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 : 44-46

Key to European species of Anaphes . Females

Note. Within Anaphes (Anaphes) , the species in couplets 11–14 are difficult to distinguish unequivocally. The features used to distinguish flavipes , medius , regulus , and stygius are weak and the synonymies proposed under each are not completely certain. There may be complexes of species within each of the three species or, alternatively, the synonyms proposed are indeed synonyms but under the wrong senior synonym. The authors are not in complete agreement with each other on which synonyms should be under which species and even whether stygius should be treated as separate from under regulus or not. Finally, we may even not have synonymized enough names under one or other of the species we consider valid. Synonyms are listed alphabetically in brackets. Characters such as presence or absence of cross striations on the inner (medial) surface of the scape and relative length of the clava compared to, in particular, fl5 + fl 6 may be useful features to separate some species but need further investigation, even though Soyka used them considerably in his descriptions and keys. Only one Anaphes (Anaphes) species, nipponicus, has been described from the eastern Palearctic region. It would key to flavipes, See Samkova et al. (2017) for morphological features to distinguish flavipes from nipponicus.

Within Anaphes (Patasson) , the last two species keyed, luna and silesicus , are very similar and difficult to distinguish from one another. We may have placed some synonyms under the wrong senior synonym. Alternatively, we may have overestimated the amount of infraspecific variation and therefore placed too many species in synonymy. No Anaphes (Patasson) species are described from the Palearctic region east of the Ural mountains.The key could be tried for that area but may not result in correct identifications. Anaphes inexpectatus and A. nitens , introduced from Australia to control Gonipterus spp. on Eucalyptus , are established in parts of southern Europe so are included in the key.

1 Clava 1-segmented (entire) [ Anaphes (Anaphes) ]......................................................... 2

- Clava 2-segmented (divided) [ Anaphes (Patasson) ]....................................................... 15

2(1) Fl2 and fl3 without mps; fore wing with posterior margin slightly outcurved (convex) or straight..................... 3

- Fl2 and fl3 with mps on one or both segments; fore wing with posterior margin slightly incurved (concave)............ 4

3(2) Fl4 without mps [fl5 and fl6 the widest segments]; ovipositor extending forward under mesosoma at most to about level of base of mesocoxa; marginal space about as long as parastigma + stigmal vein......................................................................................................... A. fuscipennis Haliday ( Figs 78 , 79 ) [= capitulatus, filicornis, maculatus, pratensis , neopratensis, stammeri]

- Fl4 with at least one mps; ovipositor extending forward under mesosoma to level of head; marginal space about as long as parastigma + stigmal vein.................................................... A. wolfsthali (Soyka) ( Fig. 87 )

4(2) Funicle with each segment shorter than pedicel length and each not more than 2.0× as long as wide.................. 5

- Funicle with at least one segment longer than pedicel and distinctly more than 2.0× as long as wide.................. 7

5(4) Fl2 with 2 mps; clava 1.6× as long as fl5 + fl6................................. A. quadraticornis (Soyka) ( Fig. 84 )

- Fl2 without mps; clava at most 1.5× as long as fl5 + fl6....................................................... 6

6(5) Fl4 with 2 mps; clava same length as fl5 + fl6.................................... A. brevicornis (Soyka) ( Fig. 74 )

- Fl4 without mps; clava 1.5× as long as fl5 + fl6................................. A. globosicornis (Soyka) ( Fig. 82 )

7(4) Fl4 without mps or (in ovipositor) with 1 mps on one or both antenna.......................................... 8

- Fl4 with 2 mps...................................................................................... 9

8(7) Fore wing 9× as long as wide, with longest marginal setae about 1.8× as long as widest wing width; microtrichia long, the apex of any one of them extending just past base of next one; metatarsomere 1 slightly longer than metatarsomere 2............................................................................... A. ensipennis (Soyka) ( Fig. 76 )

