seta, V-2-IX short, stout seta. Terminal process (Fig. 4J) triangular, divergent, tip pointed, base wide two setae, one long, thin seta on small rounded base, other seta on rounded tubercle; length 0.03 mm.
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3.2.3. Description male pupa (Fig. 4A, C–E, K; 6E, H) Similar to female with usual sexual differences. Habitus as in Fig. 4A. Total length 2.45 mm. Exuviae yellowish, except cephalotorax pale brown. Dorsal apotome (Fig. 4C) with DAL 0.10–0.11 (0.105, n = 2) mm; DAW 0.25–0.27 (0.26, n = 2) mm, DAW/DAL 2.27–2.70 (2.48, n = 2). Cephalothoracic sensilla as in Fig. 4D–E; 6E. Respiratory organ, RO length 0.22–0.23 (0.225, n = 2) mm, RO width 0.02 (n = 2). Cephalotorax: length 1.11–1.23 (1.17, n = 2) mm, width 0.78 (n = 2) mm. Segment 4 as in Fig. 6H. Segment 9 (Fig. 4K) length 0.17–18 (0.175, n = 2) mm, width 0.19–12 (0.195, n = 2) mm; genital lobe moderately wide and globose extending under posterior margin of segment. Terminal process (Fig. 4K) length 0.04 (n = 2) mm. 3.2.4. Description male adult (Figs. 5 A–D, 6 A–D) Similar to female with usual sexual differences. Flagellum collapsed in the available specimens. Clypeus with 5 pairs of setae (Fig. 5A). Palpus (Fig. 5B) with third segment bearing scattered sensilla; PR 2.30–2.62 (2.46, n = 2). Scutellum with 6–8 large, 1–2 smaller setae. Wing (Fig. 5C) length 1.22–1.25 (1.23, n = 2) mm, width 0.42 (n = 2) mm, CR 0.49–0.50 (0.49, n = 2). Genitalia (Figs. 5 D, 6 A), tergite 9 truncated distally, produced beyond level of apex of gonocoxites; apicolateral process elongate, slender, with stout apical seta; cercus small with 1–2 setae; sternite 9 (Fig. 6B) 0.38 X longer than greatest width, posteromedian margin slightly concave. Gonocoxite stout, 1.80 X longer than greatest width, with pointed anteromedian process; gonostylus 0.70 X longer than gonocoxite, stout, pilose, wide base, nearly straight, blunt tip. Paramere and gonocoxal apodemes forming an asymmetrical structure (Fig. 6C); gonocoxal apodeme stout, curved; left apodeme slightly contacting paramere, right widely contacting paramere, latter stout, tapering to pointed tip. Aedeagus (Fig. 6D) 0.72 length of greatest width; anterior margin slightly concave, basal arch extending to 0.18 of total length; basal arms slender, directed posterolaterally; posterolateral arms stout, straight, slightly convergent, each arm with distal portion abruptly recurved ventrad, ending in slender filament . 3.2.5. Description female adult (Fig. 5E–J) Head dark brown (Fig. 5E). Eyes contiguous by width of 3–4 ommatidia. Antennal flagellum (Fig. 5E–F) brown; AR 0.93. Frontal sclerite (Fig. 5G) 0.54 longer than wide, broad, elliptical, with long, slender projection. Clypeus with 5 pairs of setae. Palpus pale brown, collapsed in the available specimen. PR not measurable. Thorax. Scutum brown as in Fig. 5H, scutellum paler, with 8 large, 3 smaller setae. Legs (Fig. 5H) pale brown, femorotibial