Anisopteromalus calandrae (Howard)

Baur, Hannes, Kranz-Baltensperger, Yvonne, Cruaud, Astrid, Rasplus, Jean-Yves, Timokhov, Alexander V. & Gokhman, Vladimir E., 2014, Morphometric analysis and taxonomic revision of Anisopteromalus Ruschka (Hymenoptera: Chalcidoidea: Pteromalidae) - an integrative approach, Systematic Entomology 39 (4), pp. 1-19 : 9-13

Anisopteromalus calandrae (Howard)

( Figs 5D, F, G, I, K, 7 A, B, D)

Pteromalus calandrae Howard in Comstock, 1881: 273 : Neotype ♀ in NMBE, here designated, labelled ‘ USA Georgia Savannah Sitophilus oryzae leg. ix. 1978 [print]; chromosomes = 7 strain lab culture e.p. iii. 2002 Sitophilus granarius A. Timokhov, Moscow [print]; 1551 Baur [print]; Neotype ♀ Anisopteromalus calandrae Howard lab. H. Baur 2014 [hand, label with red left and right border]’ (entire; glued on a card rectangle); type locality: USA: Georgia, Savannah (examined by Baur). For discussion see under Remarks.

Pteromalus oryzae Cameron, 1891: 184–185: Syntypes ♂♂ and ♀♀ lost ( Bouček et al., 1979: 435), but see also Cotes (1896: 11); type locality: INDIA. Synonymized with A. calandrae by Bouček et al. (1979: 435).

Because the original description is inconclusive (Cameron mainly described some colour characters that apply to many Anisopteromalus species), we are inclined to accept the synonymy of Bouček et al. (1979). Moreover, the species described here as new can be ruled out: first, A. cornis sp.n. occurs in the Afrotropical region while P. oryzae was described from India; second, A. quinarius sp.n. naturally attacks anobiid beetles, while P. oryzae was recorded as a parasite of the rice weevil Sitophilus oryzae.

Meraporus vandinei Tucker, 1910: 343–344: Lectotype ♀ in USNM, catalogue no. 13389, here designated, labelled ‘Plano TX [print] 7. 27. 0 9 [hand]; emerged on [print] XI. 23. 0 9 [hand]; Hunter No [print] 1821 [hand]; ESTucker collector [print]; Paratype No. [print] 13389 [hand] U.S. N.M. [print]’ (entire; glued on a card point, with a card rectangle put underneath by Baur); type locality: USA: Texas, Plano (examined by Baur). Synonymized with A. calandrae by Mani (1938: 103), synonymy confirmed by MRA (see above).

Tucker (1910: 343) stated that his new species was based on 45 specimens collected by himself from Plano and by Van Dine from some other places. He then singled out one female and one male as ‘Type’ and considered the rest to be ‘Paratypes’. However, following the International Code of Zoological Nomenclature (ICZN) all specimens must be considered as syntypes. We received only 6♂ and 33♀ that stood under this name in the USNM. As lectotype we have taken a female with most body parts intact, and not the female with the additional label ‘Meraporus vandinei (n. sp.)

Tucker [hand]’ and that emerged on September 11, 1909. This specimen was heavily damaged and lacked the head and wings, but otherwise it was conspecific with the lectotype, as well as all the other paralectotypes we have seen.

Anisopteromalus mollis Ruschka, 1912: 243 –245: Lectotype ♀ in NHMV, here designated, labelled ‘2298 Mehlfruchtbörse [hand]; ex Laemophloeus ferrugineus [hand]; Anisopteromalus mollis m. type. female [hand]; Anisoptero = malus [sic] mollis [hand] det. Ruschka [print]; Type [print, red]’ (entire; lectotype the uppermost specimen on the pin; lectotype and one paralectotype ♀ remounted by Baur on card rectangles); type locality: AUSTRIA: Vienna (examined by Baur). Synonymized with A. calandrae by Graham (1969: 435), synonymy confirmed by MRA (see above).

