Anebomyia Borkent, 2014

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, Zootaxa 3879 (1), pp. 1-327 : 92-95

Anebomyia Borkent new genus

( Figs. 21L , 27G , 30T , 39F , 45T , 52E , 68A , 76K–L )

Anebomyia Borkent, 2014 : in this work.

Type species: Mallochohelea atripes Wirth, 1962: 281 , by present designation. Included species and taxonomic discussion: given below.

Etymology: Anebos-myia (beardless-fly) refers to the lack of sternite 8 tufts on female adults of species in this genus.

DIAGNOSIS: Only pupa of Ceratopogonidae with the dorsal apotome with two pairs of tubercles ( Fig. 21L ).

DESCRIPTION: Total length = 3.41–4.91 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16B , 33B ). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17C , 79H ). Head: Dorsal apotome ( Fig. 21L ), with ventral line of weakness, with additional dorsal pair dorsomedial tubercles, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 27G ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 39F ) barely posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 21L )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 27G )—2 slender setae; oculars ( Fig. 27G )—2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 27G ) wide, well-developed but narrow dorsolaterally, not extending to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 52E ); respiratory organ ( Fig. 45T ) length/width = 2.91–5.40, moderately elongate, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface with some wrinkles, with short, wide pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing ( Fig. 39F ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33A ) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs ( Fig. 39F ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33I ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—2 elongate setae (as in Figs. 31L–M ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 30T )—D-1-T, D-2-T, D-4-T, D-5-T setae, D-3-T campaniform sensillum, D-3-T anterolateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 52E )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: pigmentation light to dark brown, with tergite 1 with 3 medial spots, tergites 2–7 with medial area with stripe, 2 anterolateral spots, sternites 3–7 with pale to well defined medial stripe, anterolateral spot, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to pointed, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Figs. 76K–L ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 52E ) with 8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D-2-I, D-3-I closely approximated, D- 7-I situated anteriorly near D-3-I; segment 4 ( Fig. 68A )—D-2-IV, D-3-IV moderately elongate setae on short tubercles; D-5-IV, D-8-IV, D-9-IV moderately elongate setae; D-5-IV on single tubercle, D-8-IV, D-9-IV on separate but closely approximated tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D-5-IV, D-4-IV, D-8-IV, D-9-IV; D-7-IV near D-3-IV; L-1-IV short seta on pointed tubercle, near L-3-IV; L-2-IV, L-3-IV, L-4-IV moderately elongate setae on pointed tubercles, V-5-IV, V-6-IV, V-7-IV moderately elongate setae on short tubercles, all closely approximated; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Figs. 76 K-L)—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Anebomyia is known from five species from the Nearctic, Afrotropical and Oriental Regions (all previously in Mallochohelea ; Borkent 2014). Immatures are known from stream and river margins, lakes and ponds. Knausenberger (1987) discusses the various habitats in which A. atripes was collected in the eastern USA. At least some pupae were on emergent vegetation (although they do not have abdominal membranous discs).

TAXONOMIC DISCUSSION: Pupae are known from two species ( Tables 2–3 ). All the species placed here in the new genus Anebomyia were previously in Mallochohelea . The new genus is recognized because the newly included species are not related to those now restricted to Mallochohelea , a genus most closely related to Jenkinshelea , Macropeza , Probezzia and Neobezzia (see phylogenetic analysis below, Fig. 81 ).

There are differences between the two genera in both male and female adults and in pupae, with most of the adult features not yet interpreted cladistically (but see character 85). They are summarized as follows:

Knausenberger (1987: 267-277) provided further differences between the two genera (as albibasis and atripes groups), including a larval feature. His information was based on Nearctic species and the character states need further testing with extraterritorial material.

As indicated above, members of Anebomyia differ from true Mallochohelea by the absence of distinctive tufts of elongate setae on sternite 8 of female adults. In A. atripes , elongate setae are absent. In A. fluminea there are elongate setae on sternite 8 but these are not arranged as a tight tuft (as is true in Mallochohelea and other related genera; see character 85). Examination of female adult M. texensis indicates that it too lacks a well defined sternite 8 tuft.

