Four new species of Alloxysta (Hym.: Cynipoidea: Figitidae: Charipinae) from Petr Starý’s collection

. After the revision of Petr Starý’s Charipinae collection, four new species of Alloxysta Förster, 1869 have been found. This collection represents a good overview of the Charipinae fauna worldwide, although the best represented area is Central Europe. Here, we describe four new species of Alloxysta : A. llumae Ferrer-Suay sp. nov., A. onae Ferrer-Suay sp. nov., A. poli Ferrer-Suay sp. nov. and A. staryi Ferrer-Suay & Pujade-Villar sp. nov. Three of these new species are based on reared specimens. Diagnosis, distribution, and description of the new species are given and illustrated.


Introduction
Current knowledge of the taxonomy and tritrophic associations of the Aphidiinae Haliday, 1833 (Hymenoptera: Baconidae) worldwide is deeply indebted to the long-term efforts by Petr Starý, a great naturalist and one of the best specialists.The Aphidiinae is a group of parasitoid wasps of aphids with great potential for use in the biological control of these important pests (Petrović 2022).Thanks to his work, including countless fi eld collections, he managed to form a large collection of Charipinae Dalla Torre & Kieffer, 1910 (Hymenoptera: Figitidae), which are hyperparasitoids of aphids via Aphidiinae, their true hosts within aphids (Ferrer-Suay et al. 2012a, 2019, 2021).
The study of Charipinae is very important because they could potentially affect aphid biological control programs, manipulating the abundance of parasitoids and modifying their behavior (Sampaio et al. 2017).For this reason, the study of the core collections and material previously collected in the fi eld is essential to understand the best way to study the biology and impacts of these species in the fi eld.The Charipinae affect effectiveness of the primary parasitoids and as a result, may increase aphid populations which cause severe yield losses in some of the most important crops (Vázquez-Navarro et al. 2016;Zapata et al. 2016).
Part of the Charipinae (Hymenoptera: Cynipoidea: Figitidae) collection of Petr Starý was previously revised (Ferrer-Suay et al. 2017), 1439 specimens were identifi ed and grouped in 36 species.In the previous work many new trophic associations were published, which is a valuable improvement on the knowledge of the ecological role of this important subfamily.This information was very useful to complete the host list of Charipinae and move forward to future specialization studies based on this subfamily (Ferrer-Suay et al. 2014).A review of Charipinae with all the relevant information about this subfamily was recently published by Ferrer-Suay et al. (2021).
Thanks to the study of this material four new species have been discovered: Alloxysta llumae Ferrer-Suay sp.nov., Alloxysta poli Ferrer-Suay sp.nov., Alloxysta onae Ferrer-Suay sp.nov.and Alloxysta staryi Ferrer-Suay & Pujade-Villar sp.nov.Their diagnoses, descriptions, biology, and illustrative plates are given.The material here described was established as undetermined in the previous work related to Starý's collection (Ferrer-Suay et al. 2017).

Material and methods
The main objective was to study the tritrophic associations (plant-aphid-parasitoid), for this reason many specimens were reared in different ecosystems and habitats.All studied material was collected through the following procedures (Petr Starý pers.com.): a piece of aphid-infested plant was gently cut with scissors and transferred into a plastic jar (250-500 cm 3 ) covered by nylon mesh.The plant was either identifi ed at the spot or taken to an herbarium.Some of the aphids, and even the attending ants, were sampled in 70% ethanol for later identifi cation in the laboratory.The jars were then transferred to the laboratory where they were kept at about room temperature or in a temperature-controlled room under +18-24°C and visited mostly daily to sample the emerged parasitoids.Samples were maintained for about the next two weeks, when the whole sample was re-visited to collect the other (hyper) parasitoids and, eventually, other natural enemies from the litter on the bottom.Each sample was numbered.Specimens of Charipinae were studied using a stereo microscope (NIKON SMZ-1) and environmental scanning electron microscope (FEI Quanta 200 ESEM) belonging to the scientifi c technical services of the University of Barcelona.The fi eld-emission gun environmental scanning electron microscope was used for high-resolution imaging without gold-coating of the specimens.
Morphological terms are taken from Paretas-Martínez et al. (2007).Measurements and abbreviations include F1-F12, fi rst and subsequent fl agellomeres.The width of the fore wing radial cell is measured from the margin of the wing to the beginning of the Rs vein.The transfacial line is measured as the distance between the inner margins of the compound eyes, measured across the face through the antennal sockets divided by the height of the eye.The malar space is measured by the distance from the lower part of the gena from the mouthparts to the ventral margin of the compound eye, divided by the height of the eye.Females and males have the same characters except where indicated.
Type material is deposited in the University of Barcelona collection (UB), Department of Evolutionary Biology, Ecology and Environmental Sciences.