- Fore wing at most about 6.2× as long as wide, with longest marginal setae about 1.4× as long as widest wing width; microtrichia shorter, the apex of any one of them at most barely extending to base of next one; metatarsomere 1 slightly shorter than metatarsomere 2................................................... A. ovipositor Soyka ( Fig. 83 ) [= brevitarsis, germaniacus (i.e., angustipennis Soyka not Debauche), tenuipennis]

9(7) Metatarsomere 1 distinctly longer than 2...................................... A. crassipennis (Soyka) ( Fig. 75 ) [= acutipennis, aries , cultripennis, dilatatus, fennicus, ferrierei, gracillimus, laticornis , luteicornis , neoserenus, pilosipennis, serenus, siegerfeldi, sulphuripes, tarsalis, thoracicus, wertaneki]

- Metatarsomere 1 at most slightly longer than metatarsomere 2, but usually equal or slightly shorter than 2............ 10

10(9) Fl2 at most 2.4× as long as wide; fl1 + fl2 together not longer than fl3.................. A. gauthieri Debauche ( Fig. 80 ) [= avalae, breviventris, flavus, leonhardwitzi, nigricornis]

- Fl2 at least 2.5× as long as wide; fl1 + fl2 together longer than fl3.............................................. 11

11(10) Fore wing at most 4.4× as long as wide, with marginal setae less than 0.8× maximum wing width; marginal space separated from medial space by double line of microtrichia; hind wing fairly wide, apparently with two irregular rows of several microtrichia............................................................. A. arenbergi Debauche ( Fig. 73 )

- Fore wing at least 4.7× as long as wide, with marginal setae at least 0.9× maximum wing width; marginal space separated from medial space by a single line of microtrichia; hind wing narrower, with at most one irregular line of a few microtrichia................................................................................................. 12

12(11) Clava at most 3.2× as long as wide.................................................................... 13

- Clava at least 3.4× as long as wide.................................................................... 14

13(12) Ovipositor sheaths at least about 1.1× as long as metatibia............................. A. medius (Soyka) ( Fig. 81 ) [= intermedius, lacensis, pulchripennis, ranalteri, speciosus]

- Ovipositor sheaths at most about 1.0× as long as metatibia............................. A. regulus Walker ( Fig. 85 ) [= autumnalis, gracilior, soykai (i.e., crassicornis Soyka not Walker)]

14(12) Fore wing with apex narrower and slightly more pointed; clava at least 3.9× as long as wide.................................................................................................... A. flavipes Förster ( Fig. 77 ) [= antoniae, auripes, ciliatus, elegans, lemae , longispinosus, mirabilis, neospecialis, pilicornis, piliscapus, spinosus, varicolor]

- Fore wing with apex wider and slightly more rounded; clava at most 3.8× as long as wide. A. stygius Debauche ( Fig. 86 )

15(1) Back of head with occipital groove short, directed medially towards occipital foramen, thus forming an angle with supraorbital trabecula and strongly diverging away from posterior margin of eye ( Fig. 33 ); fore wing posterior margin with either a short hyaline section subapically, separating distal dark margin from proximal dark margin or entire posterior margin hyaline, contrasting with dark anterior margin.............................................................. 16

- Back of head with occipital groove longer, not directed medially towards occipital foramen, thus in line with supraorbital trabecula and weakly diverging from posterior margin of eye ( Figs 4 , 90c ); fore wing posterior margin almost always without a short hyaline section subapically (e.g. Figs 31 , 39 , 94 ), the entire posterior and apical margin uniformly dark..... 19

16(15) Marginal and medial spaces small, and medial space delimited posterobasally by at least 3 lines of microtrichia; fl2–fl6 with J- shaped mps (fig. 7 in Huber & Prinsloo 1990; Fig. 60 ); fore wing wide, with apex distinctly truncate (fig. 1 in Huber & Prinsloo 1990; Fig. 72 )..................................................... A. nitens (Girault) ( Figs 60–72 ) [= gonipteri]

- Marginal and medial spaces larger and more distinct, and medial space delimited posterobasally by 1 line of microtrichia; fl5–fl6 with straight mps (e.g., fig. 9 in Huber & Prinsloo 1990, and figs 43 and 46 in Huber 1992); fore wing narrower, with apex more evenly rounded (e.g., fig. 3 in Huber & Prinsloo 1990, and figs 27 and 28 in Huber 1992)................ 17