Wirth (1962), in erecting Mallochohelea , pointed out that some of the included species such as M. atripes , M. spinipes and M. texensis differed considerably from the genotype ( M. albibasis ) and "may, eventually need a separate category". He further pointed out in a strikingly prescient statement "The discovery of the pupa of this group should also serve greatly to clarify its relationships". The pupae have indeed provided the synapomorphies to clarify the relationship of these species.

Although Wirth (1962) considered M. spinipes as closely related to A. atripes , I have not included it in Anebomyia . I have not examined material of this species and know only that the adult female has spines on its femora; the sternite 8 tufts are not described and the male remains unknown. Like this and a number of other species presently in Mallochohelea , reexamination of specimens is needed before the names are potentially transferred to the new genus. Of course, knowing the pupae of further species will certainly clarify their taxonomic position.

All European species of Mallochohelea fit the characterization of this genus (Patrycja Dominiak, pers. comm.). The female of the European/Asian species M. tianshanica , however, has spines on the femora and welldeveloped sternite 8 setal tufts (Patrycja Dominiak, pers. comm.) and the male has basally fused parameres ( Remm 1980). This suggests that M. tianshanica is a true Mallochohelea but is the only species known with femoral spines. Debenham's (1974) description of Mallochohelea and the included Australian species suggests that they all actually belong within Mallochohelea . The presence of femoral spines on the Afrotropical species M. kirki ( de Meillon & Wirth 1981) may indicate they are members of Anebomyia but the presence or absence of the female sternite 8 setal tufts is unknown and males are undescribed, making its placement uncertain. The African species M. siricis has a female with spines on the forefemur and the male has separate parameres; although the condition of the female sternite 8 seta tufts is uncertain, I consider this sufficient evidence to place it in Anebomyia . The Chinese species M. yunnana , known only as a female, has femora spines and lacks the sternite 8 setal tufts, indicating it too is an Anebomyia .

Included species:

Anebomyia atripes (Wirth), 1962: 281 ( Mallochohelea ). Type locality: New Brunswick, New Jersey, USA. new combination.

Anebomyia fluminea (de Meillon & Wirth), 1981: 550 ( Mallochohelea ). Type locality: Sand River between White River and Hazyview, E. Transvaal, South Africa. new combination.

Anebomyia siricis (de Meillon), 1961: 50 ( Sphaeromias ). Type locality: Fenerive, Madagascar. new combination.

Anebomyia texensis (Wirth), 1962: 283 ( Mallochohelea ). Type locality: Kerrville, Texas, USA. new combination.

Anebomyia yunnana (Yu & Zou) , in Yu et al. 2005: 1503 ( Mallochohelea ). Malipo, Yunnan Province, China. new combination.

MATERIAL EXAMINED: A. atripes : 1 pupal exuviae, Black Lake, North Burgess Township, Ontario, Canada, 30-VI-1971 (CNCI); 1 pupal exuviae, Patuxent Wildlife Research Center, Prince George County, Maryland, USA, 4-VI-1976 (USNM); 2 pupal exuviae, as previous locality, 24-V-1977 (USNM); 3 pupal exuviae, as previous locality, 13-VI-1977 (USNM); 2 pupal exuviae, as previous locality, 1977 (USNM); 1 pupal exuviae, Lower 3 Runs Creek, Savannah R. Plantation, Barnwell County, South Carolina, USA, 16-II-1979 (WLGC). A. fluminea : 1 pupal exuviae (of holotype), Sand River between White River and Hazyview, East Transvaal, South Africa, 2-XII-1973 (NMSA).

Figures

FIGURE 13. Anterior portion of heads and thoraces, in anterior or anterodorsal view. A. Leptoconops kerteszi, ecdysial sutures of head enhanced to show position. B. Culicoides denticulatus. C. Ceratopogon nr. abstrusus. D. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). E. Alluaudomyia bella, dorsal apotome absent. F. Clinohelea curriei. G. Pellucidomyia leei. H. Mallochohelea caudellii. I. Bezzia sp. (from Bolean Lake, BC, Canada).

FIGURE 16. Anterior portion of heads. A. Parabezzia huberti, in ventral view (distorted slide specimen). B. Parabezzia sp. (5 km. E. Danby, Vermont, USA), ventrolateral view. C. Clinohelea curriei, in ventral view. D. Heteromyia clavata, in ventral view, dorsal apotome not present. E. Heteromyia wokei, in ventral, slightly lateral view. F. Pellucidomyia leei, in ventral view, dorsal apotome not present.

