Aegilips chilensis (Hymenoptera: Cynipoidea: Figitidae: Anacharitinae): Redescription and Biogeographic Considerations

Aegilips chilensis Bréthes, 1918 is redescribed and illustrated. Aegilips chilensis is considered an endemic species of the Andean region, characterized for having anteroposterior cephalic processes, resembling spines, formed from the postgenal carina. This and other diagnostic characters are diagnosed and illustrated, and morphological affinities of Aegilips Haliday, 1835 with other Anacharitinae genera are discussed. Redescription and photographs of Aegilips chilensis are given.

While Anacharitinae have a cosmopolitan distribution, being present on all continents except for Antarctica, the bulk of diversity is located in the Neotropical region, where six genera are present, three of them endemic (Acanthaegilips, Calofigites, and Solenofigites), three other genera are Aegilips, Anacharis, and Xyalaspis.Aegilips in the Andean region is only represented by only one species, Aegilips chilensis Bréthes, 1918 known from Chile and Argentina.More recent studies on American Aegilips (unpublished data) have shown a greater diversity of this genus in the Neotropical region.Examination of A. chilensis-type material has shown the presence of some characters that separate it from other Aegilips species, and the cephalic projections are unique among Anacharitinae.

Material and Methods
For this study, specimens of A. chilensis deposited in Museo Argentino de Ciencias Naturales "Bernardino Rivadavia" (MACN, Buenos Aires, Argentina) and Museo de La Plata (MLP, La Plata, Argentina) have been studied, along with undetermined specimens deposited in the Canadian National Collection of Insects, Arachnids and Nematodes (CNC, Ottawa, Canada), the Natural History Museum (NHM, London, UK), and the United States National Museum (USNM, Smithsonian Institution, Washington DC, USA) collections.
Morphological terms used are those of Richards (1977), Ronquist (1995), andRos-Farré et al (2003).All measurements are relative except for the body length.Measurements and abbreviations include the following: F1-F12, first and subsequent flagellomeres; POL (post-ocellar distance) is the distance between the inner margins of the posterior ocelli; OOL (ocular-ocellar distance) is the shortest distance between the inner margin of the compound eye and the outer edge of the posterior ocellus; LOL (lateral-frontal ocellar distance) is the distance between the edges of the lateral and frontal ocelli.Antennal formula includes scape, pedicel, and flagellomere length and relative width in brackets.Biogeographical regions are according to Morrone (2001Morrone ( , 2015)).
Images were taken at the "Serveis Científico-Tècnics" of the University of Barcelona.The field-emission gun environmental scanning electron microscope (Quanta 200 ESEM) was used for high-resolution imaging, under a low voltage (12.0 kV) and without gold-coating of the specimens in order to preserve the material.

Aegilips chilensis Haliday, 1835
Diagnosis.Aegilips lacks a mesopleural groove, unlike Anacharis, Calofigites, Hexacharis, Proanacharis, and Solenofigites thus suggesting a complex of closely related genera formed by Acanthaegilopsis, Aegilips, and Xyalaspis.In the case of Aegilips, the scutellar margin is surrounded by a circumscutellar carina and lacks a scutellar spine, a character that separates this genus from Acanthaegilopsis and Xyalaspis, although this character is not always consistent due to some specimens of Xyalaspis presenting a weak spine.

