Anaphes Haliday 1833

Huber, John T. & Thuróczy, Csaba, 2018, Review of Anaphes Haliday (Hymenoptera: Mymaridae) with key to species in Europe and a world catalogue, Zootaxa 4376 (1), pp. 1-104 : 43-44

Anaphes Haliday 1833

Synonymy. Given in Huber (1992); the junior synonyms may also be determined from Tables 2, 3, and 6.

Type species. The type species is Anaphes fuscipennis Haliday ( ICZN 2017). Huber et al. (2011, 2014) had petitioned successfully to have the type species changed.

Diagnosis. Both sexes. Body almost always uniform black or dark brown, exceptionally with metasoma almost yellow; appendages usually lighter brown, often with off-white apices of femora, tibiae and tarsi except apical tarsomere ( Figs 45 , 56 ). Head. Face without subantennal grooves ( Fig. 3 ); malar sulcus absent, at most indicated exteriorly by slight change in sculpture ( Fig. 5 ), but visible internally in cleared slide mounts ( Figs 34, 35 , 50 , 52, 53 , 60 ); vertex without stemmaticum but with a groove from anterior margin of posterior ocellus to supraorbital trabecula ( Fig. 6 ); occiput with occipital groove [evidently an extension onto the occiput of the supraorbital suture of the vertex, incorrectly named the occipital suture in Huber (2006, fig. 32)] usually in line with supraorbital suture and extending ventrally to about level of lower margin of occipital foramen ( Fig. 4 ) but sometimes short and angled inwardly towards occipital foramen ( Figs 33 , 62 ). Mandible with 3 teeth, the dorsal tooth sometimes subdivided, appearing serrate, i.e. with 2 or 3 small apical teeth ( Figs 7, 8 ) but exceptionally (quinquearticulatus) with 2 large ventral teeth distinctly contrasting with 3 small, dorsal teeth ( Figs 24, 25 ); maxilla and labium as in Figs 26 and 27 . Wings. Wings fully developed, with posterior margin straight to slightly in basal half and slightly outcurved in apical half ( Figs 39 , 57 , 72 79 ) to distinctly ( Fig. 31 ) incurved, with apex slightly asymmetrical (anterior margin usually more curved than posterior margin), and with both dorsal and ventral surfaces almost always uniformly covered with microtrichia except for two more or less distinct bare areas (the marginal and medial spaces, small in Fig. 72 ) separated by an oblique row of microtrichia extending from just behind apex of stigmal vein towards wing apex ( Figs 39 , 57 ); wing retinaculum with a socketed seta near apex ( Fig. 39 ). Mesosoma. Anterior scutellum ( Figs 29, 30 , 54 , 63 ) without setae, with campaniform sensilla separated by about 2 or 3× their own diameter from each other and well separated from transscutal articulation, with fenestra a small triangle with rounded apices ( Fig. 43 , faintly indicated since below surface), and with frenum entire ( Figs 13 , 29 ); propodeum with a median longitudinal groove ( Figs 13 , 29 , 43 , 46 , 54 , 58, 59 , 63 ). Metasoma. Petiole a thin, narrow and almost vertical crescent ( Figs 40, 41 , 48 ) longitudinally divided medially ( Figs 46, 47 ); gaster with gt1 divided medially by a longitudinal groove ( Figs 17 , 29 , 43 , 46 ).

Female. Antenna. Clava 1- or 2-segmented, funicle 6-segmented ( Fig. 37 ) or, in one species, 5-segmented ( Fig. 28 ); when 6-segmented, fl1 usually the shortest segment ( Figs 9, 10 ). Genitalia. Ovipositor often projecting anteriorly under mesosoma, apparently enclosed in a somewhat membranous anterior extension or sac of gaster perhaps consisting mainly of gs6 ( Figs 15 , 18 , 56 , 67–70 ).

Male. Antenna. Flagellum distinctly 11-segmented only in species placed in Anaphes (Yungaburra) whereas in most nominal species flagellum appearing 10-segmented because fl1 minute and either with only 1 seta ( Figs 11, 12 , 38 ) or with a seta and 1 transverse mps ( Fig. 61 , Huber & Prinsloo 1990, figs 13, 14). Genitalia. Parameres thin, each almost as long as aedeagus and with a long seta more or less midway along paramere length ( Figs 22, 23 , 47 , 49 , 59 , 71 ).

Variation. A teratological specimen ( UCRC) from Greece with 4 funicle segments on one antenna and 5 on the other is known. Both antennae have fl1 + fl2 almost completely fused and one antenna also has fl4 + fl5 almost completely fused ( Fig. 103 ). All the tarsi are clearly 3-segmented ( Fig. 102 ). Otherwise the wings ( Fig. 104 ) and body are normal for an Anaphes specimen.

