Research Article |
Corresponding author: Danielle Anjos-Santos ( danianjos_santos@comahue-conicet.gob.ar ) Academic editor: Art Borkent
© 2018 Danielle Anjos-Santos, Florentina Díaz, Gustavo Ricardo Spinelli, María Marcela Ronderos.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Anjos-Santos D, Díaz F, Spinelli GR, Ronderos MM (2018) The immatures of Bezzia chilensis Spinelli & Ronderos, 2001 (Diptera, Ceratopogonidae). ZooKeys 803: 141-154. https://doi.org/10.3897/zookeys.803.29024
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The fourth instar larva and the pupa of Bezzia chilensis Spinelli & Ronderos, 2001 are described for the first time. The immature stages were collected from macrophytes and filamentous algae in streams of the Patagonian steppe, in the provinces of Neuquén and Chubut, Argentina. The described stages were photographed and illustrated with a phase-contrast microscope and scanning electron microscope. Data on the bionomics of the species, new records and tables for characters of the known larvae and pupae of Bezzia Kieffer, 1899 from the Neotropical region are provided.
Aquatic, biting midges, immature stages, Neotropical region, Palpomyiini
Bezzia Kieffer, 1899, a worldwide genus of the tribe Palpomyiini, includes 322 species of which 48 inhabit the Neotropical region, 46 of them recorded by
Bezzia chilensis Spinelli & Ronderos, 2001 is a member of the venustula species group in the subgenus Homobezzia Macfie, 1932, distributed in Valparaiso Province (Chile), and Salta and Río Negro provinces (Argentina) (
Larvae and pupae were collected on the bordering vegetation in three streams on the Patagonian steppe in the provinces of Neuquén and Chubut. The substrate was removed with the aid of a strainer and transferred to a white tray where larvae and pupae were collected with a pipette. Further substrate samples were carried to the laboratory to search for more specimens. Larvae were placed in individual containers with water and substrate from their natural environment. Pupae were 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 in ethanol to ensure their complete pigmentation. For detailed examination with a phase-contrast microscope, larval and pupal exuviae and adults were mounted in Canada balsam following the technique described by
Bezzia
chilensis
:
(Figs
Bezzia chilensis Spinelli & Ronderos, 2001, fourth instar larva (SEM) a Head chaetotaxy, ventrolateral view b Head chaetotaxy, dorsolateral view c Head capsule detail, oblique anteroventral view d Head capsule detail, ventral view e Mouthpart, ventral view f Hypostoma and mouthparts, ventral view. Antennae (AN); collar (CO); fossa mandibularis (MF); galeolacinea (GL); hypostoma (HY); labrum (LB); lacinial sclerite 1 (LC1); lacinial sclerite 2 (LC2); mandible (MD); messors (MS); maxilla (MX); maxillary palpus (MP); palatal bar (PB); palatum (PL); sensilla campaniformia (SCa); sensilla styloconica (SS); sensilla trichoidea (ST); Head capsule chaetotaxy: o, parahypostomal setae; p, posterior perifrontal setae; q, postfrontal setae; s, anteroperifrontal setae; t, prefrontal setae; u, mesolateral setae; v, posterolateral setae; w, anterolateral setae; x, parantennal setae; y, ventral setae.
Bezzia chilensis Spinelli & Ronderos, 2001, fourth instar larva a Head capsule detail, ventral view b Caudal segment, ventral view c Epipharynx, ventral view d Left mandible, ventral view. Dorsal comb (DC); epipharynx (epy); fossa mandibularis (MF); hypopharynx (hyp); mandible (MD); messors (MS); ventral comb (VC). Caudal segment chaetotaxy: i, inner setae; l1, first lateral seta; o, outer setae. Scale bars: 0.05 mm (a–c), 0.01 mm (d).
Female (Figs
Bezzia chilensis Spinelli & Ronderos, 2001, male pupa (a–d), female pupa (e–g) a Habitus, ventral view b, f Dorsal apotome, dorsal view c, g Segment 9, ventral view d Respiratory organ, anterodorsal view (SEM) e Respiratory organ ventral view. Antenna (AN); dorsal apotome sensilla (DA-1-H, DA-2-H); genital lobe (GL); pore (p); pedicel (P); respiratory organ (MD); segment 1 (1st seg.); segment 4 (4th seg.); segment 9 (Seg. 9); terminal process (TP). Scale bar: 0.05 mm.