Aegilips chilensis Bréthes, 1918
Aegilips chilensis Bréthes, 1918: 164 (female) Diagnosis.Aegilips chilensis can be distinguished from other Aegilips for having a smooth malar area without the coriaceous sculpture always present in Aegilips and a postoccipital carina projected backwards and forming two conical anteroposterior processes; this character is absent in other Anacharitinae.
Head.Head triangular-shaped in the front view; as high as wide in the front view and 2.2 to 2.5 times its length in dorsal view.Face smooth and shiny with piliferous punctures, covered by short hairs uniformly distributed.Malar area smooth, absence of malar sulcus.Transfacial line 1.2 times the height of the compound eye.Diameter of the toruli shorter than intertorular distance but equal to eye-torulus distance.Clypeus shortly defined, convex, smooth, and glabrous.Occipital carina absent; postgenal carina present, forming two spiky occipital anteroposterior processes (Fig. 1 a-d): one behind the compound eye and another in the rear malar area.Compound eyes completely glabrous.Ocelli almost arranged in line; POL:OOL:LOL ratio = 5.5:6.0:2.0,being the ocelli diameter 2; Frons smooth and shiny, almost glabrous except for some pubescence near the margin of the compound eye.
Mesosoma.Pronotal plate alutaceous and covered by short pubescence.Pronotum (Fig. 1a) without microsculpture but carinated: lower region obliquely carinated, while upper pronotum presents vertical carinae; pubescence density variable.Mesoscutum (Fig. 1b) width 1.2 to 1.3 its length in dorsal view.Mesoscutum smooth and almost glabrous in its central region, pubescence reduced to the margins of the mesoscutum.Notauli complete but weakening in as they reach the anterior edge, not internally carinated; median mesoscutal furrow absent.Parapsidal signum present but weak; parascutal sulcus absent.Scutellum (Fig. 1b) smooth and glabrous, surrounded by a circumscutellar carina.Scutellum 0.7 to 0.9 times the length of the mesoscutum in dorsal view.Scutellar foveae big and smooth, not basally defined; in some specimens basal carinae of scutellar foveae completely disappeared, making the foveae undistinguishable from scutellar surface.Lateral pits of scutellar foveae absent.Interfoveal line incomplete, short and weak, disappeared in some specimens.Mesopleuron (Fig. 1a) smooth, glabrous and shiny, crossed in the anterior half by some weak oblique ridges.Mesopleural triangle smooth and shiny, covered with sparse hyaline pubescence.Propodeum (Fig. 1a) alutaceous and pubescent; central area glabrous divided by a longitudinal median carina.
Wings.Pubescent.Radial cell of forewing closed, 2.8 times longer than wide.Marginal pubescence of the wing denser at the apical third.
Metasoma.Petiole two times longer than wide, heavily carinated in all sides.Third abdominal tergum 3 times longer than the fourth abdominal tergum in dorsal view.Fifth, sixth and seventh abdominal terga are visible in dorsal view.Metasoma glabrous and smooth; anterior region of the abdominal tergum punctuated from fifth to seventh terga.
Comments.In the original description of Aegilips chilensis, Bréthes did not designate holotype.In Loiacono and Diaz (1977), holotype was designed, which according to article 73.1.3 of the ICZN is incorrect.Due to this evidence, holotype designated by Loiácono and Díaz has been designated as lectotype, and the other specimen belonging to the same series as paralectotype.

Discussion
Weld (1952) separated Anacharitinae in two groups: one characterized by the presence of scutellar spine which included genera Acanthaegilips and Xyalaspis, and another group lacking this feature and including the remaining genera.As mentioned by Restrepo-Ortiz and Pujade-Villar (2010), this character is quite confusing and does not reflect monophyly (Ros-Farré et al 2000, Buffington et al 2007Buffington et al , 2012)).Instead, a new division based on the presence or absence of an oblique groove in the mesopleuron was proposed, thus separating Acanthaegilips and Solenofigites from the rest of Anachari ti na e. Anacharis , Cal ofigi te s, Hexacharis, and Proanacharis also possess a mesopleural groove, although it is not oblique but transverse.It should be taken into account that this character is not synapomorphic: Proanacharis is considered a basal Anacharitinae genus (Ros-Farré et al 2000), while C a l o f i g i t e s i s c l o s e r t o A c a n t h a e g i l i p s a n d Solenofigites; Anacharis and Hexacharis are sister taxa (Buffington et al 2007(Buffington et al , 2012)).The aspect of the groove a l s o v a r i e s b e t w e e n g e n e r a : A n a c h a r i s a n d Proanacharis have a deeply excavated groove with internal carination, while the groove in Calofigites and Hexacharis is weaker and never internally carinated (even effaced in some Hexacharis).
The remaining Anacharitinae genera-Acanthaegilopsis, Aegilips, and Xyalaspis-do not have any groove, but do have a smooth mesopleuron with some wrinkled anterior sculpture which is covered by coriaceous sculpture in Acanthaegilopsis.These three genera are morphologically very similar and their monophyly is supported by phylogenetic studies (Ros-Farré et al 2000, Buffington et al 2007Buffington et al , 2012)).As mentioned before, cephalic features are rare among the anacharitines; only Solenofigites has a postoccipital carina.Aegilips chilensis is a rarity among anacharitines due to the occipital processes.
Taxa from the Andean, Cape, Australian, and Antarctic (fossils) regions are phylogenetically more closely related to each other than they are to those of either region (Morrone 2015).This pattern is also followed by Anacharitinae (Mata-Casanova et al 2015).Moreover, in the case of Neotropical fauna, the southernmost regions of South America and adjacent islands-like Malvinas/Falklands-form a distinct biogeographical entity known as the Subantarctican subregion, which is closely related to the Chilean Central subregion.The distribution pattern of A. chilensis perfectly fits these biogeographical affinities, being endemic to Central Chile, Patagonia, and Malvinas/Falklands.

Figure 1
Figure 1 Aegilips chilensis a Mesosoma in lateral view.b Mesosoma in dorsal view.c Head from rear view.d Head in dorsal view