Subgeneric concepts. As mentioned above, Debauche (1948) proposed two subgenera, A. ( Anaphes ) and A. ( Patasson ), for females with a 1- or 2-segmented clava, respectively, and provided a discussion on how they differ in other, more subtle ways. Ogloblin (1962) proposed A. ( Austranaphes ) for species from Argentina with a 2- segmented clava and males with a clearly 11-segmented flagellum. Huber (1992) treated both Debauche subgenera as species groups within Anaphes sensu stricto and also proposed two subgenera, A. ( Anaphes ), and A. ( Yungaburra ) for most of the southern hemisphere species, with Ogloblin’s A. ( Austranaphes ) treated as one species group in A. ( Yungaburra ). A major problem with Huber’s classification is that the number of flagellomeres bearing mps in the male antenna must be known to classify a species into one or other subgenus. Males are so rarely associated with females that it is better to return to Debauche’s subgeneric classification and reclassify the morphologically diverse species that Huber classified in A. ( Yungaburra ) into one or other of Debauche’s subgenera. This, incidentally, would also be preferable for biological control workers, where Patasson , either as a genus or a subgenus, was used extensively prior to Huber (1992). Patasson should only be used as a subgenus of Anaphes , not a genus.

Alternatively, one could eliminate subgenera entirely and retain only species groups within Anaphes , reclassifying all the species previously placed in A. ( Yungaburra ) into them. If classified within the fuscipennis species group, for example, the species previously placed in A. ( Yungaburra ) could perhaps be distinguished from other species in the group using features such as size of the marginal and medial spaces (often considerably reduced, e.g., in nitens ), mps shape in females (often curved apically, e.g., in nitens ) and sculpture of the head and mesosoma (often with isodiametric reticulations, as in quinquearticulatus).

Finally, one could propose an entirely new species group classification but the problem would be to delimit them properly so they are easily identifiable. Huber’s subgenera cannot be distinguished based on the number of claval segments in females. Some undescribed Anaphes from Australia and one from Indonesia, as well as saintpierrei, have a 1-segmented clava in contrast to other, undescribed Australian region species and nitens , which have a 2-segmented clava. The former would key to the amplipennis group of Anaphes (Yungaburra) which includes the South American species only ( Huber 1992) and nitens obviously would key to his nitens group of A. ( Yungaburra ), in which he included only the Australian-region species. We emphasize again that the subgeneric classification of Anaphes proposed by Huber (1992) is therefore rejected in favour of a return to that of Debauche (1948), i.e., with the subgenera A. ( Anaphes ) and A. ( Patasson ). The species classified previously in A. ( Yungaburra ), are reclassified as follows: amplipennis, fijiensis, nunezi and pucarobius into A. ( Anaphes ), and galtoni, inexpectatus , nitens , and tasmaniae into A. ( Patasson ) ( Tables 3 and 6).

Figures

FIGURES58, 59. Anaphes inexpectatus, paratype male,same data as in Figs 50, 51. 58, metasoma, dorsal (slightly damaged); 59, mesosoma dorsal, genitalia as seen through mesosoma.Scale bars = 100 µm.

FIGURES71, 72. Anaphes nitens, male, same data as inFigs 60–62.71, metasoma, ventral,as seen dorsally through body; 72, wings. Scale bars = 200 µm.

FIGURES 73, 74. Anaphes spp., habitus and type slide (body length cited from original description). 73, arenbergi (0.699 mm); 74, brevicornis (0.594 mm).

FIGURES 75, 76. Anaphes spp., habitus and type slide (body length cited from original description). 75, crassipennis (0.82 mm); 76, ensipennis (0.72 mm).

FIGURE 77. Anaphes flavipes, habitus and neotype slide.Scale bar = 500 µm.

FIGURE 78. Anaphes fuscipennis, lectotype habitus (body length cited from original description = 0.025 inches [= 0.635 mm]).

FIGURE 79. Anaphes fuscipennis, habitus (specimen faded to brown), Canada, Nova Scotia, Sydney, 46°08.712'N 60°10.962'W, 14.vi.2004, sweeping fallow field,H.Goulet, C. Boudreault,A.Badiss.Scale bar = 500 µm.

FIGURES 44, 45. Anaphes brevis, male, same collection data as Figs 32–35. 44, mesosoma, lateral; 45, legs, lateraltarsomere lengths measured from one insertion to the next, as indicated by arrows.Scale bars = 100 µm.

FIGURES 56, 57. Anaphes inexpectatus, paratype female, same data as in Figs 50, 51. 56, habitus, lateral; 57, wings. Scale bars = 200 µm.

FIGURES 3–8. Anaphes sp., micrographs of head of unidentified specimens from Canada. 3, anterior; 4, posterior; 5, lateral; 6, dorsal; 7,ventral; 8, mouthparts.See Table 8 for morphological terms. Scale bars for 3–7 = 50 µm, for 8 = 10 µm.

FIGURES 32–35. Anaphes brevis, head, specimen from Spain, Catalonia, Navata, emerged 21.ix.2000 ex. Lygus eggs on Chenopodium (in cages 14–21.ix.2000), D. Coutinot & J. Lopez. 32, anterior; 33, posterior—arrow indicates vertexal suture (incorrectly named the occipital suture in Huber 2006, fig. 32); 34, slightly ventrolateral; 35, slightly dorsolateral. Scale bars = 100 µm.

