Forcipomyia Meigen
Creators: name: Borkent, Art
Publisher: name: Zenodo
Year: 2014
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Diptera
Ceratopogonidae
Forcipomyia

description: Forcipomyia Meigen
(Figs. 2A–B, 8D–H, 9A–H, 10A–H, 11A–C, 14C–D, 18C–D, 23C–D, 29D–F, 32C, 34C, 42E–R, 47C–D, 55C, 56A, 72F–I)
DIAGNOSIS: Only pupa of Ceratopogonidae without a prothoracic extension (Figs. 23C–D), with the labium not divided medially by an elongate suture (as is present in Fig. 23F) and with the terminal processes either closely abutting basally (Figs. 8D–H, 9A–H, 10A–H, 11A–D, 72F–H) or the terminal process spicules (if present) directed posteriorly to posterolaterally (not anteriorly); combination of retaining larval exuviae on the abdomen or with very elongate tubercles on the thorax (Figs. 9A, C, F, 10A–E) and/or abdomen, and the terminal process with spicules (if present) directed posteriorly to posterolaterally (not anteriorly) is unique to many species.
DESCRIPTION: Habitus as in Figs. 8D–H, 9A–H, 10A–H, 11A–C. Total length = 1.69–3.03 mm. With (as in Fig. 11F) or without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face (Fig. 14D). Ecdysial tear to anterior margin of base of antenna (Figs. 14C–D, 79B); along posterolateral portion of eye. Head: Dorsal apotome (Figs. 18C–D), without ventral line of weakness, with or without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13B) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Figs. 23C–D) with mandible, lacinia well-developed, not overlapping apically; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 34C) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Figs. 18C–D)—absent or 1 very short seta; dorsolateral cephalic sclerite sensilla—absent or 1 short seta, 1 campaniform sensillum; clypeal-labrals (Figs. 23C–D)—1 seta, 1 campaniform sensillum; oculars (Figs. 23C–D)—1 campaniform sensillum. Thorax: Prothoracic extension (Figs. 23C–D) absent; mesonotum with or without elongate tubercles, extending posteromedially, completely dividing metathorax medially (Figs. 47C–D); respiratory organ (Figs. 42E–R) length/width = 1.05–4.75, highly variable, knob-like to elongate, slender to nearly spherical, with or without posterior basal swelling or lobe, somewhat flattened laterally or circular in cross-section, with pores closely abutting at apex of respiratory organ, arranged in single line of varying shapes or two widely spaced rows, outer surface smooth or with spicules, without or with short pedicel, base without posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, smooth or with spirals restricted to base; wing (Fig. 34C) with apical tubercle or angle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (Figs. 10C, 32C) broadly to narrowly separate; halter apex abutting anterolateral edge of tergite 2; legs (Fig. 34C) with lateral margin of foreleg near midlength of wing slightly sinuous to evenly curved; hind leg visible at lateral margin of wing (Fig. 32C); with apex of foreleg moderately to well anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—0–2 campaniform sensilla; anterolaterals—with or without 1 short or elongate seta or 1 campaniform sensillum; dorsal setae (Figs. 29D–F)—D-1-T seta, D-2-T seta present or absent, D-3-T campaniform sensillum; D-2-T, D-3-T on single tubercle in some, or closely associated (if D-2-T present); supraalar 2—campaniform sensillum; metathoracics (Figs. 47C–D)—0, 2 campaniform sensilla; M-3-T near anterior margin of metathorax. Abdomen: without pigmentation pattern, segment 2 as wide or somewhat wider than segment 3, segments with undivided, thin to thick setae, without tubercles or segments 2–8 with spines or thick tubercles or segments 1–5 with branched or thick spined elongate tubercles, tubercles rounded to pointed, tergites or sternites entire, each without membranous disc; segment 9 (Figs. 72F–I) highly variable in shape, terminal processes widely separated to closely approximated basally, each projecting posteriorly to laterally, tapering to pointed apex or tapering near apex to point; sensilla: tergite 1 (Figs. 47C–D) with 3–5 setae, 1–2 campaniform sensilla, including 1–2 lateral sensilla, D-2-I, D-3-I (if present) well separated, D-7-I absent or situated posteriorly near D-8-I; segment 4 (Figs. 55C, 56A)—D-2-IV peg-like to slender seta, D-3-IV slender seta, without tubercles or very short separate ones; D-8-IV, when present, a moderately elongate seta; D-5-IV present or absent, D-7-IV, D-9-IV absent; D-4-IV without to on moderately elongate tubercle, posterior dorsal sensilla with D-4-IV, D-8-IV lateral to each other or D-8-IV absent, L-1-IV short to elongate seta present on short to long tubercle, close to L-2-IV, L-2-IV, L-3-IV, L-4-IV on separate rounded to elongate tubercles, in some with L-2-IV, L-3-IV closely approximated, V-5-IV, V-6-IV, V-7-IV short to moderately elongate setae, V-5-IV present or absent, on separate tubercles or V-6-IV, V-7-IV on single tubercle; segment 8 without D-3-VIII, without L-1-VIII; segment 9 (Figs. 72F–I)—D-5-IX campaniform sensillum, D-6-IX campaniform sensillum or short or elongate seta.