Etymology
This species is dedicated to the daughter of the fi rst author, Llum Garrido Ferrer.

Material examined
Only known from the holotype female.COLOURATION.Head, mesosoma and metasoma brown.Scape, pedicel, F1-F2 dark yellow, F4-F11 yellowish brown.Legs yellow and veins yellowish brown.HEAD.Transversally ovate, smooth, and shiny, slightly wider than high in frontal view.Few scattered setae above toruli.Scattered setae on vertex and many setae on face.Transfacial line 1.1 times the height of compound eye.Malar space 0.6 times the height of compound eye.
MESOSOMA.Pronotum completely covered by scattered setae, with two thin carinae sometimes diffi cult to see under the pubescence (Fig. 1B).Mesoscutum smooth and shiny, round in dorsal view with few scattered setae.Scutellum smooth and shiny with scattered setae, more abundant on apex of scutellum.Propodeum completely covered by abundant pubescence, without carinae present (Fig. 1C).FORE WING.Longer than body, 1.4 times as long as mesosoma and metasoma together.Covered with dense pubescence; marginal setae present and very long (Fig. 1A).Partially open radial cell (nearly closed), 2.3 times as long as wide in female.R1 short and curved; Rs long and slightly curved nearly reachn R1 (Fig. 1F).
METASOMA.Anterior part with an incomplete ring of setae, glabrous at centre, wider laterally.Metasoma smooth and shiny, T3 and T4 clearly distinguished.

Host
Unknown.

Etymology
This species is dedicated to the daughter of the fi rst author, Ona Garrido Ferrer.

Holotype
METASOMA.Anterior part with an incomplete ring of setae, glabrous at centre, wider laterally.Metasoma smooth and shiny, T3 and T4 clearly distinguished.
Alloxysta poli Ferrer-Suay sp.nov.urn:lsid:zoobank.org:act:FD5BACE6-9E86-4FB2-AA01-2C90E9BCF8ACFig. 3 Diagnosis Alloxysta poli sp.nov. is characterized by the presence of pronotal carinae and a propodeal plate, radial cell partially open, 2.3 times as long as wide, rhinaria and club begin on F2, F1 longer than pedicel and subequal to F2, F2 longer than F3.It is closely related to A. macrophadna (Hartig, 1841) but they can be differentiated by the short radial cell, 3.0 times as long as wide in A. macrophadna while it is 2.3 times as wide in A. poli.

Etymology
This species is dedicated to the son of the fi rst author, Pol Garrido Ferrer.

Material examined
Known from seven males and twenty-eight females.
MESOSOMA.Pronotum covered by sparse setae, being less in the distolateral corners and abundant in the anterior margins, with two long and think carinae clearly visible (Fig. 3C).Mesoscutum smooth and shiny, round in dorsal view with few scattered setae.Scutellum smooth and shiny with scattered setae, more abundant on apex of scutellum.Propodeum covered by abundant setae, without carinae and without setae where carinae usually present (Fig. 3D).FORE WING.Longer than body, 1.5 times as long as mesosoma and metasoma together.Covered with dense pubescence; marginal setae present and very long.Partially open radial cell, 2.3 times as long as wide in female paratypes (Fig. 3A), 2.8 times in male, type and paratypes (Fig. 3B).R1 short and curved; Rs long and slightly curved.
METASOMA.Anterior part with an incomplete ring of setae, glabrous at centre, wider laterally.Metasoma smooth and shiny, T3 and T4 clearly distinguished.
Alloxysta staryi Ferrer-Suay & Pujade-Villar sp.nov.urn:lsid:zoobank.org:act:7D3238EA-F499-4FEC-896F-E81A1EF0962BFig. 4 Diagnosis Alloxysta staryi sp.nov. is characterized by the presence of pronotal carinae and propodeal plate, radial cell partially open being 2.3 times as long as wide, rhinaria and club begin on F2, F1 longer than pedicel and subequal to F2, F2 longer than F3.It is similar to A. melanogaster (Hartig, 1840) male but they can be differentiated by the beginning of rhinaria and club: F2 in A. staryi but F3 in A. melanogaster; and the proportion between fl agellomeres: F1 longer than pedicel and subequal to F2, F2 longer than F3 in A. staryi while pedicel-F3 subequal in length in A. melanogaster.