Aplastomorpha pratti Crawford, 1913: 252–253: Lectotype ♀ in USNM, catalogue no. 15314, here designated, labelled ‘Dallas TX [print], Nov., 1906 [hand]; U.S. D. A. No. [print] 6076 [hand]; Bred from No [print] 3715 [hand]; WDHunter collector [print]; Paratype No. [print] 15314 [hand] U.S. N.M. [print]’ (right antenna beyond third anellus and parts of mid and hind legs lacking; glued on a card point, with a card rectangle put underneath by Baur); type locality: USA: Texas, Dallas (examined by Baur). Synonymized with A. calandrae by Mani (1938: 103), synonymy confirmed by MRA (see above).

We consider the entire series of 2♂ and 9♀ housed in the USNM to be syntypes, because no specimen was fixed as holotype in the original description (compare Crawford, 1913: 252–253). One female belonged to Dinarmus, as also stated by an identification label of Gahan. The rest of the specimens were clearly conspecific. We have taken as lectotype a female with most body parts intact, and not the female with the additional label ‘Aplastomorpha pratti Type Crfd [hand]’. This specimen was in a slightly worse condition, lacking for instance the metatibiae that constitute one of the characters used in our morphometric analyses. Neocatolaccus australiensis Girault, 1913: 306: Lectotype ♀ in QMB, catalogue no. 1956, here designated, labelled ‘ TYPE [print, orange label]; Syntype 1♂ 2♀♀ Hy. 1956 [hand, red label]; Neocatolaccus australiensis Gir., ♀ types [last word red underlined, Girault hand]’ (head lacking, wings glued to card point and crumpled; glued at tip of a card point; on the same card point two male paralectotypes, one with head missing. Mounted separately on a slide two female and two male heads labelled ‘ TYPE [print] Hy/1956 [hand] A. A. Girault [print]; Queensland Museum. [print] Neocatolac ♀ -cus. [sic] australiensis ♂ wing [hand]’; one of the heads might belong to the lectotype, the other one to a paralectotype female mounted on a separate pin; association of those heads with specimens was impossible); type locality: AUS- TRALIA: Queensland, Nelson (? = Gordonvale) (examined by Baur). Synonymized with A. calandrae by Bouček (1988: 414), synonymy confirmed by MRA (see above). As pointed out by Dahms (1983: 102), Girault (1913) published the catalogue number as ‘1596’ which is wrong. The correct number is ‘1956’ as indicated above.

Bruchobius medius Masi, 1917: 176–177: Holotype ♀ in BMNH, catalogue no. 5.699, labelled ’64 [hand, on card point]; Mahe, ’08-9 Seychelles Exp. [print]; Holo- [hand] Type [print; round label with red rim]; Percy Sladen Trust. Exped. B.M. 1913-170. [print]; Brucho = bius medius ♀ Masi [hand]; B.M. TYPE HYM. [print] 5.699; ♀ Anisopteromalus calandrae (How.) [hand] det. Z. Bouček, 197 [print] 5 syn. nov. [hand]’ (right flagellum beyond first segment and left hindwing lacking; card point mounted with micro-pin on rectangular tag; head remounted on card point by Baur); type locality: SEYCHELLES: Mahé, Cascade Estate (examined by Baur). Synonymized with A. calandrae by Bouček (1976: 22), synonymy confirmed by MRA (see above).

Neocatolaccus mamezophagus Ishii & Nagasawa, 1942: 67–68: Syntypes ♂♂ and ♀♀ in Tokyo College of Agriculture and Forestry; type locality: JAPAN. Synonymized with A. calandrae by Tachikawa (1966: 99).

The type material could not be traced, even with the help of Japanese colleagues. The original description clearly rules out that N. mamezophagus is the same as any of the new species described here. First, females of A. cornis sp.n. have the pedicel distinctly shorter than first funicular segment, while it is stated by Ishii & Nagasawa that the pedicel is slightly longer than the first funicular segment; second, males of A. quinarius sp.n. have the first funicular segment at least as long and usually longer than the second, while the first segment of N. mamezophagus is said to be slightly shorter than the second. The character states of N. mamezophagus match specimens of A. calandrae as defined here, hence we accept the synonymy of Tachikawa (1966).