DISTRIBUTION AND HABITAT: The genus Forcipomyia is known from 1150 species in 35 subgenera from every Region worldwide (Borkent 2014). The larvae and pupae are found in many semi-terrestrial to moist terrestrial habitats, including under loose bark of logs, under decaying leaves and manure, decomposing vegetable matter and fungi, moss on soil, rotting wood or rocks, sap at tree wounds, in and above the water of various phytotelmata, mud along streams, bogs, and shallow water ponds.
TAXONOMIC DISCUSSION: There are 146 species of Forcipomyia, in 14 subgenera, known as pupae (Tables 2–3). The pupae of this genus vary tremendously morphologically and are rich in numerous derived features, including overall body form, chaetotaxy, shape of respiratory organs and more. The group is ripe for a cladistic analysis and combined with their various behaviours and diverse niches, are certain to provide a rich opportunity to examine diversification within the genus. A few features were put into a phylogenetic context by Chan & LeRoux (1971c).
Although earlier work had shown that pupae of Forcipomyia have a variety of interesting modifications (e.g. Long 1902, Speiser 1910, Carter 1919, Rieth 1915), it was not until L.G. Saunders (1924, 1925, 1957, 1959, 1964), in a series of remarkable papers, studied the group that the wealth of features and their importance in their classification became apparent. He described exemplars from the 10 available subgenera (out of 14 for which immatures are known today), showing that the larvae, pupae and adults had congruent and distinctive features, indicating a solid basis for their classification (although derived features were not interpreted). He provided a consistent format for the description of the pupae and laid the groundwork for all future studies. Work by Chan & Saunders (1965), Chan & LeRoux (1965, 1970, 1971a, 1971b) and Chan & Linley (1989) added significantly with further detailed descriptions of Forcipomyia pupae.
The pupae are distinctive enough that a key to the subgenera is possible (see above).
Further differences exist at the species level and early work indicated these also could be keyed (Malloch 1915 — Illinois; Lenz 1934 —Europe; Wirth & Howarth 1982 —Hawaiian F. (Euprojoannisia); Lewańczyk et al. 2010 —European species of F. (Forcipomyia )).
Wirth & Howarth (1982), Chen et al. (1980), Spinelli et al. (2995) and Marino et al. (2010) provided SEM photomicrographs of pupae of Forcipomyia.
I initially thought that the subgenera F. (Lepidohelea) and F. (Schizoforcipomyia) could be separated on the basis of the relative size of the medial tubercle on the dorsal apotome. Previous descriptions of species of F. (Lepidohelea) show these to be long. The only species of F. (Schizoforcipomyia) to be described for this feature is F. borbonica by Chan & LeRoux (1971b, as F. petersoni), who illustrate the pupa as having a short, rounded tubercle on the middle of the dorsal apotome, although in the text they indicate this to be "elongate". I examined the two paratype pupal exuviae of F. petersoni and found both to have an elongate tubercle, indistinguishable from those of species of F. (Lepidohelea).
Pupae of Forcipomyia have reduced numbers of thoracic sensilla, as well as fewer on the dorsum of the abdominal segments and the exact naming of these was uncertain. D-3-T and D-4-IV are likely correct as unique campaniform sensilla (in these areas).
