The pupa of <i>Stilobezzia (Eukraiohelea) elegantula</i> (Johannsen) (Diptera: Ceratopogonidae)

DÍAZ, Florentina; AYALA, Mahia M.; LEGUIZAMÓN, Alan; FERREIRA-KEPPLER, Ruth L.; SPINELLI, Gustavo R.

doi:10.1590/1809-4392202303471

ABSTRACT

The pupa of Stilobezzia (Eukraiohelea) elegantula is described and illustrated, being the first description of a pupa for a species belonging to this subgenus. The studied material was collected large banks of floating aquatic macrophytes in the Marchantaria Island, in the municipality of Iranduba, Amazonas, Brazil, located at 15 km from Manaus, and surrounded by the white waters of the Solimões river. The collected pupae were isolated and observed in the laboratory until the adult emergence and then identified as S. (E.) elegantula. The described pupa is compared with those of Neotropical species Stilobezzia (Stilobezzia) punctulata and S. (S.) rabelloi.

KEYWORDS:
Immatures; Predaceous midges; taxonomy

RESUMEN

Se describe e ilustra la pupa de Stilobezzia (Eukraiohelea) elegantula siendo la primera descripción de una pupa de una especie perteneciente al subgénero. El material estudiado fue colectado en la Isla Marchantaria sobre grandes bancos de macrófitas acuáticos flotantes, municipio de Iranduba, Amazonas, Brasil, localizada a 15 km de Manaus, y rodeada por las aguas del río Solimões. Las pupas colectadas fueron aisladas y observadas en el laboratorio hasta la emergencia del adulto y luego identificadas como S. (E.) elegantula. La pupa descrita se compara con las especies neotropicales Stilobezzia (Stilobezzia) punctulata y S. (S.) rabelloi.

PALABRAS CLAVE:
Inmaduros; Mosquitas depredadoras; taxonomía

INTRODUCTION

The predaceous midge genus StilobezziaKieffer, 1911 is a large and diverse group of Ceratopogonidae with about 361 species, occurring throughout the zoogeographical regions described worldwide (Borkent and Dominiak 2020; Borkent et al. 2022; Cazorla and Da Silva, 2023; Da Silva et al. 2023), except Antarctica (Borkent 2014).

Adult females are important predators on other small insects, and the immature stages are found in a wide variety of aquatic and semiaquatic habitats, including streams, lake and pond margins, puddles, swamps, rice fields, rock pools and tree holes, and feed on microorganisms (De Meillon and Wirth 1991; Cazorla et al. 2006). Four subgenera are recognized within the genus, the cosmopolitan S. (Acanthohelea) Kieffer, 1917 and S. (Stilobezzia) sensu strictu with 338 extant species described, S. (Debenhamia) Wirth and Grogan 1988, known by two species from Australia, and S. (Eukraiohelea) Ingram and Macfie 1921, with 14 species from Africa, America and southeastern Asia (Borkent and Dominiak 2020; Borkent et al. 2022; Cazorla and Da Silva 2023; Da Silva et al. 2023). Borkent (2014) listed 29 species of Stilobezzia for which the pupae are known worldwide, six of them in the subgenus S. (Acanthohelea) and 23 in the subgenus S. (Stilobezzia), and subsequently Anjos-Santos et al. (2017) described the pupa of S. (A.) megatheca Cazorla and Spinelli from Patagonia.

Currently, the subgenus S. (Eukraiohelea) is represented in the Neotropical region by five species recently reviewed by Cazorla et al. (2017): S. amnigena (Macfie) and S. elegantula (Johannsen) from southern and eastern USA to northeastern Argentina, and S. dorsofasciata (Lutz), S. proxima Cazorla and Felippe-Bauer and S. quasielegantula Cazorla and Felippe-Bauer, restricted to Rio de Janeiro, Brazil. The only described immature within the subgenus is the weak, very superficial description of the egg of S. dorsofasciata by Lutz (1914).

During a field sampling program in the vicinity of Manaus, Brazil, pupae of Stilobezzia sp. were collected. They were isolated and observed in the laboratory until the adult emergence and then identified as S. (E.) elegantula. The pupa of this species is herein described and illustrated, being this the first description for one species in this subgenus.

MATERIAL AND METHODS

Study area

The present study was conducted at Camaleão Lake on the Marchantaria Island, in the municipality of Iranduba, Amazonas, Brazil. This island at 15 km from the city of Manaus is surrounded by the white water of the Solimões river. The area is directly exposed to the flood-pulse by the water level fluctuations of the Amazon and its tributaries (Wittmann et al. 2010). The climate is equatorial-neotropical, with monthly mean minimum temperatures of 24-26°C and maximums of 28°C (Salati and Vose 1984).