Diagnosis, female. Head and mesosoma olive-green with slight bronze tinges in places, setae whitish, inconspicuous ( Fig. 5 K). Gena terete, not carinate near mouth margin. Flagellum distinctly clavate ( Fig. 5 I), first funicular segment subconical, basally slightly broader than third anellus, provided with 1–2 rows of longitudinal sensilla ( Fig. 5 F). Scutellum projecting at level of anterior margin of dorsellum ( Fig. 5 D), in lateral view weakly curved. Forewing with setae on wing disc dark. Speculum bare, closed in distal and proximal part. Anterior plica of propodeum short and evenly curved, joining sometimes an indistinct costula. Posterior margin of first gastral tergite curving backwards and strongly produced ( Fig. 5 G). Head breadth 1.24–1.51× metatibia length and 1.38–1.52× eye distance; head height 2.46–3.39× eye breadth; eye height 1.09–1.30× scutellum length; pedicel plus flagellum 1.77–2.25× eye height; mesosoma length 5.96–8.14× OOL; scutellum length 1.48–1.96× stigmal vein; metatibia length 1.66–2.21× marginal vein; gaster length 7.78–12.02× OOL.

Description, female. Antenna with scape testaceous, pedicel testaceous, fuscous on upper side, anelli fuscous and rest of flagellum testaceous, fuscous on upper side. Legs with tibiae yellowish.

Head 1.06–1.19× as broad as mesoscutum. POL 1.22–1.87× OOL. Eyes 1.42–1.80× as high as broad, separated by 1.29–1.59× their height. Malar space 0.46–0.63× eye height. Head in frontal view 1.10–1.30× as broad as high. Antenna with scape 0.77–1.00× as long as eye height. Combined length of pedicel plus flagellum 0.86–1.03× head breadth. First and second anellus strongly transverse, third transverse, about as long as second anellus.

Mesosoma 1.16–1.36× as long as broad. Mesoscutum 1.84–2.42× as broad as long. Hind margin of scutellum broadly rounded. Upper mesepimeron strongly narrowing below, reaching at most basal third of mesopleuron. Basal setal line complete. Costal cell with dorsal surface with at most a short row of setae distally, lower surface with a patch of setae in distal half and a single row of setae running to proximal part, costal setal line complete. Forewing disc moderately pilose. Marginal vein 1.35–1.96× as long as stigmal vein. Stigma subcircular to oval, medium-sized. Propodeum 0.42–0.60× as long as scutellum. Median carina fine, straight. Median area in anterior part evenly reticulate with inner corner of anterior plica with a rather deep and reticulate depression along the anterior plicae. Nucha subglobose, not distinctly separated from rest of propodeum, often at least with some traces of alutaceous sculpture.

Metasoma: Gaster 1.41–2.54× as long as broad, 1.13–1.66× as long as mesosoma, and 0.69–1.22× as long as mesoscutum. Posterior margin of second gastral tergite incised medially to almost straight. Posterior margin of third gastral tergite straight.

Material examined. Beside the above-mentioned namebearing types we examined the following specimens: Paralectotypes, Anisopteromalus mollis , 2♂, 1 ♀, same data as lectotype; Aplastomorpha pratti, USA: 2♂, 7 ♀, Texas, Dallas, xi.1906 (W.D. Hunter), paratype no. 15314 ( USNM); USA: 1 ♀, Texas, Dallas, xi.1906 (W.D. Hunter), type no. 15314 ( USNM); Meraporus vandinei, USA: 1♂, 1 ♀, Texas, El Campo, 22.vi.1909 (D.L. Van Dine), paratype ( USNM); USA: 3 ♂, 2 6♀, Texas, Plano, 26.vii.1909 ( E.S. Tucker), paratype no. 13389 ( USNM); USA: 1♂, 1♀, Texas, Plano, 26.vii.1909 ( E.S. Tucker), type no. 13389 ( USNM); USA: 2♀, Louisiana, Welsh, 2.viii.1909 (D. L. Van Dine), paratype ( USNM); USA: 1♂, 2♀, Louisiana, Lake Arthur, 29.vii.–2.viii.1909 (Rosenfeld and D.L. Van Dine), paratype ( USNM); Neocatolaccus australiensis, 2♂, 1♀, same data as lectotype. Further, non-type material is listed in Appendix S1.