17(16) Ovipositor extending forward almost to head, trombone shaped; fl4 without mps (occasionally with 1 mps), the segment shorter and narrower than fl3 or fl5............................................... A. collinus Walker ( Fig. 90 ) [= aestivus, aterrimus, declinatus, discolor, discolorsimilis, latus, minor, nigerrimus, ornatus, ovatus, pallidus, pallipes, relictus, stubaiensis, tenuis , trijohanni ]

- Ovipositor extending forward at most to level of mesocoxa; fl4 with 2 mps (occasionally with 1 mps in inexpectatus or no mps in brevis , the segment usually as long and wide as fl3 or fl5, but sometimes shorter ( Figs 36, 37 )................. 18

18(17) Funicle segments longer, length/width of fl2–fl5 each at least 3.1...................... A. brevis Walker ( Figs 32–49 ) [= brunneus, latipterus, maculicornis, malchinensis, maradonae , pectoralis, pinguicornis, rotundipennis]

- Funicle segments shorter, Length/width of fl2–fl5 each at most 2.9...... A. inexpectatus Huber & Prinsloo ( Figs 50–59 )

19(15) Fl2–fl4 without mps; fore wing narrow and parallel sided, its width near apex equal to width at apex of venation............................................................................. A. angustipennis Debauche ( Fig. 88 )

- Fl2–fl4 with 1 or 2 mps on at least one of the segments, usually on two or three of them........................... 20

20(19) Fl2 with 1 or 2 mps................................................................................. 21

- Fl2 without mps................................................................................... 24

21(20) Fl2 with 1 mps.......................................................... A. debilipennis (Soyka) ( Fig. 93 ) [superadditus]

- Fl2 with 2 mps.................................................................................... 22

22(21) Fl6 at most 2.2× as long as wide................................................ A. maialis Debauche ( Fig. 99 ) [= aequus, devius, semiflavus, vulgarisimilis]

- Fl6 at least 3.0× as long as wide....................................................................... 23

23(22) Fore wing narrow, at least 6.0× as long as wide; ovipositor shorter than gaster, at most 0.8× as long as metatibia, not extending under mesosoma; metatarsomere 1 distinctly longer than metatarsomere 2 [scape narrow, with ventral margin not very convex; fore wing with marginal space long]................................... A. lineipennis (Soyka) ( Fig. 95 ) [= chrysomelae , dessarti, longipennis, obscurus, semimedius]

- Fore wing wider, at most 5.6× as long as wide; metatarsomere 1 at most only slightly longer than metatarsomere 2; ovipositor at least as long as gaster, at least 1.1× as long as metatibia........ A. longicornis Walker (part) ( Figs 96, 97 [male]) [= acutiventris , additus , archettii, ater, balteatus, campestris, communis, consimilis, distinctus, dubius, duplicatus, elongatus, eulongicornis , fabarius, falsus, flavitarsis, kressbachi, leptoceras , longior, malkwitzi, neodistinctus, neoflavus, pallidicornis, pannonicus, piceicornis, rufus, timidus]

24(20) Ovipositor not extending forward under mesosoma, shorter than almost globular and apically rounded gaster and less than 0.7× as long as metatibia; fl2 shorter than pedicel and shorter than fl3, at most 1.7× as long as fl1.................................................................................................. A. diana (Girault) ( Fig. 94 ) [= apilosus, compressus, globosus, heterotomus, lameerei , minimus, niger, parciventris, reductus, valkenburgicus]

- Ovipositor extending at least slightly forward under mesosoma and at least 1.1× as long as metatibia; fl2 almost always at least as long as pedicel, at least as long as fl3 and at least 2.0× as long as fl1.................................... 25

25(24) Metatarsomere 1 distinctly longer than metatarsomere 2................................................... 26

- Metatarsomere 1 shorter, equal to or, at most, barely longer than metatarsomere 2............................... 27