FIGURE 17. Anterior portion of heads. A. Jenkinshelea sp., in ventral view (from Ottawa, ON, Canada). B. Mallochohelea caudellii., in ventral view. C. Sphaeromias longipennis, in ventral view. D. Bezzia sp., in ventral view (6 km E Salmon Arm, BC, Canada). E. Palpomyia sp. (Black Lake, Quebec, Canada), in ventral view. F. Phaenobezzia sp., in ventral view (from Skulsuza, South Africa).

FIGURE 21. Dorsal apotomes, anterior view. A. Dibezzia prominens (from Mayer 1934c). B. Hebetula tonnoiri (from Elson- Harris 1987). C. Jenkinshelea albaria (from Wirth & Grogan 1979). D. Macropeza albitarsis (from Szadziewski & Dominiak, 2007). E. Mallochohelea termophila (from Spinelli 1984, modified). F. Probezzia pallida (from Wirth & Grogan 1979). G. Neobezzia fittkaui (from Ronderos et al. 2011, modified). H. Nilobezzia basispinigera (from Elson-Harris 1987). I. Nilobezzia virago (from Elson-Harris 1987). J. Johannsenomyia argentata (from Wirth & Grogan 1979). K. Lanatomyia electra (from Elson-Harris 1987). L. Anebomyia atripes (from Wirth & Grogan 1979, modified).

FIGURE 27. Posterior portion of heads, in ventral view. A. Mallochohelea caudellii. B. Probezzia sabroskyi. C. Neobezzia fittkaui. D. Nilobezzia curticornis. E. Johannsenomyia argentata. F. Lanatomyia electra. G. Anebomyia atripes. H. Homohelea delanoe.

FIGURE 30. Dorsal setae from left side of thoraces. A. Serromyia sp. B. Allohelea japonica. C. Monohelea obscura. D. Atyphohelea macroneura. E. Parabezzia huberti. F. Echinohelea lanei. G. Clinohelea curriei. H. Heteromyia clavata. I. Pellucidomyia leei. J. Dibezzia debenhamae (dirty specimen, D-5-T perhaps broken setal socket). K. Hebetula tonnoiri. L. Jenkinshelea magnipennis. M. Macropeza natalensis. N. Mallochohelea caudellii. O. Probezzia sabroskyi. P. Neobezzia fittkaui (from Ronderos et al. 2011, modified). Q. Nilobezzia curticornis. R. Johannsenomyia argentata. S. Lanatomyia miles. T. Anebomyia atripes.

FIGURE 31. Dorsal setae from left side of thoraces; anteromedial setae. A. Homohelea delanoe. B. Leehelea hispida. C. Sphaeromias longipennis. D. Xenohelea galatea. E. Bezzia nobilis. F. Clastrieromyia dycei. G. Pachyhelea pachymera. H. Palpomyia jonesi. I. Phaenobezzia mashonensis. J. Paryphoconus oliveirai. K. Echinohelea lanei, partial lateral view (distorted slide specimen). L. Palpomyia (JAD685), dorsal view. M. Palpomyia (JAD685), dorsolateral view. N. Probezzia seminigra, dorsal view of left side.

FIGURE 33. Cephalothoraces and abdominal segments 1–3, in lateral view. A. Parabezzia sp. (5 km. E. Danby, Vermont, USA). B. Clinohelea curriei. C. Pellucidomyia leei. D. Hebetula tonnoiri. E. Jenkinshelea sp. (from Ottawa, ON, Canada). F. Mallochohelea caudellii. G. Probezzia seminigra. H. Nilobezzia sp. (from Kruger NP, South Africa). I. Sphaeromias longipennis. J. Bezzia sp. (6 km E. Salmon Arm, BC, Canada) K. Palpomyia sp. (Black Lake, Quebec, Canada). L. Phaenobezzia sp. (from Skulsuza, South Africa).

FIGURE 39. Legs, wings and apices of the antennae, in ventral view. A. Probezzia sabroskyi. B. Neobezzia fittkaui. C. Nilobezzia sp. (from Australia). D. Johannsenomyia argentata. E. Lanatomyia electra. F. Anebomyia atripes.