joints darkish, tarsi slightly infuscated; hind tibial comb with 6 spines; prothoracic TR 2.61, mesotoracic TR 2.48, metathoracic TR 2.52. Wing (Fig. 5I), length 1.08 mm, width 0.48 mm, CR 0.53; membrane hyaline, densely covered with macrotrichia; 2nd radial cell reduced to a suture; fork of CuA1 at level of end of 1th radial cell. Halter pale brown. Abdomen. Pale brown. Subgenital plate (Fig. 5J) arrow-shaped, anterior portion broad basally, narrowed at midlength, distally broad with blunt tip; posterolateral arms stout, straight, slightly curved. Spermatheca partially collapsed, rounded (Fig. 5J), neck conical, long, stout, measuring 22.5 . 3.2.6. Distribution Argentina (Tucumán). 3.2.7. Type material Holotype male (with pupal exuvium) on microscope slide, labeled: “Holotype” Dasyhelea yunga Díaz. Argentina: Tucumán prov., Departamento San Isidro de Lules, Potrero de Las Tablas,
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Fig. 4. Dasyhelea yunga Díaz, (B, F–J) female pupa, (A, C–E, K) male pupa. (A) Entire pupa. (B-C) Dorsal apotome. (D) Dorsolateral cephalic sclerite sensilla. (E) Cephalothoracic sensilla (dorsal view). (F) Respiratory organ. (G) Clypeal/labral sensilla and ocular sensilla. (H) Dorsal sensilla. (I) Metathorax and tergite 1. (J–K) Segment 9 (ventral view). Anterolateral sensilla (AL-1-T, AL-2-T, AL-3-T); anteromedial sensilla (AM-1-T, AM-2-T); dorsal apotome sensilla (DA-1-H, DA-2-H); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H, DL-3-H); dorsal sensilla (D-1-T, D-2-T, D-3-T); genital lobe (GL); metathoracic sensilla (M-2-T, M-3-T); ocular sensillum (O); respiratory organ (RO); segment 9 sensilla (D-5-IX, V-1-IX, V-2-IX); supraalar campaniform sensillum (SA-2-T); tergite 1 sensilla (D-2-I, D-4-I, D-7-I, D-8-I, L-1-I, L-2-I, L-3-I); terminal process (TP).
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Fig. 5. Dasyhelea yunga Díaz, (A–D) adult male, (E–J) adult female. (A) Clypeus. (B) Palpus. (C, I) Wing. (D) Gentalia. (E) Head. (F) Flagellum. (G) Frontal sclerite. (H) Thorax and legs. (J) Abdominal segment 8–9.
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Fig. 6. Dasyhelea yunga Díaz, (A–D) adult male, (E–G) female pupa, (H) male pupa. (A) Genitalia. (B) Sternite 9. (C) Paramere and gonocoxal apodemes. (D) Aedeagus. (E) Cephalothoracic sensilla (dorsal view). (F) Dorsal sensilla. (G) Metathorax and tergite 1. (H) Segment 4. Anterolateral sensilla (AL-1-T, AL-2-T, AL-3-T); anteromedial sensilla (AM-1-T, AM-2-T); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H, Dl-3-H); dorsal sensilla (D1-T, D-2-T, D-3-T); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H, DL-3-H); metathoracic sensilla (M-2-T, M-3-T); segment 4 sensilla (D-1-IV, D-2-IV, D-3-IV, D-4-IV, D-5-IV, D-7-IV, D-8-IV, L-1-IV, L-2-IV, L-3-IV, L-4-IV, V-1-IV, V-5-IV,V-6-IV, V7-IV), supraalar campaniform sensillum (SA-2-T); tergite 1 sensilla (D-2-I, D-4-I, D-7-I, D-8-I, L-1-I, L-2-I, L-3-I).