Specimens were collected between April and June 2011 during the high-water period of the Rio Solimões (Junk 1989) in large banks of floating aquatic macrophytes (Figure 1), with a predominance of Eichhornia crassipes (Mart.) Solms (03°15’20.4’’S, 59°57’55.9’’W) or Pistia stratiotes L. (03°14’27.5’’S, 59°57’30.4’’W). In these environments, the water temperatura at the plant root level was 29.8°C, the conductivity 243.8 µS/cm and the pH 9.76.

Figure 1
Geographical location of Lago Camaleão in wetlands at Ilha da Marchantaria, Iranduba, Amazonas, Brasil. B-E show the collection sites of the immature stages of Stilobezzia.

The pupae were collected with a metallic ladle and separated manually in white plastic trays using plastic pipettes, into smaller glass cups with river-water and transported to the Laboratory of Cytotaxonomy of Aquatic Insects- LACIA in a box thermal plastic container. 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, pupal exuviae and adults were mounted in Canada balsam following the technique described by Borkent and Spinelli (2007). Photomicrographs were taken with a digital camera Micrometrics SE Premium, through a Nikon Eclipse E200 microscope. For terminology see Borkent (2014) and Anjos-Santos et al. (2017).

The material examined is deposited in the following collections: INPA- Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil; MLPA- División Entomología, Museo de La Plata, Argentina.

RESULTS

Stilobezzia (Eukraiohelea) elegantula (Johannsen)

Zoobank identifier: urn:lsid:zoobank.org:pub:85069F97-F8AC-42C1-9116-4CF002492721

Material examined

BRAZIL, AMAZONAS, Iranduba, Ilha da Marchantaria, Lago Camaleão, 03°15’20.4’’S; 59°57’55.9’’W, 18-iv-2011, Torreias & Ferreira-Keppler lgt., 2 ♂ (with ‘ex pupa’), (1 in INPA; 1 in MLPA); 1 ♀ (with ‘ex pupa’) (INPA); same date except 03°14’27.5’’S; 59°57’30.4’’W 27-vi-2011, 1 ♀ (with ‘ex pupa’) (MLPA).

Description of female pupa

(Figure 2B, E-F, H; 3A, E). Exuviae overall pale brown. Total length 2.87-3.00 (2.94, n = 2) mm. Head: Dorsal apotome (Figure 2B) smooth, anterior margin rounded with small, hyaline notched; distal margin slightly concave with a few wrinkles; DAL 0.11-0.13 (0.205, n = 2) mm; DAW 0.14-0.16 (0.15, n = 2) mm; DAW/DAL 1.07-1.45 (1.26, n = 2); mouthparts with mandible well developed; palpus extending to posterolateral margin of labium; labium separated medially by labrum; apex of labrum rounded. Sensilla: dorsal apotomals (Figure 2B): DA-1-H elongate, thin seta, located on small tubercle, DA-2-H campaniform sensillum at tubercle base; one clypeal/labrals: CL-1-H long, thin seta; oculars: O-2-H campaniform sensillum. Thorax: Cephalothorax rectangular, surface smooth with small wrinkles on all surfaces. Length of cephalothorax 0.95-1.02 (0.98, n=2) mm, width 0.85-0.90 (0.97, n = 2) mm. Respiratory organ (Figure 2E) brown, smooth, medium-sized, narrowing gradually to pointed apex, with 28-30 pores in single row; pedicel slender (Figure 2E, H-G), P 0.04-0.05 (0.045, n = 2) mm; RO length 0.27-0.30 (0.28, n = 2) mm, RO width 0.04-0.06(0.05, n = 2) mm; P/RO 0.04-0.06 (0.05, n = 2). Sensilla: two dorsolateral cephalic sclerite (Figure 2F): DL-1-H long, thin seta, DL-2-H campaniform sensillum; two anteromedial setae: AM-1-T, AM-2-T long, thin setae; two anterolaterals: AL-1-T, AL-2-T very long, stout setae, AL-2-T shorter than AL-1-T; dorsals (Figure 2H): D-1-T long, stout seta; D-2-T, D-4-T long, thin setae, D-3-T campaniform sensillum, D-1-T, D-2-T, D-4-T on small tubercle, closely approximated; SA-1-T campaniform sensillum; metathoracics: M-1-T very long, thin seta, M-2-T campaniform sensillum. Abdomen: segments 3-9 covered with very small spicules on anterior margin; tergite 1 (Figure 3A) with two anteromesals: D-2-I, D-3-I long, thin setae; D-4-I, D-7-I campaniform sensilla, D-7-I near to D-3-I, D-8-I, D-9-I long, thin setae, D-8-I stouter than D-9-I; three lateral sensilla L-1-I, L-3-I long, stout setae, L-2-I long, thin seta, all on stout tubercle; segment 4: D-2-IV medium-sized, thin seta, D-4-I, D-7-IV campaniform sensilla, D-7-IV near to D-2-IV, D-5-IV, D-9-IV long, thin setae, D-8-IV long, stout seta; L-1-IV, L-2-IV, L-3-IV, L-4-IV long, stout setae, all on cylindrical tubercles; V-5-IV very long, stout seta, V-6-IV, V-7-IV long, thin setae; segment 9 (Figure 3E) approximately 1.27 times as long as width, length 0.60-0.62 (0.61, n = 2) mm, width 0.15 (n = 2) mm; terminal process very elongate, parallel, length 0.45-0.50 (0.47, n = 2) mm, with D-5-IX campaniform sensillum.

Figure 2
Morphological details pupa of the Stilobezzia (Eukraiohelea) elegantula (Johannsen) pupa. A, C-D, G, I-J: Male pupa. B, E-F, H: Female pupa. A- Entire pupa. B-C dorsal apotome. D - clypeal/labral sensillum and ocular sensillum. E - respiratory organ. F - dorsolateral cephalic sclerite sensilla. G - Anterolateral sensilla and anteromedial sensilla. H - dorsal sensilla. I - metathorax and tergite 1. J - metathoracic sensilla. Scale bars: 0.05 mm. Abbreviations: Anterolateral sensilla (AL-1-T, AL-2-T); anteromedial sensilla (AM-1-T, AM-2-T); clypeal labral sensillum (CL-1-H); dorsal apotome (DA); dorsal apotome sensilla (DA-1-H, DA-2-H); dorsolateral cephalic sclerite sensilla (DL-1-H, DL-2-H); dorsal sensilla (D-1-T, D-2-T, D-3-T, D-4-T); genital lobe (GL); metathoracic sensilla (M-2-T, M-3-T); oculars sensillum (O-2-H); pedicel (P); respiratory organ (RO); tergite 1 sensilla (D-2-I, D-3-1, D-4-I, D-7-I, L-1-I, L-2-I, L-3-I); terminal process (TP). Scale bars = 0.05 mm

Figure 3
Morphological detail of the Stilobezzia (Eukraiohelea) elegantula (Johannsen) pupa. A, E: Female pupa. B-D, F: Male pupa. A - tergite 1. B Segment 4 dorsal sensilla. C - Segment 4 lateral sensilla. D - Segment 4 ventral sensilla. E-F - Segment 9, ventral view. Scale bars: 5 mm. Abbreviations: Genital lobe (GL); Segment 4 sensilla (D-3-IV, D-4-IV, D-5-IV, D-7-IV, L-1-IV, L-2-IV, L-3-IV, L-4-IV, V-5-IV, V-6-IV, V-7-IV); tergite 1 sensilla (D-2-I, D-3-I, D-4-I, D-7-I, D-8-I, D-9-I, L-1-I, L-2-I, L-3-I); terminal processes (TP).

Description of male pupa

(Figure 2A, C-D, G, I-J; 3B-D, F). Habitus as in Figure 2A. Similar to female with usual sexual differences: Total length 2.80-3.00 (2.90, n = 2) mm. Dorsal apotome (Figure 2C) with DAL 0.19-0.22 (0.205, n = 2) mm; DAW 0.09-0.10 (0.095, n = 2) mm; DAW/DAL 1.90-2.44 (2.17, n = 2), clypeal-labrals as in Figure 2D. Respiratory organ, P 0.024-0.045 (n = 2) mm; RO length 0.28-0.30 (0.29, n = 2) mm, RO width 0.05 (n = 2) mm; P/RO 0.16-0.18 (0.17, n = 2) mm; anteromedials and anterolaterals as in Figure 2G. Cephalothorax: length 0.80 mm, width 1.02 mm, metathoracis and tergite 1 as in Figure 2I-J; segment 4 (Figure 3B-D). Segment 9 (Figure 3F) length 0.57-0.67 (0.62, n = 2) mm, width 0.15-0.20 (0.17, n = 2) mm; genital lobe (Figure 3F) elongate, just reaching to posterior margin of segment; terminal process length 0.42-0.45 (0.43n = 2) mm.

DISCUSSION

Several species of Ceratopogonidae breed in macrophytes, for example the Neotropical S. (S.) punctulata Lane and S. (S.) rabelloi Lane and other genera, such as Dasyhelea Kieffer, Forcipomyia Kieffer, Culicoides Latreille, Heteromyia Say and Bezzia Kieffer. As pointed out by Borkent and Craig (2001) and Cazorla and Marino (2004), the pupae share characters related with the breeding site, as the respiratory organ narrowing gradually to pointed apex, smooth dorsal apotome, the tubercles of the abdominal segment with simple or branched setae, and very elongated terminal processes.

Stilobezzia (S.) punctulata is readily distinguished from S. (E.) elegantula by the pale brown cepahalothorax and darker abdomen, the dorsal apotome with DA-1-H stout seta, presence of two clypeal/ labrals and two oculars, respiratory organ with two rows of pores (basal row with 5 pores and apicolateral row with 18 pores), the sensilla of segment 4 with bifurcated setae except D-2-IV and L-1-IV, which are simple setae, and male ninth segment with genital lobe not reaching its posterior margin.

Stilobezzia (S.) rabelloi can be distinguished from S. (E.) elegantula by the general pale brown coloration except the apical portion of segment 7 and segments 8-9, which are slightly darker the dorsal apotome with DA-1-H represented by a stout seta, DL-1-H stout seta, presence of two clypeal/labrals and two oculars, respiratory organ with 5 basal and 15 apical pores, AL-1-T and AL-2-T thick setae, dorsals D-1-T; D-2-T and D-4-T with long thin setae, segment 4 with D-2-IV pore, and D-5-IV, D-8-IV, D-9-IV, V-5-IV, V-6-IV are bifurcated with plumose apices.

CONCLUSIONS

Of the 111 extant genera of Ceratopogonidae, the immature stages of only 45 of them are known at present. Therefore, there is still a need for larger collection efforts to document the immature stages of this family and their breeding environments. Because of this, it is necessary to obtain more associated material in order to have a complete idea of the preimagos of the genera of the family and to be able to know the different environments where they breed. Finally, knowing the habitats of the immatures and the precise identification of these immatures is a valuable tool to be able to recommend control measures and avoid outbreaks of the species suspected to be vectors of viruses such as Bluetongue or Oropuche, , which affect both animals and humans.

ACKNOWLEDGEMENTS

Financial support was provided by Agencia Nacional de Promoción Científica y Técnica, Argentina (PICT No. 2021-I-A 01008) and MCTI/INPA/PRJ. 12.272. RLFK received research grants 306081/2013-0 and 307039/2016-1 from Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq). This study was partially funded by Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) -Finance Code 001 and Fundação de Amparo à Pesquisa do Estado do Amazonas (FAPEAM).

REFERENCES

Anjos-Santos, D.; Cazorla, C. G.; Ronderos, M. M. 2017. First description of immature stages ad adult females of Stilobezzia (Acanthohelea) megatheca (Diptera: Ceratopogonidae) from Patagonia. Annales de Limnologie, International Journal of Limnology 53: 175-188.
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.
Borkent, A.; Craig, D. A. 2001. Submerged Stilobezzia rabelloi Lane (Diptera: Ceratopogonidae) pupae obtain oxygen from the aquatic fern Salvinia minima Baker. Proceedings of the Entomological Society of Washington 103: 655-665.
Borkent, A.; Dominiak, P. 2020. Catalog of the biting midges of the World (Diptera: Ceratopogonidae). Zootaxa 4787: 1-377.
Borkent, A.; Grogan, W. 2009. Catalog of the New World Biting Midges North of Mexico (Diptera: Ceratopogonidae). Zootaxa 2273: 1-48.
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 Vol. 4. Pensoft, Sofia-Moscow, p. 1-198.
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CITE AS:
Díaz, F.; Ayala, M.M.; Leguizamón, A.; Ferreira-Keppler, R.L.; Spinelli, G.R. 2025. The pupa of Stilobezzia (Eukraiohelea) elegantula (Johannsen) (Diptera: Ceratopogonidae). Acta Amazonica 55: e55bc23347.

Data availability

The data that support the findings of this study were published in this article.

Edited by

ASSOCIATE EDITOR:
Pitágoras da Conceição Bispo https://orcid.org/0000-0002-7356-8882

Publication Dates

Publication in this collection
17 Nov 2025
Date of issue
2025

History

Received
21 Nov 2023
Accepted
26 Nov 2024

This is an open-access article distributed under the terms of the Creative Commons Attribution License

[1] Anjos-Santos, D.; Cazorla, C. G.; Ronderos, M. M. 2017. First description of immature stages ad adult females of Stilobezzia (Acanthohelea) megatheca (Diptera: Ceratopogonidae) from Patagonia. Annales de Limnologie, International Journal of Limnology 53: 175-188.

[2] 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.

[3] Borkent, A.; Craig, D. A. 2001. Submerged Stilobezzia rabelloi Lane (Diptera: Ceratopogonidae) pupae obtain oxygen from the aquatic fern Salvinia minima Baker. Proceedings of the Entomological Society of Washington 103: 655-665.

[4] Borkent, A.; Dominiak, P. 2020. Catalog of the biting midges of the World (Diptera: Ceratopogonidae). Zootaxa 4787: 1-377.

[5] Borkent, A.; Grogan, W. 2009. Catalog of the New World Biting Midges North of Mexico (Diptera: Ceratopogonidae). Zootaxa 2273: 1-48.

[6] 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 Vol. 4. Pensoft, Sofia-Moscow, p. 1-198.

[7] Borkent, A.; Dominiak, P.; Díaz, F. 2022. An update and errata for the Catalog of the biting midges of the World (Diptera: Ceratopogonidae) Zootaxa 5120: 53-64.

[8] Cazorla, C.G.; Da Silva, I.M. 2023. Neotropical Stilobezzia Kieffer, 1911 (Diptera: Ceratopogonidae): a new species, redescriptions, new records, and synonymy. Annales Zoologici 73: 341-354.

[9] Cazorla, C.G.; Marino, P.I. (2004) The pupa of Stilobezzia punctulata Lane from Peruvian Amazonia. (Diptera: Ceratopogonidae). Amazoniana, 18, 75-80.

[10] Cazorla, C.G.; Díaz, F.; Ronderos, M.M. 2006. Redescription of pupa and adult of Stilobezzia fiebrigi Kieffer 1917 (Diptera: Ceratopogonidae). Transactions of the American Entomological Society 132: 111-119.

[11] Cazorla, C.G.; Cardoso, E.A.; Felippe-Bauer, M.L. 2017. Contributions to the knowledge of predaceous midges of the subgenus Eukraiohelea Ingram & Macfie of Stilobezzia Kieffer (Diptera: Ceratopogonidae), from Brazil. Zootaxa 4324: 557-570.

[12] Da Silva, I.M; Ferreira-Keppler, R.L.; Cazorla, C.G. 2023: Stilobezzia Kieffer (Diptera: Ceratopogonidae) from the Brazilian Amazon: three new species, redescription of S. (Stilobezzia) maculata Lane and new records. Zootaxa 5249 (4): 485-499.

[13] De Meillon, B.; Wirth, W.W. 1991. The genera and subgenera (excluding Culicoides) of the Afrotropical biting midges (Diptera: Ceratopogonidae). Annals of the Natal Museum 32: 27-147.

[14] Ingram, A; Macfie, JWS. 1931. Ceratopogonidae. In: Diptera of Patagonia and South Chile Part II. Fasc. 4 (London: British Museum): 155-232.

[15] Junk, W. 1989. Flood tolerance and tree distribution in central Amazonian floodplains. In: Holm-Nielsen, L.B.; Nielsen, I.C.; Balslev, H. (Eds.): Tropical Forest: botanical dynamics, speciation and diversity Academy Press, London. p. 47-64

[16] Kieffer, J.J. 1911. Nouvelles descriptions de chironomides obtenus d’éclosion. Bulletin de la Société de Moselle (Metz) 27: 1-60.

[17] Kieffer, J.J. 1917. Chironomides d’Amérique conservés au Musée National Hongrois de Budapest. Annales Historico-Naturales Musei Nationalis Hungarici 15: 175-228.

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[19] Salati, E.; Vose, P.B. 1984. Amazon Basin: a system in equilibrium. Science 225: 129-138.

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The pupa of Stilobezzia (Eukraiohelea) elegantula (Johannsen) (Diptera: Ceratopogonidae)

A pupa de Stilobezzia (Eukraiohelea) elegantula (Johannsen) (Diptera: Ceratopogonidae)