Achrysocharoides Girault (Hymenoptera, Eulophidae) new to tropical America, with eight new species

Abstract The genus Achrysocharoides Girault is here reported for the first time from tropical America. Included are ten species, eight newly described: Achrysocharoides asperulus, Achrysocharoides callisetosus, Achrysocharoides cuspidatus, Achrysocharoides foveatus, Achrysocharoides infuscus, Achrysocharoides mediocarinatus, Achrysocharoides purpureus, Achrysocharoides sulcatus, and two already known: Achrysocharoides ecuadorensis (Hansson) and Achrysocharoides gliricidiae (Hansson & Cave). All species are included in an identification key, diagnosed, described and illustrated. Only one of the species, Achrysocharoides gliricidiae, has a host record, an endoparasitoid in a leafmining Gracillariidae (Lepidoptera) on Gliricidia sepium (Fabaceae), thus conforming to the biology of extralimital Achrysocharoides species. The genus Kratoysma Bouček is here established as a junior synonym of Achrysocharoides, and the following species previously in Kratoysma are here recombined to Achrysocharoides: Kratoysma citri Bouček, Kratoysma ecuadorensis Hansson, Kratoysma gliricidiae Hansson & Cave, Kratoysma longifacies Hansson, Kratoysma nepalensis Hansson, Derostenus usticrus Erdös.


Introduction
Species of Achrysocharoides Girault are unusually host/host plant specific (e.g. Askew and Ruse 1974). They develop as koinobiont endoparasitoids on larvae of leafmining microlepidoptera, mainly on species in the genus Phyllonorycter Hübner (Lepido-ptera: Gracillariidae) (e.g. Bryan 1980a). They have species-specific, and diverse, sex allocation strategies (West et al. 1999), courtship behaviour (Bryan 1980b), and sexspecific size differences (Askew and Ruse 1974). Furthermore, Hansson et al. (ms submitted) demonstrated that species of Achrysocharoides co-occurring simultaneously on the same plant, i.e. sympatric species, showed reproductive character displacement of newly discovered visual signaling attributes, i.e. strongly indicating them as devices for reproductive isolation. These visual signals featured wing interference patterns (WIPs), a new type of character recently discovered by . Thus Achrysocharoides possesses physical attributes and behaviours that make it very interesting as a model group for evolutionary studies.
Knowledge of Achrysocharoides is almost exclusively confined to the temperate parts of the Northern Hemisphere , and references therein), with 22 species known from Europe (now 23 with the recombined A. usticrus (Erdös)), 22 from North America and 10 from Japan. At least nominally no species were previously recorded from the Neotropical region. However, with the synonymization of Kratoysma with Achrysocharoides, in this paper, two species are actually known from tropical America. One of these species is with a host record and similar to northern temperate species it is an endoparasitoid in leafmining Gracillariidae. With the addition of the eight new species described here from tropical America the basic knowledge of this biologically interesting group has increased considerably, opening up the possibility of further research of the evolution of this group in a tropical area. The ratios, summarized in Table 1, are based on the holotype and one of the paratypes (if present) of the other sex.

Results and discussion
The analysis, based on external morphological characters, of the material at hand resulted in ten species -two already described and eight undescribed species. Wing interference patterns (WIPs) was one character-set included in the analysis. However, these patterns had no useful information for species separation as they were very similar between species (as in Fig. 26). Hansson et al. (manuscript, submitted) called this pattern ancestral and it was present in all allopatric species of Achrysocharoides in Europe. This pattern is with a narrow band in the forewing, from the stigmal vein to the posterior margin of the wing, with a thick membrane inside and a thin membrane outside the band. Applying the results from Hansson et al., the prediction is that all species included here are allopatric.