Etymology
This species is named after Petr Starý, collector of the material, and without whom this material would not have been possible to be studied.

Holotype
MESOSOMA.Pronotum covered by disperse setae, being less in the distolateral corners and abundant in the anterior margins, with two small carinae present (Fig. 4B).Mesoscutum smooth and shiny, round in dorsal view with few scattered setae.Scutellum smooth and shiny with scattered setae, more abundant on apex of scutellum.Propodeum covered by abundant long pubescence, with protruding plate, this plate has a few setae on fi rst half and curved sides (Fig. 4D).FORE WING.Longer than body, 1.4 times as long as mesosoma and metasoma together.Covered with dense pubescence; marginal setae present and very long (Fig. 4A).Partially open radial cell, 2.3 times as long as wide.R1 short and curved; Rs long and slightly curved (Fig. 4A).
METASOMA.Anterior part with an incomplete ring of setae, glabrous at centre, wider laterally.Metasoma smooth and shiny, T3 and T4 clearly distinguished.

Discussion
The group of Hymenoptera Parasitica is hyper diverse and there are many species waiting to be discovered.As is well known, the study of entomological collections could give us the opportunity to fi nd new species and in this way improve our knowledge about this subfamily with data about their distribution patterns as well as their host relationships.Especially in Charipinae, there are few data about their trophic relations, most of the material originating from fi eld catches or by Malaise traps.There is also very limited information about the grade of specialization of species of Charipinae; this should be addressed in future work on this subfamily, for example by dedicated rearing efforts, documenting species and their ecology.
Petr Starý collected many specimens, being a specialist of the subfamily Aphidiinae, and usually he collected mummies.Sometimes, instead of Aphidiinae, Charipinae appear, because they are parasitoids of the aphidiines.This collection has around 1500 specimens from around the world; in Ferrer-Suay et al. (2017) Central European material was studied, but there is also material collected from other geographical regions (Ferrer-Suay et al. in prep.).
Thanks to the study of this collection, we have found four new species of Alloxysta here described, three from Czech Republic (A. llumae sp. nov., A. onae sp. nov. and A. staryi sp. nov.) and another from the Czech Republic, Slovakia and Cuba (Alloxysta poli sp.nov.).
Although type material is scarce for the new species described from Czech Republic, we have no doubt that they are new species since the distinctive characters are different from any of the Alloxysta known species (Ferrer-Suay et al. 2018).
Alloxysta poli sp.nov.has a Holarctic distribution having been collected in the Czech Republic and Cuba.It is not uncommon in Charipinae for this to happen, similar cases having been cited in other species, for example Phaenoglyphis americana Baker, 1896 was described from the Nearctic region, and has been cited from the Palaearctic (France and Spain) by Ferrer-Suay et al. (2012b, 2015).Globalization, with transport of aphids, and the fact that its host associations are not very specifi c, can result in a very extensive distribution.
With these species described here, there are now 109 described Alloxysta (61 in Europe, 34 in America and 14 on both continents).
The study of the Petr Starý collection has so far provided a multitude of biological data and an extension of the distribution areas of many species of Charipinae (Ferrer-Suay et al. 2017).Additionally, with the description of the new species, we are still investigating possible new records from other countries.
DiagnosisAlloxysta onae sp.nov. is characterized by the partially open radial cell, pronotal carinae absent, propodeal carinae present, forming a plate and antennae with rhinaria beginning in F2 and club shape in F3, female with F1 longer than pedicel and F2, F2 longer than F3 and F3 subequal to F4.It is closely related to A. longipennis (Hartig, 1841) but they can be differentiated by the presence of pronotal carinae, absent in A. onae while they are present in A. longipennis.