MATERIAL EXAMINED: F. aeria: 2 pupal exuviae (of paratype), Mayaquez, Puerto Rico, 18-III-1953 (USNM). F. anabaenae: 1 pupal exuviae (of paratype), Singapore, 25-III-1962 (CNCI). F. antiguensis: 1 pupal exuviae (of paratype), Falmouth Harbour, Antigua, 6-V-1953 (CNCI). F. ashantii: 1 pupal exuviae, Tafo, Ghana, 17-V-1963 (ANIC); Berger's Hall, Hazyview, East Transvaal, South Africa, 3-XII-1973 (ANIC). F. attenuata: 1 pupal exuviae (of paratype), La Lola, Costa Rica, 2-VI-1956 (CNCI). F. bipunctata: 7 pupa, 23 pupal exuviae, 4 mi. W., 3 mi. S Schuyler, Nebraska, USA, various dates, 1990-1991 (CNCI). F. blantoni: 2 pupal exuviae, Turrialba, Costa Rica, 2-IX-1964 (USNM); 2 pupal exuviae, as previous locality, 26-XI-1964 (USNM); 1 pupal exuviae, as previous locality, 21-XII-1964 (USNM); 1 pupal exuviae, as previous locality, 29-XII-1964 (USNM). F. borbonica: 1 pupal exuviae (labeled as paratype of an undescribed species F. lerouxi), Wallace Way, Singapore, 1961 (LEMQ). F. brasiliensis: 1 pupal exuviae, Maracas, Trinidad, 18-VII-1957 (USNM). F. bromeliae: 1 pupal exuviae (of paratype), Rio de Janeiro, Brazil, 23-VIII-1923 (CNCI). F. bystraki: 1 pupal exuviae, Calvert, Cecil County, Maryland, USA, 29-III-1969 (USNM); 2 pupal exuviae, Headwaters, Highland County, Virginia, USA, VI-1969 (USNM). F. caerulea: 1 pupal exuviae (of paratype), Rio de Janeiro, Brazil, 8-VIII-1923 (CNCI). F. canadensis: 2 pupae (of paratypes), Pike Lake, Saskatchewan, Canada, X-1957 (USNM). F. caribbeana: 1 pupal exuviae (of paratype), Trinidad, 12-V-1953 (CNCI). F. cerifera: 1 pupal exuviae, Rio de Janeiro, Brazil, 1-VIII- 1923 (CNCI). F. cornuta: 1 pupal exuviae (of paratype), Costa Rica, 4-VI-1956 (CNCI). F. crinita: 1 pupal exuviae (of paratype), Emma Lake, Saskatoon, Saskatchewan, Canada, 19-IV-1957 (CNCI). F. edmistoni: 1 pupal exuviae (of paratype), Allegany State Park, New York, USA, 28-V-3-VI-1963 (USNM); 1 pupal exuviae (of paratype), Bittinger 4-H Campground, Garrett County, Maryland, USA, 5-V-1960 (USNM); 1 pupa (of paratype), 1 pupal exuviae (of paratype), Patuxent Wildlife Refuge, Maryland, USA, 28-VII-1979 (USNM). F. elegantula: 1 pupal exuviae, Saskatoon, Saskatchewan, Canada, 3-IX-1950 (USNM); 1 pupa, 1 pupal exuviae, Plummer's Island, Maryland, USA, 3-VI-1976 (USNM); 1 pupal exuviae, as previous locality, 10-VI-1976 (USNM); 1 pupa, 2 pupal exuviae (in glycerin), as previous locality, 10-VI-1976 (WLGC). F. flavescens: 1 pupal exuviae (of paratype), Luna, P.I., 25-X-1961 (CNCI). F. harpegonata: 1 pupa (of paratype), Puerto Rica, 23-I-1953 (USNM). F. herediae: 1 pupal exuviae (of paratype), Siquirres, Costa Rica, 1-VI-1956 (USNM). F. hutsoni: 1 pupa (of paratype), South Island, Aldabra, 1-17-II-1968 (USNM). F. hygrophila: 3 pupae, Sidney, British Columbia, Canada, 20-VIII-1947 (USNM); 2 pupal exuviae, no locality data (USNM). F. intermedia: 1 pupal exuviae (of paratype), nr. Siquirres, Costa Rica, 15-V-1956 (CNCI). F. jocosa: 1 pupal exuviae (of paratype), Mt. St. Benedict, Trinidad, 13-V-1953 (CNCI). F. lesliei: 2 pupae, Clark Hall, Dominica, 9-I-1965 (USNM); 1 pupa (of paratype of F. bicolor), 2 pupal exuviae (of paratypes of F. bicolor), Mayaquez, Puerto Rico, 3-II-1953 (USNM). F. luteigenua: 1 pupa (of paratype), 1 pupal exuviae (of paratype), Finca la Tigra, nr. La Virgen, Heredia Province, Costa