Anagrus (Anagrus) bakkendorfi Soyka, 1946

(Figs 1–13)

Anagrus armatus var. nigriceps Girault 1915: 276 . Type locality: Corvallis, Benton County, Oregon, USA. Lectotype female [USNM], designated by Chiappini et al. (1996: 578), on slide; examined (Chiappini et al. 1996: 578).

Anagrus bakkendorfi Soyka 1946: 40 . Type locality: Valkenburg, Limburg, the Netherlands. Oldest available replacement name for A. nigriceps Girault 1915: 276 ( A. armatus var. nigriceps) nec A. nigriceps (Smits van Burgst 1914: 125–127) ( Litus nigriceps). Holotype female [NHMW] on W. Soyka’s slide #340; examined (Chiappini & Triapitsyn 1999: 121).

Anagrus latipennis Soyka 1956: 24 . Type locality (of the lectotype female on W. Soyka’s slide # 417 in NHMW, effectively designated by Chiappini 1989: 107 [an invalid designation of a holotype]): Jettchenshof [as “Jettchens Hof”; a farm adjacent to the woods, ca. 1 km E of Pisede, ca. 53°46’N 12°46’E, 12 m, formerly in Landkreis Demmin], Malchin, Mecklenburgische Seenplatte, Mecklenburg-Western Pomerania, Germany; examined (Chiappini & Triapitsyn 1999: 121). Synonymy with A. bakkendorfi by Chiappini 1989: 106–107.

Anagrus avalae Soyka 1956: 24 . Type locality (of the lectotype female on W. Soyka’s slide # 338 in NHMW, effectively designated by Chiappini 1989: 108 [an invalid designation of a holotype]): Mt. Avala, Belgrade, Serbia; Chiappini & Triapitsyn, 1999: 120–124 (then oldest available replacement name for A. nigriceps Girault 1915: 276 ( A. armatus var. nigriceps) nec Anagrus nigriceps (Smits van Burgst 1914: 125–127) ( Litus nigriceps); examined (Chiappini & Triapitsyn 1999: 120). Syn. n.

Anagrus arcuatus Soyka 1956: 24 . Type locality (of the lectotype female on W. Soyka’s slide # 335 in NHMW, effectively designated by Chiappini 1989: 108 [an invalid designation of a holotype]): Europe (possibly an unspecified locality in Burgenland, Austria because of an abbreviation “Bgld.” on the original label); examined (Chiappini & Triapitsyn 1999: 121). Synonymy with A. avalae by Chiappini & Triapitsyn 1999: 120–124. Syn. n.

Anagrus valkenburgensis Soyka 1956: 24 . Type locality (of the lectotype female on W. Soyka’s slide # 494 in NHMW, effectively designated by Chiappini 1989: 107 [an invalid designation of a holotype]): Valkenburg, Limburg, the Netherlands; examined (Chiappini & Triapitsyn 1999: 121). Synonymy with A. avalae by Chiappini & Triapitsyn 1999: 120–124. Syn. n.

Anagrus diversicornis Soyka 1956: 24 . Type locality (of the lectotype female on W. Soyka’s slide # 362 in NHMW, effectively designated by Chiappini 1989: 108 [an invalid designation of a holotype]): Valkenburg, Limburg, the Netherlands; examined (Chiappini & Triapitsyn 1999: 121). Synonymy with A. avalae by Chiappini 1989: 108. Syn. n.

Anagrus incarnatus ssp. fuscus Boţoc 1963: 99, figs 5a–5d. Type locality (of the lost type (s), not specified): Cluj-Napoca, Cluj, Romania; not examined. Synonymy with A. bakkendorfi by Triapitsyn & Berezovskiy 2004: 29.

Anagrus nigriceps: Burks 1979: 1023 (catalog).

Anagrus (Anagrus) bakkendorfi: Chiappini 1989:106–107 (synonymy,redescription, type information); Triapitsyn & Berezovskiy 2004: 29–30 (synonymy, distribution, comments); Triapitsyn 2015: 12 (key), 27 (coded redescription, distribution, hosts), 42 (checklist, synonyms); Triapitsyn et al. 2020b: 568 (distribution).