26(25) Fore wing at least 8.0× as long as wide.................................... A. parallelipennis (Soyka) ( Fig. 100 ) [= rectipennis]

- Fore wing at most 6.0× as long as wide.......................................... A. longicornis Walker (part) [gabitzi, medioacutus, sibbei]

27(25) Ovipositor at least 1.7× as long as metatibia length................................. A. crassus (Soyka) ( Fig. 89 ) [= stratipennis]

- Ovipositor at most 1.6× as long as metatibia length....................................................... 28

28(27) Fore wing at least 9.0× as long as wide......................................... A. crassipilis (Soyka) ( Fig. 92 ) [linearis]

- Fore wing at most 7.6× as long as wide................................................................. 29

29(28) Fore wing with microtrichia on dorsal surface of membrane long, the apex of microtrichia usually overlapping the base of the next............................................................... A. crassicornis (Walker) ( Fig. 91 ) [arcuatus, fortipennis, longipilis, nemorosus, lucidus, serbicus, variatus, weidenhofi]

- Fore wing with microtrichia on dorsal surface of membrane shorter, the apex of each not usually extending to base of the next............................................................................................. 30

30(29) Metatarsomere 1 as long as metatarsomere 2; fl2 usually as long as fl3..................... A. luna (Girault) ( Fig. 98 ) [aequipennatus, albipes, bicolor , brachygaster , devillei , differens, dorcas , euryale , hundsheimensis, neobrevior, neustadti, ordinarius, palliditarsis, pilosissimus, speciosior, swiedecki, vulgaris, wratislawensis]

- Metatarsomere 1 shorter than metatarsomere 2; fl2 usually shorter than fl3.............. A. silesicus (Soyka) ( Fig. 101 ) [brevior, calvescens , exiguus, exiguosimilis, flavicornis, neobrevior, obsoletus, pellucens, similis]

Figures

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.

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.

FIGURE 82. Anaphes globosicornis, habitus. 82a, body without head; 82b, head + antennae and type slide. Body length cited from original description = 0.63 mm.

FIGURES83, 84. Anaphes spp., habitus and typeslide (body length cited from originaldescription).83, ovipositor (0.39mm); 84, quadraticornis (0.60 mm).

FIGURE 85. Anaphes regulus, lectotype, habitus (0.47 mm—mesosoma + metasoma only, cited from Graham [1982]).

FIGURE 90. Anaphes collinus, lectotype. 90a, habitus (0.4 mm, cited from Graham’s [1982] description); 90b, antenna; 90c, wings.

FIGURE 91. Anaphes crassicornis, lectotype, habitus (= 0.4 mm, cited from Graham’s [1982] description). Scale bar = 500 µm.

FIGURES92, 93. Anaphes spp., habitus and typeslide(body length cited from original description).92, crassipilis (0.36 mm); 93, debilipennis(0.54 mm).

FIGURES 86, 87. Anaphes spp., habitus and type slide (body length cited from original description). 86, stygius (0.706 mm); 87, wolfsthali (0.75 mm).

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).

FIGURES 80, 81. Anaphes spp., habitus and type slide (body length cited from original description). 80, gauthieri (0.473 mm); 81, medius (0.83 mm).

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 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 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 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 94, 95. Anaphes spp., habitus and type slide. 94, diana; 95, lineipennis (0.63 mm, body length cited from original description).Scale bar for 94= 500 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.

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

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.

FIGURES 88, 89. Anaphes spp., habitus and type slide (body length cited from original description).88, angustipennis (0.431 mm); 89, crassus (0.63 mm).

FIGURES 98, 99. Anaphes spp., habitus and type slide (body length cited from original description). 98, luna; 99, maialis (0.595mm). Scale bar for 98 = 500 µm.

FIGURES 96, 97. Anaphes longicornis, habitus. 96a, lectotype (0.75 mm, cited from Graham’s (1982) description); 96b, lectotype, antenna; 97, paralectotype, male.

FIGURES 100, 101. Anaphes spp., habitus and type slide (body length cited from original description). 100, parallelipennis (0.612mm); 101, silesicus (0.43 mm).