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arroyo Los Tablaos y RP. 341, 26.855429, 65.431505, 690 m, 14-XI2014, Donato-Siri. Paratype. 1 male (with pupal exuvium), 1 female (with pupal exuvium), same data as holotype. 3.2.8. Etymology The specific epithet “yunga” is a reference to the collected area. 3.3. Dasyhelea allegrae Díaz sp. n. 3.3.1. Diagnosis The only Neotropical species of the mutabilis group with the following combination of characters: males with legs entirely pale brown, apicolateral process of tergite 9 bifid at extreme tip, paramere with J-shaped distal portion, and aedeagus lacking posteromedian projection, posterolateral arms divergent, each with recurved pointed tip; females unknown. Pupa with respiratory organ paddle-shaped, with labyrinthic ducts opened by uncountable pores and basal portion dark brown, slender and annulate. 3.3.2. Description male pupa (Fig. 7A–H; 9D–G) Habitus as in Fig. 7A. General coloration of exuviae pale brown, except basal portion of respiratory organ dark brown. Total length 2.60 mm. Head. Dorsal apotome (Fig. 7B) 2.17 X broader than long, apex rounded, surface covered with small rounded tubercles, anterior margin deeply notched, with 2 pairs of raised, smooth areas; apex of posterior margin rounded, tip hyaline, with stout, dorsomedial conspicuous, elongated tubercle; DAL 0.11 mm; DAW 0.25 mm; DAW/DAL 2.17; sensilla: dorsal apotome sensilla(Fig. 7B): DA1-H medium-sized, stout seta, DA-2-H campaniform sensillum, lateral to DA-1-H. Mouthparts (Fig. 7D) with mandible, lacinia; palpus extending posterior to posterolateral margin of labium; labium divided medially by two halves slightly overlapping. Sensilla: three dorsolateral cephalic sclerite sensilla (Figs. 7 C, 9 D): DL-1-H, DL-2-H short, thin setae, DL-3-H campaniform sensillum; two clypeal/labrals (Fig. 7D): CL-1-H, CL-2-H medium-sized, thin setae; two ocular sensilla (Fig. 7D): O-1-H medium-sized, thin seta, O-2-H campaniform sensillum. Cephalothorax surface smooth or with small rounded tubercles, length 1.10 mm, width 0.67 mm. Thorax. prothoracic extension absent; respiratory organ long (Fig. 7A, E), apical portion pale brown, paddle-shaped, with labyrinthic ducts opened by uncountable pores, basal portion dark brown, slender, annulate; RO length 0.46 mm, RO width 0.10 mm; without pedicel. Sensilla: two anteromedials (Fig. 7F, 9D): AM1-T long, thin seta, AM-2-T short, stout seta; three anterolaterals (Figs. 7 F, 9 D): AL-1-T long, thin seta, AL-2-T short, stout seta, AL3-T campaniform sensillum; dorsals (Fig. 9E): D-1-T, D-2-T short, thin setae, D-3-T campaniform sensillum, supraalar (SA-2-T) campaniform sensillum. Metathoracics (Figs. 7 G, 9 F): M-2-T, M-3-T campaniform sensilla, M-3-T near anterior margin of metathorax.
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spinules with V-1-IX, V-2-IX long, thin setae, V-1-IX longer than V2-IX; length 0.21 mm, width 0.14 mm; genital lobe moderately long and globose extending under posterior margin of segment. Terminal processes (Fig. 7A, H) triangular, divergent, tip pointed length 0.11 mm. 3.3.3. Description male adult (Figs. 8 A–E, 9 A–C) Head brown (Fig. 8A). Eyes contiguous by width of 2–3 ommatidia. Antenna with flagellum (Fig. 8A) brown; AR 1.11. Clypeus (Fig. 8B) with 4 pairs of setae. Palpus (Fig. 8C) brown, collapsed in the available specimen; segments 2–5 in proportion of 20-58-2126, PR not measurable. Thorax brown; scutellum paler, with 7–8 large setae, 2–3 thinner ones. Legs entirely pale brown; apex of hind tibial with 8 spines; prothoracic tarsal ratio 1.66, mesothoracic tarsal ratio 1.77, metathoracic tarsal ratio 1.59. Wing (Fig. 8D) length 1.23 mm, width 0.40 mm, costal ratio 0.43; membrane hyaline, densely covered with macrotrichia; 2nd radial cell with lumen; fork of CuA1 at level of end of 2nd radial cell. Halter missing in the available specimen. Abdomen brown. Genitalia (Figs. 8 E, 9 A) tergite 9 slightly tapering, distal margin rounded; extending to apex of gonocoxites; apicolateral process stout, short, bifid at extreme tip, outer tooth larger than inner one, each one with thin seta; cercus not visible in the available specimen; sternite 9 not discernible due mounting process. Gonocoxite stout, 1.5x longer than greatest width with small, well sclerotized, pointed posteromedian process; gonostylus 0.9 length of gonocoxite, base moderately broad, slightly curved, tapering distally, tip pointed. Paramere and gonocoxal apodemes forming an asymmetrical structure (Fig. 9B); left apodeme stouter, broadly connected to paramere, right slender, slightly connected to paramere; latter stout, J-shaped, tip pointed. Aedeagus (Fig. 9C) as long as of greatest width; W/L ratio 0.10, basal arch stout, heavily sclerotized, concave; basal arms stout, recurved; without posteromedian projection; posterolateral arms broad basally, tapering distally, divergent, with recurved pointed tip . 3.3.4. Female Unknown. 3.3.5. Distribution Argentina (Catamarca). 3.3.6. Type material Holotype male (with pupal exuvium) on microscope slide, labeled: “Holotype” Dasyhelea allegrae Díaz. Argentina: Catamarca: Dpto Andalgalá, Río Las Quintas y RP. 341, 27,3684296, 65,9812444, 1557 m, 11-XI-2014, Donato-Siri (MLPA). 3.3.7. Etymology This species is named after Allegra Haded, goddaughter of Florentina Díaz. 4. Taxonomic discussion
Abdomen. Abdominal segments covered with small spinules. Sensilla: tergite 1(Figs. 7 G, 9 F): with setae as follows: D-2-I peg; D-4-I, D-7-I campaniform sensilla; L-1-I long, thin seta, L-2-I, L3-I short, thin setae. Second abdominal segment similar to the first one; segment 4 with sensillar pattern (Fig. 9G) as follows: D-2-IV medium-sized, stout seta on notorious triangular tubercle, D-I-IV, D-4-IV, D-7-IV campaniform sensilla, D-8-IV medium-sized, thin seta, all located on triangular tubercles; L-1-IV long, thin seta, L-2-IV, L-3-IV, L-4-IV short, stout setae, all located on triangular tubercles; V-6-IV medium-sized, thin seta, V-7-IV short, stout seta, both on flattened tubercles. Segment 9 (Fig. 7A, H) 1.50 X longer than width, dorsal surface with D-5-IX campaniform sensillum, D6-IX not visible in specimen examined; ventral surface with many
Two of the described species, Dasyhelea azteca and Dasyhelea yunga n. sp., belong to the grisea group as defined by Waugh and Wirth (1976), and this group is equivalent to the subgenus Dasyhelea s. str. (Dominiak, 2012). The larva of D. azteca is very similar to its Neotropical congeners D. eloyi Díaz & Ronderos and D. necrophila Spinelli & Rodríguez by virtue of the pale brown head capsule, whitish body and caudal segment bearing a ring of short spines anteriorly; however, D. eloyi differs by the palatum bearing 3 pairs of sensilla campaniformia and 3 pairs of sensilla coeloconica, the scopae armed with 4–6 teeth, the mandibule with 4 stout teeth, the hypostoma crenulated with 11–12 strong teeth and the epipahrynx lacking auxiliary sclerites; D. necrophila differs by the palatum with
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Fig. 7. Dasyhelea allegrae Díaz, (A–H) male pupa. (A) Entire pupa. (B) Dorsal apotome. (C) Dorsolateral cephalic sclerite sensilla. (D) Clypeal/labral sensilla and ocular sensilla. (E) Respiratory organ. (F) Cephalothoracic sensilla (dorsal view). (G) Metathorax and tergite 1. (H) Segment 9 (ventral view). Anterolateral sensilla (AL-1-T, AL-2-T, AL-3-T); anteromedial sensilla (AM-1-T, AM-2-T); dorsal apotome sensilla (DA-1-H, DA-2-H); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H, DL-3-H); genital lobe (GL); metathoracic sensilla (M-2-T, M-3-T); respiratory organ (RO); segment 9 sensilla (D-5-IX, V-1-IX, V-2-IX); tergite 1 sensilla (D-2-I, D-4-I, D-7-I, D-8-I, L-1-I, L-2-I, L-3-I); terminal process (TP).