Biology. The species has a relatively broad host range. It prefers to parasitize Sitophilus (Dryophthoridae) and Callosobruchus spp. (Chrysomelidae: Bruchinae); however, it can easily be reared (e.g. under laboratory conditions) on certain Anobiidae, like Lasioderma serricorne (see Gokhman & Timokhov, 2002; Timokhov & Gokhman, 2003). Many other host records (see Noyes, 2013) need to be verified.

Distribution. Cosmopolitan ( Noyes, 2013).

Remarks. The females of A. calandrae can be most easily recognized by the characters given in the key and diagnosis. The species is most similar to A. cornis sp.n. From A. quinarius sp.n. it is further distinguished by the pilosity of the forewing, for which we refer to the description of each species.

In order to assess the taxonomic status of A. calandrae , it is necessary to concentrate on the other cosmopolitan species, A. quinarius sp.n. Not only have the two species been confused over a long time, but they both are also likely to occur in human dwellings. However, these species usually occupy somewhat different habitats due to their different host preferences ( Gokhman & Timokhov, 2002; Timokhov & Gokhman, 2003): A. calandrae inhabits mills and grain bins where it is usually associated with Sitophilus spp.; A. quinarius sp.n. inhabits houses and warehouses (e.g. containing stored fruit or tobacco) being associated with Stegobium or Lasioderma beetles there. We believe that these differences were an important factor that hampered the discovery of A. quinarius sp.n. (see Discussion).

A further complication is that the male holotype of A. calandrae is now lost ( Peck, 1963: 733), which was recently confirmed by the curator of the Chalcidoidea collections at the USNM, M. Gates (personal communication). We have therefore studied characters of the males of the two species that were relevant with respect to Howard’s original description. They can be distinguished as follows:

• A. calandrae: First funicular (third flagellar) segment always shorter than the second ( Fig. 7 A), in small specimens only about as long as combined length of first and second anellus ( Fig. 7 B). Pale band in proximal half of gaster dirty yellow ( Fig. 7 D).

• A. quinarius sp.n.: First funicular (third flagellar) segment about as long as the second, always much longer than combined length of first and second anellus ( Fig. 7 C). Pale band in proximal half of gaster bright yellow ( Fig. 7 E).

Now, Howard in Comstock (1881: 273), who apparently examined a very small specimen, noted in his brief but concise description ‘joint 5 [corresponding to the third flagellar segment] small, equal in length to the two ring joints’ and ‘abdomen [corresponding to the gaster] yellow-brown at base’. A comparison of Howard’s statements with our description of the male characters should make it clear that both concepts of A. calandrae match almost perfectly.

With regard to the overall morphological similarity with the other cosmopolitan species, A. quinarius sp.n., we designated a neotype for A. calandrae in order to guarantee stability in the application of the name. This is even more important, because A. calandrae is the type species of Aplastomorpha Crawford, the only synonym of Anisopteromalus (see Noyes, 2013). We selected a specimen from Savannah ( Georgia), which might appear to be quite far away from Hempstead, Waller County (Texas), the place where the holotype of A. calandrae was collected (Howard in Comstock, 1881: 273). Given that the species is very easily disseminated from one site to another by transportation of stored products, the distance between the two localities is certainly not a problem here. Taking material of the laboratory culture from Savannah also had the advantage that a specimen in perfect condition coming from the centre of the analysed size and shape space was available (cf. Fig. 1 B). The Savannah strain could furthermore be included in our genetic analyses ( Fig. 4 ). Finally, in both cases the specimens were originally reared from Sitophilus oryzae -infested crop – that is, both demonstrably had the same host preference.

Figures

Fig. 4. ML trees from the analyses of ITS 2 and Cytb sequences of several strains of the OTUs Anisopteromalus calandrae and A. quinarius. Bootstrap supports higher than 70 % are indicated at nodes. Scale bar indicates substitution per site for both trees.