Bezzia chilensis Spinelli & Ronderos, 2001, female pupa a Mouthparts, ventral view b Anterolateral, anteromedial and dorsal lateral sensilla, ventral view c Dorsal sensilla and supraalar sensillum, dorsal view d Metathoracics chaetotaxy, dorsal view e Tergite 1 chaetotaxy, dorsal view f Segment 4 chaetotaxy, dorsal and ventral view. Anterolateral sensillum (AL-1-T); anteromedial sensilla (AM-1-T, AM-2-T, AM-3-T) ; clypeal/labral sensilla (CL-1-H, CL-2-H); dorsal sensilla (D-1-T, D-2-T, D-3-T, D-4-T, D-5-T); dorsal sensilla of segment 1 (D-2-I, D-3-I, D-4-I, D-5-I, D-7-I, D-8-I, D-9-I); dorsal sensilla of segment 4 (D-2-IV, D-3-I, D-4-I, D-5-IV, D-7-IV, D-8-IV, D-9-IV); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H); lateral sensilla of segment 1 (L-1-I, L-2-I, L-3-I); lateral sensilla of segment 4 (L-1-IV, L-2-IV, L-3-IV, L-4-IV); ocular sensilla (O-1-H, O-2-H, O-3-H); metathoracic sensillum (M-3-T); supraalar sensillum (SA-2-T); ventral sensilla of segment 4 (V-5-IV, V-6-IV, V-7-IV). Scale bar: 0.05 mm.
Male (Fig.
Argentina, Neuquén Province, Parque Nacional Nahuel Huapi, río Cuyín Manzano, 40°44'13"S, 71°09'17"W, alt. 760 m, 06-II-2009, A. Siri, 2 females and 1 male (with pupal exuviae). Argentina, Chubut Province, Ruta Nacional 40, arroyo La Cancha, 42°45'35.9"S, 71°06'28.4"W, alt. 860 m, 13-II-2015, adults emerged in laboratory 14-II-2015, D. Anjos-Santos and P. Pessacq, 1 female, 3 males (with pupal exuviae); same data except adult emerged 15-II-2015, 1 male (with pupal exuviae); same data except adults emerged 17-II-2015, 2 females, 1 male (with pupal exuviae); same data except adults emerged 18-II-2015, 2 females, 1 male (with pupal exuviae); same data except adults emerged 17-II-2015, 1 female, 1 male (with pupal exuviae); same data except pupa emerged in laboratory 26-II-2015, adult emerged 28-II-2015, 1 female (with larval and pupal exuviae); same data except adult emerged 03-III-2015, 1 female (with pupal exuviae); Argentina, Chubut prov., Ruta Nacional 40, arroyo Madera, 42°39'57.59"S, 71°04'19.72"W, alt 930 m, 18-II-2015, pupa emerged in laboratory 23-II-2015, adult emerged 28-II-2015, D. Anjos-Santos and P. Pessacq, 1 male (with larval and pupal exuviae); same data except pupae emerged 23-II-2015, adults emerged 02-III-2015, 2 females (with larval and pupal exuviae); same data except pupa emerged 24-II-2015, 1 male (with pupal exuviae); same data except pupa emerged 26-II-2015, 1 male (with pupal exuviae); Argentina, Chubut Province, arroyo Montoso, 42°42'01.26"S; 70°48'12.36"W, alt. 630 m, 14-I-2016, pupa emerged in laboratory 16-I-2016, D. Anjos-Santos and P. Pessacq, 1 male (with pupal exuviae); same data except pupa emerged 17-I-2016, adult emerged 21-I-2016, 1 female (with larval and pupal exuviae); same data except pupa emerged 17-I-2016, adult emerged 22-I-2016, 1 male (with larval and pupal exuviae); same data except pupa emerged 20-I-2016, adult emerged 25-I-2016, 1 male (with larval and pupal exuviae).
Argentina, Chubut Province, Ruta Nacional 40, arroyo Madera, 42°39'57.59"S, 71°04'19.72"W, alt. 930 m, 18-II-2015, D. Anjos-Santos and P. Pessacq, 3 larvae, 1 male pupa.
Argentina (Salta, Neuquén, Río Negro and Chubut provinces); Chile (Valparaiso Province).
The immature described here were collected in northwestern Argentine Patagonian steppe (Fig.
In a series of contributions reviewing the Neotropical Bezzia,
The immatures of Bezzia chilensis are herein compared with four species belonging to the subgenus Homobezzia; these four are the only ones which have a complete description: B. blantoni (described by
The larva of Bezzia chilensis shares with B. blantoni, B. galesa and B. roldani features typical of predatory larvae: hooked mandibles with fossa mandibularis, epipharynx less massive with 2 combs and cylindrical maxillary palpus (
Main diagnostic characters for the known larvae of Neotropical species of Bezzia.