FIGURES 50, 51. Anaphes inexpectatus, paratype female from Australia, Tasmania, near Hobart, ex. lab. culture in Pretoria, South Africa, i.1989, N.J. van Rensburg, ex.Gonipterus scutellatus [misidentification,probably G. platensis]. 50, head,lateral; 51, antenna.Scale bars = 100 µm.

FIGURES 52, 53. Anaphes inexpectatus, paratype male, same Data as in Figs 50, 51. 52, head + antennae; 53, head, posterior, seen through head from anterior. Scale bars = 100 µm.

FIGURES 60–62. Anaphes nitens, specimens from South Africa, Cape Province, Sassveld forestry Station near George, vii.1986, ex Gonipterus scutellatus. 60, female,head + antenna, anterior; 61, male antenna;62, male head,posterior.Scale bars = 200 µm.

FIGURES24–28. Anaphes quinquearticulatus, paratype female.24, head,anterior; 25, mouthparts, left mandible in focus; 26, mouthparts, maxillae in focus; 27, mouthparts,labium in focus;28, antenna.Scale bars for 24 and28 = 100 µm, for 25–27 = 50 µm.

FIGURES 36–39. Anaphes brevis, Same collection data as in Figs 32–35. (except Fig. 39—France, Yvelines, Béhoust, vi.1987, lab. culture ex. Lygus). 36, female, left antenna (outer surface); 37, female, right antenna (inner [medial] surface); 38, male, left antenna; 39, wings—arrow indicates socketed seta at apex of retinaculum.Scale bars = 100 µm.

FIGURES 29–31. Anaphes quinquearticulatus. 29, paratype female, mesosoma + metasoma, dorsal surface; 30, paratype female, mesosoma + metasoma, genitalia as seen throughbody; 31, holotype wings.Scale bars = 100 µm.

FIGURES 54, 55. Anaphes inexpectatus, paratype, same data as in Figs 50, 51. 54, male mesosoma, dorsal; 55, female mesosoma, lateral.Scale bars = 100 µm.

FIGURES 63, 64. Anaphes nitens, female, same Data as in Figs 60–62. 63, mesosoma, dorsal; 64, mesosoma, lateral. Scale bars = 200 µm.

FIGURES 42, 43. Anaphes brevis, same collection data as Figs 32–35. 42, female, metasoma, lateral, showing genitalia; 42, male, mesosoma, dorsal. Scale bars = 100 µm.

FIGURES 9–16. Anaphes sp., micrographs of unidentified specimens from Canada. 9, female, scape–base of fl 3, inner (medial) surface; 10, female, scape–base of fl 3, outer (lateral) surface; 11, male, scape–base of fl 4, inner (medial) surface; 12, male, scape–base of fl 4, outer (lateral) surface; 13, mesosoma, dorsal; 14, mesosoma, lateral; 15, mesosoma, ventral; 16, mesosoma, anterior.See Table 8 (p. 50) for morphological terms. Scale bars = 50 µm.

FIGURES 46, 47. Anaphes brevis, male, same collection data as Figs 32–35. 46, metasoma, dorsal; 47, genitalia, dorsal, as seen through metasoma.Scale bars = 100 µm.

FIGURES 40, 41. Anaphes brevis, female, same collection data as Figs 32–35. 40, mesosoma, lateral; 41, metasoma, lateral. Scale bars = 100 µm.

FIGURES 48, 49. Anaphes brevis, male, same collection data as Figs 32–35. 48, metasoma, lateral; 49, genitalia lateral, as seen through metasoma.Scale bars = 100 µm.

FIGURES 17–23. Anaphes sp., micrographs of unidentified specimens from Canada. 17, female, metasoma, dorsal; 18, female, metasoma, lateral; 19, female, metasoma, ventral; 20, male, apex of metasoma, dorsal; 21, male, genitalia, ventral; 22, male, apex of gaster, lateral; 23, male, apex of gaster, posterior.Scale bars = 50 µm.

FIGURES 67, 68. Anaphes nitens, female, same data as in Figs 60–62. 67, metasoma, lateral; 68, metasoma, midsagittal (= median) plane, showing internal structure of genitalia.Scale bars = 200 µm.

FIGURES69, 70. Anaphesnitens, female,same data as inFigs 60–62.69, metasoma,dorsal, genitalia as seen dorsallythrough body; 70, metasoma, ventral, as seen through body.Scale bars = 200 µm.

FIGURE 102–104. Anaphes sp., teratological specimen from Greece, Central Macedonia, Lake Kerkini 41°08'15.6"N 23°13'01.2"E, 65 m, 23–29.v.2006, G. Ramel (UCRC); 102, dorsal habitus, note apparently 3-segmented tarsi, enlarged in insets; 103, antennae, both with fl 1 + fl 2 fused and one (lower image) also with fl 4 + fl 5 fused; 104, wings. Fore wing length 585 µm, antennae length from apex of clava to base of pedicel 320 µm (4-segmented funicle) and 344 µm (5-segmented funicle). Body length ca. 0.58 mm.