2 pairs of sensilla campaniformia and 2 pairs of sensilla coeloconica, the mandibule with 4 stout teeth, the hypostoma crenulated bearing 3 medial and 8 lateral strong teeth. The pupa of D. azteca is very similar to D. yunga by virtue of the single ocular sensillum and the tergite of the first abdominal segment with the L-1-I represented by a long and thin seta; however, D. yunga differs by the yellowish general coloration, the surface of the dorsal apotome smooth and bearing 3 tubercles, the absence of clypeal/labral sensilla and the respiratory organ with 18–20 apical and 3–4 lateral pores. The pupae of D. eloyi and D. necrophila differs from the two species herein described by the longer and thinner clypeal labrals, the number of pores in the respiratory organ (16–18 apical and 6–8 lateral pores in the D. eloyi, 30 apical and 4 lateral in D. necrophila),
the shorter lateral sensilla of the first abdominal segment in D. eloyi, and by the anterolateral sensilla AL-3-T represented by a peg in D. necrophila. The pupa of another Neotropical species, D. correntina Ronderos & Díaz, differs from D. azteca and D. yunga by the larger size (total length 3.48 mm), the exuviae dark brown, the respiratory organ with 10–12 apical and 6–7 lateral pores and the shorter L-1-I sensillum of the first abdominal tergite. The adults of D. yunga are very similar to D. eloyi by virtue of the fork of CuA1 at level of end of 1th radial cell and the distal portion of the aeadeagal posterolateral arms abruptly recurved ventrad and ending in a slender filament. However, D. eloyi can be distinguished from D. yunga by the frontal sclerite trilobed, the clypeus with 8 setae, the halter dark brown, the posterolateral arms of the female subgenital plate curved, the neck
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Fig. 8. Dasyhelea allegrae Díaz, (A–E) adut male. (A) Head. (B) Clypeus. (C) Palpus. (D) Wing. (E) Genitalia.
of the spermatheca long, stout and oblique, the pointed gonostili and the male sternite 9 rounded distally. The third species herein described, Dasyhelea allegrae n. sp., belongs to the mutabilis species group. Its adult male is very similar to the Patagonian species D. ingrami Díaz & Spinelli by virtue of the tergite 9 rounded distally, the paramere and gonocoxal apodemes forming an asymmetrical structure and the aedeagus lacking a posteromedian projection and with the posterolateral arms single, divergent and with recurved pointed tips; however, the male of D. ingrami differs by the apicolateral process triangular, the gonostili with wide base and tapering abruptly at midlength,
its tip hook-like and the stouter paramere with blunt tip. Apart from the peculiar respiratory organ, the pupa of this new species is very similar to D. pseudolacustris Díaz & Spinelli and D. serrana Díaz & Spinelli, both recently described by Diaz et al. (2014) from Argentinean Patagonia; however, D. pseudolacustris differs by the rounded tubercle of the posterior margin of the dorsal apotome with 6–7 strong spicules, the clypeal/labrals short and hyaline, the smooth surface of the cephalotorax and the respiratory organ slender with 42–45 apical and 6–8 lateral pores; a detailed revision of the pupa of D. pseudolacustris reveals that it has two dorsal apotome sensilla and three dorsolateral sensilla instead of one and two
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Fig. 9. Dasyhelea allegrae Díaz, (A–C) adult male, (D–G) male pupa. (A) Genitalia. (B) Paramere and gonocoxal apodemes. (C) Aedeagus. (D) Cephalothoracic sensilla (dorsal view). (E) Dorsal sensilla. (F) Metathorax and tergite 1. (G) Segment 4. Anterolateral sensilla (AL-1-T, AL-2-T, AL-3-T); anteromedial sensilla (AM-1-T, AM-2-T); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H, Dl-3-H); dorsal sensilla (D1-T, D-2-T, D-3-T); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H, DL-3-H); metathoracic sensilla (M-2-T, M-3-T); segment 4 sensilla (D-1-IV, D-2-IV, D-3-IV, D-4-IV, D-5-IV, D-7-IV, D-8-IV, L-1-IV, L-2-IV, L-3-IV, L-4-IV, V-5-IV,V-6-IV, V7-IV), supraalar campaniform sensillum (SA-2-T); tergite 1 sensilla (D-2-I, D-4-I, D-7-I, D-8-I, L-1-I, L-2-I, L-3-I).