Subgenus | species | Head capsule ratio | Head capsule setae | Hypostoma | Fossa mandibularis | Scopae | Maxillary palpus | Hypopharyngeal fringe | Caudal segment | Reference |
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Bezzia | bivittata | ? | ? | Finely toothed | Deep | Absent | 3 subapical papillae | Absent | ? |
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nobilis | 2.7 times as long as wide | Medium-sized to long | Finely toothed | Deep | Absent | ? | Absent | 2.5 times as long as wide |
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Homobezzia | blantoni | 2.6–3.4 times as long as wide | Minute | Finely toothed, not flanked by stout teeth | Shallow | Present, with 5 teeth | 2–3 subapical papillae | Present | 4 times as long as wide |
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chilensis | 2 times as long as wide | Medium-sized to long | Finely toothed flanked by 6–8 stout teeth | Deep | Absent | 4 subapical papillae | Absent | 2 times as long as wide | This study | |
galesa | 2.8 times as long as wide | Minute | Finely toothed, with strong lateral teeth | Deep | Absent | 4 subapical papillae | Absent | 5–6 times as long as wide |
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glabra | 4 times as long as wide | ? | ? | ? | ? | ? | ? | 10 times as long as wide |
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roldani | 3.4 times as long as wide | Minute | Finely toothed, not flanked by stout teeth | Shallow | Absent | 2–3 subapical papillae | Absent | 5 times as long as wide |
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With regard to the pupa, besides the features given in Table
Main diagnostic characters for the known pupae of Neotropical species of Bezzia.
Subgenus | Species | Dorsal apotomals sensilla | Dorsolateral cephalic sclerite sensilla | Clypeal/labral sensilla | Oculars sensilla | Respiratory organ apex | Respiratory organ pores | Sensillum D-7-IV | Terminal process of segment 9 | References |
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Bezzia | nobilis | 1 seta, 2 campaniform | 3 setae | 2 setae | 1 seta, 1 campaniform | Rounded | 16–25 on distal 1/4 | Present | Short |
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gibbera | 1 seta, 1 campaniform | ? | ? | ? | ? | ? | ? | ? |
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Homobezzia | blantoni | 1 seta, 2 campaniform | 1 seta | 2 setae | 2 setae | Rounded | 31–35 on distal apex | Absent | Long |
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brevicornis | 1 seta, 1 campaniform | ? | ? | ? | Asymmetrical (outer side curved, inner side straight) | 23–25 on distal 1/3 | ? | Short |
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bromeliae | 1 seta | ? | ? | ? | Asymmetrical (outer side curved, inner side straight) | 4 on distal apex | ? | Short |
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chilensis | 1 seta, 1 campaniform | 1 seta, 1 campaniform | 2 setae | 2 setae, 1 campaniform | Rounded | 31–41 on distal apex and apicolateral 1/4 | Present | Short | This study | |
galesa | 1 seta, 2 campaniform | 1 seta | 2 setae | 2 setae, 1 campaniform | Rounded | 50–60 on distal apex, 10–11 lateral ones | Absent | Long |
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glabra | 1 seta, 2 campaniform | ? | ? | ? | Rounded | 50–60 on distal 1/3 | ? | Long |
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pulchripes | 1 seta, 2 campaniform | ? | ? | ? | Bilobed | 40–52 on distal apex | Absent | Long | Mayer (1959) | |
roldani | 1 seta, 2 campaniform | 2 setae | 2–3 setae | 2 setae | Rounded | 50–60 on distal 1/3 | Absent | Long |
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snowi | 1 seta, 1 campaniform | ? | ? | ? | Rounded | 11–12 on distal 1/4 | ? | Short |
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ventanensis | 1 seta, 1 campaniform | 2 setae | 2 setae | 1 seta, 1 campaniform | Asymmetrical (outer side curved, inner side straight) | 13–15 on distal 1/3 | Absent | Short |
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In addition, a detailed revision during this study revealed that D-4-T of B. galesa is a seta and was erroneously described as campaniform sensillum by
The pupae of Bezzia chilensis and the other Neotropical known pupae of Bezzia share the features of the generic description given by
We are grateful to Lic. Luz Manzo for technical assistance and Dr Miguel Archangelsky for the critical reading of the manuscript. This work was funded by the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET, Argentina), PIP114 201101 00405 and PIP 0305. This is the contribution number 131 of LIESA (CIEMEP).