Nine of the species included here belong to the gahani species-group, characterized by having an edge along occipital margin, a transverse carina close to posterior margin Table 1. Ratios, for an explanation of the morphological abbreviations see above. of dorsal pronotum, two submedian carinae on propodeum, and with a row of foveae laterally on scutellum. Achrysocharoides gliricidiae lacks propodeal median carinae (Fig.  31), and A. mediocarinatus has just a single median carina (Fig. 42). However, both species have the other characters for the group, and they are thus best placed in the gahani-group. The placement of A. foveatus into a species-group is problematic, and it is left as unplaced. This species has a sharp edge along the occipital margin (Fig. 56) and foveae on lateral part of scutellum (Fig. 59), but lacks a pronotal carina and longitudinal carinae on propodeum (Fig. 57). Apart from the gahani-group three other species-groups may have pits on the scutellum (Kamijo 1991): titiani-, clypeatus-, and latreilleii-groups, but none of these groups have a carinate occipital margin, submedian carinae on propodeum, or -except some species in the titiani-group -a transverse carina on pronotum. When describing Kratoysma Bouček (1965) compared it with subgenus Kratochviliana Malač (of genus Chrysocharis Förster), with Enaysma Delucchi (now a synonym of Achrysocharoides) -hence the name Krato(chviliana)-(Ena)ysma -and with Pediobius Walker. He found the "general aspect" of Kratoysma to be like that of Kratochviliana, the frons with a raised cross-line (= raised frons above frontal suture) was shared with Enaysma, while two other characters, wing venation and presence of lateral plicae on the propodeum, were shared with Pediobius. Some of the features mentioned by Bouček are hard to define: "general aspect" and unspecified "wing venation" are open to any interpretation. The raised frontal suture and the presence of propodeal plicae are more definite features. The raised frons above frontal suture (in the female) is an apomorphy also for Achrysocharoides, while the propodeal plicae are present in several other genera of Entedoninae, e.g. Pediobius and in some species of Achrysocharoides. Achrysocharoides has another apomorphy unknown to Bouček at the time of the description of Kratoysma: a longitudinal carina on the lateral downsloping part of pronotum (Gumovsky 2007) (Figs 60-63). This longitudinal carina is also present in Kratoysma usticrus, the type species for Kratoysma (Fig. 61). Hence there are two unique apomorphies for Achrysocharoides+Kratoysma, the raised frontal suture and the longitudinal carina on lateral pronotum. The differences between these genera at the time when Kratoysma was described were mainly two: presence in Kratoysma of a transverse pronotal carina and propodeal plicae, with the absence of both in Achrysocharoides. However, the addition of new species of Achrysocharoides from Japan (Kamijo 1990a(Kamijo , 1990b and North America (Kamijo 1991) has demonstrated a variation in both characters, a variation that obscures the borderline, based on morphological characters, between these two genera and which presents major difficulties for the definition of them. Hence there are no longer any derived characters to keep Achrysocharoides and Kratoysma as separate genera, and Kratoysma, being the junior name, is hereby synonymized with Achrysocharoides. The following species, previously placed in Kratoysma, are hereby transferred to Achrysocharoides: A. citri (Bouček), comb. n., A. ecuadorensis (Hansson), comb. n., A. gliricidiae (Hansson & Cave), comb. n., A. longifacies (Hansson), comb. n., A. nepalensis (Hansson), comb. n., A. usticrus (Erdös), comb. n.
The classification into species-groups. The subdivision of Achrysocharoides was initiated by Graham (1959) who divided the European species into two subgenera, Enaysma Delucchi and Pentenaysma Graham. These correspond with the two speciesgroups, atys-and latreilleii-groups, which Bryan (1980a) introduced for the European species, thus abandoning the formal subdivision into subgenera. Yoshimoto (1977) divided the Nearctic species into two species-groups, the gahani-and guizoti-groups. Kamijo (1991) transferred some of the Nearctic species from the guizoti-group to either of the two newly erected clypeatus-and titiani-groups, and removed the remaining species in the guizoti-group to the latreilleii-group, thus terminating the guizoti-group. Kamijo (1990b) established the crassinervis-group for two species from Japan and one undescribed species from Nepal. Hence there are currently six species-groups in Achrysocharoides: atys-, clypeatus-, crassinervis-, gahani-, latreilleii-, and titiani-groups. See Kamijo (1991) for group-diagnostic characters. Diagnosis. Eyes densely pubescent (e.g. Fig. 3); females with frontal suture as a raised carina (i.e. with frons just above frontal suture protruding), straight (e.g. Fig.  3); males with frontal suture straight to slightly V-shaped, sometimes missing; females with antennal scrobes indistinct (i.e. not as narrow grooves) joining below frontal suture (e.g. Fig. 3); lateral downsloping part of pronotum with a longitudinal carina (Figs 60-63); postmarginal vein short, 0.5-1.5× as long as stigmal vein. Achrysocharoides is similar to Apleurotropis, but has a short postmarginal vein (in Apleurotropis postmarginal vein is 2.8-3.7× as long as the stigmal vein), with antennal scrobes in female joining below frontal suture (antennal scrobes join the frontal suture separately in Apleurotropis).