Anagrus (Anagrus) avalae: Chiappini 1989: 108 (synonymy, redescription, type information); Triapitsyn 2001: 282–284 (taxonomic history, identification, distribution and hosts in Australia and New Zealand); Triapitsyn & Berezovskiy 2004: 24–25 (distribution, host associations); Triapitsyn 2015: 12 (key), 27 (coded redescription, distribution, hosts), 41 (checklist, synonyms); Triapitsyn et al. 2019: 89 (molecular voucher), 94, 96 (genetic analysis); Triapitsyn et al. 2020a: 141 (host in Japan); Triapitsyn et al. 2020b: 568 (distribution).

Anagrus oregonensis Triapitsyn in Chiappini et al. 1996: 578–579, then replacement name for A. nigriceps Girault 1915: 276 ( A. armatus var. nigriceps) nec Anagrus nigriceps (Smits van Burgst 1914: 125–127) ( Litus nigriceps); synonymy with A. avalae by Chiappini & Triapitsyn 1999: 120–124. Syn. n.

Anagrus avalae: Chiappini & Triapitsyn 1999: 120–124 (taxonomic history, synonymy, type information, distribution, redescription of both sexes, illustrations, diagnosis, host associations).

Anagrus bakkendorfi: Chiappini & Triapitsyn 1999: 121 (type information including for A. latipennis), 124 (diagnosis, comments).

Material examined. NEARCTIC. CANADA. BRITISH COLUMBIA: Oliver, Eggert vineyard (Fairview Cellars winery), collected 5.ii.1999, emerged in Summerland 1.iii.1999, T. Lowery, K. Todd, from leafhopper eggs on wild rose [1 ♀ (Fig. 3), 1 ³ (Fig. 4), UCRC] (identified as A. avalae by S. V. Triapitsyn in 1999). Summerland, PARC Entomological Orchard, collected 1.vii.1999, emerged in 28.vii.1999, T. Lowery, K. Todd, from leafhopper eggs on dogwood [1 ♀, UCRC] (identified as A. avalae by S. V. Triapitsyn in 1999). USA. CALIFORNIA: Alameda County, Albany, Gill Tract, 37°53’09’’N 122°17’58’’W, 10 m, S.H. Wilson, collected 28.viii.2013, emerged 30.viii.2013 from unknown host eggs on an unknown tree [1 ♀, UCRC] (identified as A. avalae by S. V. Triapitsyn in 2014). Mendocino County, Fetzer, 13244 Old River Road, 38°59’22’’N 123°06’15’’W, 153 m, S.H. Wilson, collected 30.viii.2013, emerged 8.ix.2013 from unknown host eggs on alder, Alnus sp. [1 ♀, UCRC] (identified as A. avalae by S. V. Triapitsyn in 2014). Sonoma County, Simi Chalk Hill, 38°38’05’’N 122°45’51’’W, 55 m, S.H. Wilson, from unknown host eggs on alder: collected 7.v.2012, emerged 8.v.2012 [1 ♀, UCRC] (identified as A. avalae by S. V. Triapitsyn in 2013); collected 3.xi.2013, emerged 9.xi.2013 [1 ♀, UCRC] (identified as A. avalae by S. V. Triapitsyn in 2015).

PALAEARCTIC. RUSSIA. MOSKOVSKAYA OBLAST’, Pushkinskiy rayon, Pushkino, Mamontovka, E.Ya. Shuvakhina, Malaise trap in garden: 10–20.vii.2000 [1 ♀, UCRC (Fig. 5)]; 20–31.vii.2000 [1 ♀, UCRC] (identified as A. avalae by S. V. Triapitsyn in 2000 and 2002, respectively). PRIMORSKIY KRAY, Ussuriyskiy rayon, Gornotayozhnoye, 43.66°N 132.25°E, 200 m, 21–31.vii.2000, M. V. Michailovskaya, Malaise trap [1 ♀, UCRC] (identified as A. avalae by S. V. Triapitsyn in 2002). SPAIN. NAVARRA, Irati Forest, Iratibizkar, E. Baquero: 1120 m (Malaise trap in Fagus sylvatica forest): 26.vii.2000 [1 ♀, MZNA], 10.viii.2000 [1 ♀, MZNA], 23.viii.2000 [11 ♀, MZNA; 5 ♀, UCRC (Fig. 1)], 7.ix.2000 [1 ³, UCRC (Fig. 2)] (identified as A. bakkendorfi by S. V. Triapitsyn in 2004 except for 6 females in MZNA, collected 23.viii.2000, identified by S. V. Triapitsyn in 2004 as A. avalae); 42.970669°N 1.114398°W, 1154 m, 4.ix.2020, E. Baquero, yellow pan traps [3 ♀, MZNA (2, including molecular voucher PR20-511, UCRC _ ENT 00541253), UCRC (1, molecular voucher PR20-512, UCRC _ ENT 00541252)] (identified as A. bakkendorfi by E. Baquero and S. V. Triapitsyn in 2020). UNITED KINGDOM. ENGLAND: Kent County, Sevenoaks, viii.2014, A. Polaszek, yellow pan traps in garden [1 ♀, UCRC, molecular voucher PR15-026, UCRC ENT 311795] (identified as A. avalae by S. V. Triapitsyn in 2015). Surrey County, Woking, 25.vi.1885, F. Enock [1 ♀, MMUE] (on F. Enock’s slide #1332, labeled with his manuscript name “ Anagrus molsoni ”) (identified as A. avalae by S. V. Triapitsyn in 2014).