Rica, 12-XI-1981 (USNM); 1 pupa (of paratype), as previous locality, 17-XI-1981 (USNM); 1 pupa (of paratype), as previous locality, 30-VII-1982 (USNM). F. malayae: 2 pupal exuviae (of holotype, paratype), Cameron’s Highlands, Malaysia, 29-I-1934 (CNCI). F. mortuifolii: 1 pupal exuviae, no locality data (USNM). F. musae: 2 pupal exuviae, km 123 Br 174, Manaus, Amazonas, Brazil, 29-VII-2005 (CNCI). F. nodosa: 1 pupal exuviae (of paratype), nr. Siquirres, Costa Rica, 15-VII-1956 (CNCI). F. oligarthra: 1 pupal exuviae (of paratype), Carcega Beach, Puerto Rico, 22-II-1953 (CNCI). F. pictoni: 1 pupa, 2 pupal exuviae, Siquirres, Costa Rica, 11-VI-1956 (USNM). F. pinicola: 2 pupal exuviae, no locality (USNM). F. pulchrithorax: 3 pupae, Cambridge, England, United Kingdom, 31-VIII-1922 (USNM). F. quasicornuta: 1 pupal exuviae (of paratype), nr. Siquirres, Costa Rica, 6-V-1050 (CNCI). F. seminole: 2 pupal exuviae, Hills, Puerto Rico, 6-II-1953 (USNM); 2 pupae, 2 pupal exuviae, Georgetown, Guyana, 20-V-1943 (USNM). F. sonora: 2 pupae, 2 pupal exuviae, Death Valley, Saratoga Springs, California, USA, 19-II-1955 (USNM). F.spinosa: 2 pupae (of paratypes), 1 pupal exuviae, Mayaquez, Puerto Rico, 3-II-1953 (USNM); 1 pupal exuviae (of paratype), as previous locality, 3-II-1953 (CNCI); 1 pupal exuviae, Ilheus, Bahia, Brazil, 20-X-1977 (USNM). F. terrestris: 1 pupal exuviae (of paratype), St. Augustine, Trinidad, 9-VII- 1957 (CNCI); 1 pupa (of paratype), 3 pupal exuviae (of paratype), Las Hermanas, Trinidad, 9-VII-1957 (USNM). F. trinidadensis: 1 pupal exuviae (of holotype), St. Augustine, Trinidad, 3-VII-1957 (CNCI). F. tuberculata: 1 pupa, 2 pupal exuviae, Hacienda Theobroma, Siquirres, Costa Rica, 1-VI-1956 (USNM); 1 pupa, Bayano Field Station, Panama Province, Panama, VI-1976 (USNM); 1 pupal exuviae (of paratype), Trinidad, 8-V-1953 (CNCI); 1 pupa (of paratype), 2 pupal exuviae (of paratypes), North Range, Trinidad, 8-V-1953 (USNM). F. usingeri: 3 pupae (of paratypes), Berkeley Hills, Alameda County, California, USA, X-1947 (USNM). F. varicolor: 1 pupal exuviae (of paratype), Rio de Janeiro, Brazil, 1-VIII-1923 (CNCI). F. wirthi: 1 pupal exuviae (of paratype), Alum Rock Park, Santa Clara County, California, USA, 8-IV-1948 (USNM). F. (Phytohelea) nr. antiguensis: 3 pupal exuviae, 1 pupal exuviae (in glycerin), Riverwoods Field Laboratory, nr. Cornwall, Highlands County, Florida, USA, 13-I-1999 (CNCI). F. (Euprojoannisia) sp.: 1 pupal exuviae, 52 km S of airport, Iquitos, Loreto, Peru, 1-II- 2003 (CNCI). F. (Phytohelea) sp.: 2 pupal exuviae, 2 pupal exuviae (in glycerin), Jardin Botanico, Rio Piedras, San Juan, Puerto Rico, 24-X-2006 (CNCI). F. sp.: 1 pupa, 1 pupal exuviae (in glycerin), Nebraska, USA (CNCI); 1 pupal exuviae, Rio Dantes, P.N. Barbilla, Cartago, Costa Rica, 10-II-2006 (CNCI); 2 pupal exuviae, Iquitos, Loreto, Peru, 1-II-2003 (CNCI); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 19-II-1967 (BPBM); 1 pupal exuviae, as previous locality, 9-12-XII-1967 (BPBM).
description: Published as part of Borkent, Art, 2014, The Pupae of the Biting Midges of the World (Diptera: Ceratopogonidae), With a Generic Key and Analysis of the Phylogenetic Relationships Between Genera, pp. 1-327 in Zootaxa 3879 (1) on pages 40-41, DOI: 10.11646/zootaxa.3879.1.1, http://zenodo.org/record/4949051