Identification. To separate Achrysocharoides from other Eulophidae genera the keys in Bouček (1988) Gibson et al. (1997) (Nearctic), Graham (1959) (Europe) are useful. To differentiate Achrysocharoides from other genera of Entedoninae in tropical America the matrix key on the website http://www.neotropicaleulophidae.com can be used.
Description. Female flagellum with a 2-segmented clava, in male with a 2-segmented clava or with all 5 flagellomeres distinctly separated; male flagellomeres with scattered setae; male scape enlarged, frequently with a species-specific shape, ventral sensory area present along entire scape; sensilla ampullacea globular, symmetric (type I sensu Hansson (1990)), present on all flagellomeres. Antenna with discoid anelli. Mandibles with two large teeth at apex, with one or several smaller teeth above large teeth. Clypeus not delimited. Malar sulcus present. Males with a more or less developed cross-ridge below antennal toruli. Frons occasionally with an indistinct groove between median ocellus and frontal suture. Female with frons just above frontal suture protruding, hence frontal suture appear to be a raised carina. Frontal suture in female straight; in male straight to slightly V-shaped, but sometimes missing. Antennal scrobes usually join below frontal suture in females, join on or below frontal suture in males, scrobes absent in males of some species. Occipital margin with raised carina or an edge, or rounded; occiput with a median fold/groove, at least close to occipital margin.
Pronotum with or without a transverse carina. Midlobe of mesoscutum with two pairs of setae, sometimes with an indistinct median groove in posterior ½; notauli more or less distinct in anterior 1/2, in posterior 1/2 present as weakly delimited depressions which are smooth to weakly reticulate. Scutellum with one pair of setae; sometimes with an anteromedian groove; with or without detached lateral foveae or rows of foveae. Transepimeral sulcus almost straight to weakly curved. Dorsellum visible in dorsal view. Forewing with costal cell usually wider than width of base of submarginal vein; postmarginal vein 0.5-1.5× as long as stigmal vein, usually about as long as stigmal vein. Propodeum without longitudinal ridges, or with a complete median carina -undivided or branched in posterior half -median carina sometimes incomplete and present only in posterior 1/3, or with two complete submedian carinae that run parallel or diverge weakly to strongly towards posterior part of propodeum.
Petiole 0.5-1.5× as long as wide, smooth and shiny or with some irregular sculpture, sometimes with anterolateral corners, ventral surface smooth. Male genitalia as in most other genera of Entedoninae, i.e. with normal volsellar setae, one parameral setae at the apex of phallobase, with two digital spines (Hansson 1996).
Petiole 0.6× as long as wide, with weak sculpture. Gaster oval-shaped. MALE. Unknown. Etymology. From the Latin asper = rough, in its diminutive form = asperulus, referring to the strong reticulation on thoracic dorsum.
Petiole 1.5× as long as wide, dorsal surface with weak sculpture. Gaster oval-shaped. MALE. Unknown. Etymology. From "propodeal callus" and the Latin setosus = bristly, referring to the numerous setae on propodeal callus.
Distribution. Costa Rica. The two available specimens have both been collected at high altitude (2600-2800 m).
Petiole as long as wide, dorsal surface smooth. Gaster slightly elongate. MALE. Unknown. Etymology. From the Latin cuspidatus = make pointed, referring to reticulate part on midlobe of mesoscutum that is pointed towards scutellum.
Petiole as long as wide, dorsal surface with weak irregular sulpture. Gaster slightly elongate.
Description. FEMALE. Length 1.2-1.8 mm. Scape yellowish-brown to pale brown, remaining antenna dark brown. Frons below frontal suture golden-green to golden-red, above metallic bluish-purple to goldengreen (Fig. 53). Vertex metallic bluish-purple to golden-green. Mesoscutum, scutellum and propodeum golden-green to metallic bluish-green (Fig. 54). Fore coxa white to dark and metallic, mid and hind coxae dark and metallic; remaining parts of legs white, except infuscate apical tarsal segment on all legs. Forewing hyaline with a weak median infuscate spot. Petiole dark brown with metallic purple tinges. First gastral tergite metallic bluish-green, remaining tergites metallic dark purple.
Petiole as long as wide to transverse, dorsal surface with weak or strong sculpture. Gaster oval-shaped.
MALE. Unknown. Etymology. From the Latin sulcus = groove, referring to strong groove on posteromedian mesoscutum.