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respectively, as they were originally described (Diaz et al., 2014); the pupa of D. serrana can be distinguished by the yellowish exuviae, the dorsal surface of the dorsal apotome covered with strong spicules and bearing one dorsal apotome sensillum, the two short clypeal/labrals, the respiratory organ with base and apex smooth and with 6–7 apical and 5–6 lateral pores and the seta D-8-IV of the fourth abdominal segment short and thin. 5. Bionomics Larvae and pupae of D. azteca were collected in March 2009 and spring 2011, 2014 from small temporary ponds near of margin of Rio de La Plata River and stream in Corrientes. Larvae lasted three days to reach to pupal stage in the laboratory and the pupae that reached the adult stage took 3–4 days to emerge. The pupae of D. yunga and D. allegrae were collected in November 2015 from arroyo Los Tablaos and Río Las Quintas respectively and lasted 2–3 days to reach the adult stage. Pupae exibit the typical semi-circular slow adbominal movement typical of other ceratopogonid pupae. Acknowledgments We are grateful to Nélida Caligaris for technical assistance and Mónica Caviglia for the English proofreading. This work was supported by Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Argentina. PIP 03051 and PIP 0568. References Borkent, A., Spinelli, G.R., 2007. Neotropical Ceratopogonidae (Diptera: Insecta). In: Adis, J., Arias, J.R., Rueda-Delgado, G., Wantzen, K.M. (Eds.), Aquatic Biodiversity in Latin America (ABLA), vol. 4. Pensoft, Sofia-Moscow, pp. 198. Borkent, A., 2014. The pupae of the biting midges of the world (Diptera: ceratopogonidae), with a generic key and analysis of the phylogenetic relationships between genera. Zootaxa 3879, 1–327.
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Borkent, A., 2016. World Species of Biting Midges (Diptera: Ceratopogonidae), http://www.inhs.uiuc.edu/research/FLYTREE/CeratopogonidaeCatalog.pdf. Last update May 16 2016. 245 pp. Brown, A.D., Grau, A., Lomáscolo, T., Gasparri, N.I., 2002. Una estrategia de ˜ (Yungas) de Argentina. conservación para las selvas subtropicales de montana Ecotrópicos 15, 147–159. Brown, A.D., Pacheco, S., Lomáscolo, T., Malizia, L., 2006. Situación ambiental de los ˜ bosques Andino Yunguenos. In: Brown, A.D., Martinez Ortiz, U., Acerbi, M., Corcuera, J. (Eds.), La Situación Ambiental Argentina 2005. Fundación Vida Silvestre Argentina, Buenos Aires, pp. 53–56. Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E., Zumbado, M.A., 2009. Manual of Central American Diptera, vol. 1. NRC Research Press, Ottawa, Ontario, Canada, 714 pp. Cabrera, A.L., Willink, A., 1973. Biogeografía de América Latina Washington D.C. OEA. Monografía n◦ 13, Serie Biología., pp. 109. Díaz, F., Ronderos, M.M., Spinelli, G.R., Ferreira-Keppler, R.L., Torreias, S.R., 2013. A new species of Dasyhelea Kieffer (Diptera: Ceratopogonidae) from brazilian amazonia. Zootaxa 3686 (1), 085–093. Diaz, F., Spinelli, G.R., Donato, M., Ronderos, M.M., 2014. A phylogenetic analysis of the mutabilis species group of Dasyhelea Kieffer, and a taxonomic revision of the Patagonian species (Diptera: ceratopogonidae). J. Nat. Hist. 48 (35–36), 2117–2175. Dominiak, P., 2012. Biting midges of the genus Dasyhelea Kieffer (Diptera: Ceratopogonidae) in Poland. Pol. J. Entomol. 81, 211–304. Huerta, H., Grogan, W.L.Jr., 2006. A new species and new record of biting midges of the genus Dasyhelea Kieffer (Diptera: Cetatopogonidae) from Morelos and Jalisco, Mexico. Proceedings of the Entomological Society of Washington 108 (4), 892–898. Kieffer, J.J., 1911. Nouvelles descriptions de chironomides obtenus d’éclosion. Bulletin de la Société de Moselle (Metz) 27, 1–60. Morrone, J.J., 2001. Biogeografía de América Latina y el Caribe. Manuales y Tesis SEA 3 Zaragoza (Spain)., pp. 148. Ronderos, M.M., Spinelli, G.R., Sarmiento, P., 2000. Preparation and mounting of biting midges of the genus Culicoides Latreille (Diptera: ceratopogonidae) to be observed with a scanning electron microscope. Trans. Am. Entomol. Soc. 126 (1), 125–132. Ronderos, M.M., Díaz, F., Sarmiento, P., 2008. A new method using acid to clean and a technique for preparation of eggs of biting midges (Diptera: Ceratopogonidae) for scanning electron microscope. Trans. Am. Entomol. Soc. 134 (3+4), 471–476. Waugh, W.T., Wirth, W.W., 1976. A revision of the genus Dasyhelea Kieffer of the eastern United States north of Florida (Diptera: Ceratopogonidae). Ann. Entomol. Soc. Am. 69 (2), 219–247.