FIGURE 45. Respiratory organs, in dorsal view, pattern on tracheal tubes shown only in some. A. Clinohelea tasmaniensis (from Elson-Harris & Kettle 1986). B. Clinohelea bimaculata (from Wirth & Grogan 1979, modified). C. Heteromyia wokei (from Wirth & Grogan 1977). D. Heteromyia clavata (from Wirth & Grogan 1977). E. Pellucidomyia leei (from Elson-Harris & Kettle 1986). F. Pellucidomyia geari (from de Meillon & Wirth 1981). G. Dibezzia prominens (from Mayer 1934c). H. Hebetula tonnoiri (from Debenham 1974). I. Jenkinshelea albaria (from Wirth & Grogan 1979). J. Macropeza albitarsis (from Szadziewski & Dominiak, 2007). K. Mallochohelea albihalter (from Wirth & Grogan 1979). L. Probezzia pallida (from Wirth & Grogan 1979). M. Probezzia xanthogaster (from Wirth & Grogan 1979). N. Neobezzia fittkaui (from Ronderos et al. 2011). O. Nilobezzia aranea (from Elson-Harris 1987). P. Nilobezzia virago (from Elson-Harris 1987). Q. Johannsenomyia argentata (from Wirth & Grogan 1979). R. Lanatomyia electra (from Debenham 1974). S. Lanatomyia miles (from Debenham 1974). T. Anebomyia atripes (from Wirth & Grogan 1979).

FIGURE 52. Metathoraces and tergites 1, in dorsal view. A. Neobezzia fittkaui. B. Nilobezzia curticornis. C. Johannsenomyia argentata. D. Lanatomyia miles. E. Anebomyia atripes. F. Homohelea delanoe (reconstructed from distorted and dirty specimen).

FIGURE 68. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Anebomyia atripes. B. Homohelea delanoe. C. Leehelea hispida (reconstructed from distorted specimen).

FIGURE 76. Abdominal segments 9. A. Macropeza albitarsis, male, in ventral view (modified from Szadziewski & Dominiak, 2007). B. Macropeza albitarsis, female, in ventral view (from Szadziewski & Dominiak, 2007). C. Mallochohelea termophila, male, in ventral view (from Spinelli 1984). D. Mallochohelea termophila, female, in dorsal view (from Spinelli 1984). E. Probezzia pallida, male, in dorsal and ventral view (from Wirth & Grogan 1979). F. Neobezzia fittkaui, male, in dorsal view (from Ronderos, et al. 2011). G. Nilobezzia virago, female, in dorsal view (from Elson-Harris 1987). H. Johannsenomyia argentata, male, in dorsal and ventral view (from Wirth & Grogan 1979). I. Johannsenomyia argentata, female, in dorsal and ventral view (from Wirth & Grogan 1979). J. Lanatomyia miles, female, in dorsal view (from Elson- Harris 1987). K. Anebomyia atripes, male, in dorsal and ventral view (from Wirth & Grogan 1979). L. Anebomyia atripes, female, in dorsal and ventral view (from Wirth & Grogan 1979).

FIGURE 79. Evolution of ecdysial sutures and splits in pupal exuviae of Ceratopogonidae, head and anterior portion of thorax in ventral view. Numbers refer to character states of characters 2 and 8 in text. Red indicates ecdysial splits. A. Leptoconopinae. B. Forcipomyiinae. C. Dasyheleinae. D. Most Ceratopogonini. E. Most Ceratopogonini, Clinohelea. F. Allohelea, Atyphohelea, Parabezzia. G. Heteromyia, Pellucidomyia, Hebetula. H. most Sphaeromiini s. lat., most Palpomyiini, Paryphoconus; some Bezzia are nearly completed fused (as in Fig. 79I), with only the dorsal apotome separate, along with an ecdysial split along the base of the antenna and the dorsal thoracic split. I. Jenkinshelea, Macropeza, Mallochohelea, Probezzia, Neobezzia.

FIGURE 81. Continuation of Majority rule (50%) cladogram of the genera of derived lineages of Ceratopogonidae. Numbers below each branch represent character state changes as described in the text and the matrix in Table 4. For characters with multiple states the character state changed to is indicated in superscript behind the character number. Bremer support values are shown above the branches. The root of this tree is a continuation from "All remaining Ceratopogonidae" as shown in Figure 80.

There are differences between the two genera in both male and female adults and in pupae, with most of the adult features not yet interpreted cladistically (but see character 85). They are summarized as follows: