Family Ceratopogonidae

MARINO P,; DÍAZ F,; RONDEROS, M. M.; FERREIRA KEPPLER R.

Key to Neotropical Hexapoda

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Año: 2018; p. 1 - 811

Ceratopogonidae is placed in the infraorder Culicomorpha (Borkent, 2012). Their common names are ?polvorines, manta blanca, chaquistes or jejenes? (common name shared with Simuliidae) in Spanish speaking countries; ?mosquito pólvora or maruim? in Brazil; and ?biting midges, no-see-ums or punkies? in English speaking countries (Spinelli & Ronderos, 2005). It is an ancestral group, with numerous fossil records in amber from 17 to around 121 million years old (Borkent & Spinelli, 2007). It is one of the most diverse family of Diptera, occurring on all continents except Antarctica and in most habitats, including deserts. They are present from the tropics to the high Arctic and from lowlands to high mountains (4,200 meters on Mount Everest, Nepal) (Borkent, 2014). Despite their great diversity and their nearly ubiquitous presence, the immatures of Ceratopogonidae are still the least known among the biting Diptera.The family includes six subfamilies, four of which are extant and worldwide in distribution, with 111 extant genera and 6,267 species. In addition, the fossil record includes another 284 extinct species and 21 extinct genera (Borkent, 2016). Four subfamilies inhabit the Neotropical region: Leptoconopinae, Forcipomyiinae, Dasyheleinae and Ceratopogoninae (in seven tribes: Culicoidini Kieffer, Ceratopogonini, Heteromyiini Wirth, Johannsenomyiini Crampton, Sphaeromiini Newman, Palpomyiini Enderlein, and Stenoxenini Coquillett) including more than 1,225 species in 52 genera, of which, 16 are cosmopolitan (Borkent, 2016; Santarém & Felippe-Bauer, 2016).Members of this family are holometabolous, with the immature stages inhabiting aquatic, semiaquatic or terrestrial habitats. Females usually lay their eggs on or in organically rich aquatic or wet habitats where larvae complete their development and pupate. Oviposition microhabitats are very diverse (Fig. 16.1.1A-H). Egg production varies from 10 to 600, depending on the genera and species (Díaz et al., 2005). Egg clutches can be laid in strings or masses that are cemented with gelatinous substances (Fig. 16.1.2A-F), depending on subfamily (Díaz et al., 2005; Ronderos et al., 2006). In Forcipomyiinae eggs are oval to elongate, black or dark brown and glossy; in Dasyheleinae (Fig. 16.1.2A-B, E,) they are horseshoe or C-shaped and enclosed in a gelatinous film. In Ceratopogoninae (Fig. 16.1.2C-D, F), eggs are laid in loose groups on moist substrate and are slender, banana shaped with a thin or often sculptured chorion, e.g. Culicoidini (Fig. 16.1.2F), or in strings or masses with gelatinous coatings e.g. Palpomyiini (Fig. 16.1.2D). Aquatic or semiaquatic larvae inhabit the superficial layer of the breeding site, usually at depths of 0-12 cm (Uslu & Dick, 2010; Gonzalez Gonzalez de Heredia & Goldarazena Lafuente, 2011). Active larvae can be recognized by their slow to very rapid serpentine. At the end of the final 4th instar, larvae develop pupal respiratory organs and also accumulate a large amount of body fat in the abdomen that distinguish late 4th instar or prepupa. Ceratopogonidae larvae have diverse feeding habits: filter feeders, detritivores, or carnivores and others feed on phytoplankton or small debris, however, many are omnivores (McCafferty, 1981). A summary of the morphology and food items of larvae of species belonging to 52 genera of Ceratopogonidae is presented in Figs. 16.1.3A-G; 16.1.5A-H. From a morpho-behavioral perspective, this variation is most evident in the size and shape of the head capsule, mandibles, epipharynx, and hypostoma.Regarding larval locomotion, ceratopogonid larvae can be grouped in two categories based on their locomotory abilities: non swimming (crawling) or swimming. Larvae of this family are unique among dipterous larvae in their adaptations for swimming. The morphological characteristics of the caudal segment are related to the environment in which they live (Fig. 16.1.8A-G).During the pupal stage, most aquatic species float on the surface of water where they rest and respire, but, if disturbed they may thrash about and/or sink. There are many questions about whether pupal character states are useful for differentiating closely related genera and species, as well as if they possess morphological characters of group and intergroup relationships. Their respiratory organs and terminal processes of segment 9 are often unique to genus, group and species. Fox (1942) studied the correlation between some morphological structures of the pupae of Culicoides and suggested that variation in the respiratory organ and the segment 9 are related to environment. Differences between genera and species could also be due to the different environments where they develop. Borkent & Craig (2001) noted associations between breeding sites and respiratory organs, length of body setae and terminal processes of the pupa of Stilobezzia rabelloi Lane; which was later confirmed for several other ceratopogonids including S. punctulata Lane, Dasyhelea paulistana Forattini & Rabello, D. pseudopollinosa Díaz & Ronderos and Culicoides lacustris Ronderos. In order to obtain air, the pupae of these five species insert the pointed respiratory organ into the floating or dangling submerged leaves of an aquatic plants (e.g. Salvinia auriculata Aubl., Salvinaceae, Azolla filiculoides Lam., Azollaceae). After emergence, adults are pale (teneral), poorly sclerotized, and require about 24h to achieve full coloration. Males typically emerge earlier than females, and they are fertile about 24h after emergence (Gonzalez Gonzalez de Heredia & Goldarazena Lafuente, 2011).