Updated diagnosis. FEMALE. Body color variable, from yellow or light brown except a darker anterior part of mesoscutum, to brown or dark brown except for pale frenum (Figs 1, 3, 5). Antennal funicle with multiporous plate sensilla on F3 (0 or 1) (absent from examined specimens with longer ovipositors that key to A. bakkendorfi in Triapitsyn 2015, Figs 7, 8), F4 (1), F5 (0 (Fig. 8) or 1 (Fig. 7)), F6 (1 or 2). Fore wing (Fig. 9) with 2 rows of setae behind and just beyond apex of venation. Ovipositor (Figs 6, 10–13) of variable length and extent of projection beyond gastral apex, 1.9–3.3× as long as protibia. Otherwise as redescribed by Chiappini & Triapitsyn (1999) for A. avalae and by Triapitsyn (2015) for both A. avalae and A. bakkendorfi .

MALE. As described and illustrated for A. avalae by Chiappini & Triapitsyn (1999). Body often notably darker (Figs 2, 4) than for some lighter colored females (Figs 1, 3).

Distribution. PALAEARCTIC (? Austria, Belgium, Bulgaria, Finland, France, Germany, Greece, Iran, Italy, Japan, Netherlands, Poland, Romania, Russia, Serbia, Spain, Sweden, United Kingdom), AUSTRALASIAN (Australia, New Zealand), NEARCTIC (Canada, USA) (Triapitsyn 2015; Triapitsyn et al. 2020b [also as A. avalae]), and possibly NEOTROPICAL (Chile), according to an unconfirmed record of Anagrus armatus nigriceps Girault by Guilleminot & Apablaza (1986), although this record far more likely results from a misidentification.

Almost certainly unintentionally introduced from Europe to Australia and New Zealand (Triapitsyn 2001 [as A. avalae]). It is unclear if its occurrence in the Nearctic region is due to its apparent natural Holarctic distribution or an accidental introduction from Europe; we would guess the latter scenario is more likely because in western North America this species is almost exclusively collected in agricultural and garden environments, which have not been well sampled in the eastern Nearctic region.

Hosts. Various Cicadellidae (Hemiptera) listed for A. avalae by Chiappini & Triapitsyn (1999), Triapitsyn (2001), Triapitsyn & Berezovskiy (2004), and Triapitsyn et al. (2020a).

Remarks. As expected, based on the morphological predictive assessment (Chiappini & Triapitsyn 1999; Triapitsyn & Berezovskiy 2004), specimens PR20-511 and PR20-512 identified as A. bakkendorfi, in which the ovipositor is 3.0× and 3.3× the length of the protibia, respectively, were found to be genetically very similar to the specimen PR15-026 identified as A. avalae, in which the ovipositor is 2.1× the length of the protibia. Sequences of the nuclear ribosomal ITS2 gene from specimens PR20-511 and PR20-512 (GenBank accessions MW633070 and MW633071, respectively) were the same length (+/- 1bp) as three cloned sequences previously obtained from PR15-026 (MK024915 –17), and were 98.7–99.6% similar, with no consistent substitutions present. Those of the mitochondrial COI gene (MW633277 and MW633278) bore 95.9 % similarity with the single COI sequence from PR15-026 (MK024808). Taken together, these levels of similarity are a strong indication of conspecificity, hence the here proposed synonymy of A. avalae (and its synonyms) with A. bakkendorfi . This corroborates the recent finding that in females of some Anagrus Haliday species, such as A. (Anagrus) atomus (L.), intraspecific variability of the relative length of the ovipositor can be far more pronounced than previously recognized (Triapitsyn et al. 2020c), so by itself it should not be used as the sole species defining feature.