Pimpla Fabricius, 1804 (Ichneumonidae, Pimplinae) from Uruguay: a replacement name, new records, and an identification key to the species

Abstract We report new faunistic records of Pimpla Fabricius, 1804 from Uruguay. The following species are reported from the country for the first time: P. albomarginata Cameron, 1846, P. caerulea Brullé, 1846, P. perssoni Gauld, 1991, and P. semirufa Brullé, 1846. In addition, we propose a replacement name for Pimpla rufipes Brullé, 1846 and provide diagnosis, digital images, and an identification key for all the Pimpla species known to occur in Uruguay.


Introduction
The parasitoid wasp family Ichneumonidae (Darwin wasps) is among the largest animal families anywhere on Earth (Klopfstein et al. 2019). It is taxonomically challenging, and many species are either rare or at least rarely collected. One exception is the genus Pimpla Fabricius, 1804 (Pimplinae, Pimplini). It is composed of moderately large (in tropical regions), often colorful species which are abundant in many entomological collections (Townes 1969;Porter 1970).
With over 200 valid species (Yu et al. 2016;Watanabe and Matsumoto 2019), this genus is among the largest genera within the subfamily Pimplinae. The species of Pimpla are known to be idiobiont endoparasitoids of prepupae and pupae of Lepidoptera (Gauld 1991). The genus is characterized by simple and large tarsal claws (females), the straight apex of the ovipositor, a weakly concave internal margin of the compound eye in front of the antennal insertion (Gauld et al. 1998), and mid tarsomere IV medioventrally with a longitudinal band of fine hair (an autapomorphy) (Gauld et al. 2002).
On account of the taxonomical works of Charles C. Porter in South America (Porter 1970) and Ian D. Gauld in Central America (Gauld 1991;Gauld et al. 1998), the genus is one of the best-known Darwin wasp genera in the Neotropical region. Porter (1970) reported 35 (21 of them new) species from South America and Gauld (1991) and Gauld et al. (1998) found 17 (six of them new) species in Costa Rica.
The aim of this paper is to provide new records of Pimpla from Uruguay. In addition, we provide diagnosis, high-quality layer-stacked photographs, and an identification key for the species currently known from the country. This work is part of a series of articles reporting new Pimpla records from South America. This work was started by Pádua et al. (2019).

Study area
The field sampling was conducted in four locations in the municipality of Castillos, Rocha Department, Uruguay, between December 2014 and December 2016 (see Fernandes et al. 2019).

Specimens studied
Pimpla specimens were collected by Malaise trapping, and the specimens are deposited in the Invertebrate Collection of Instituto Nacional de Pesquisas da Amazônia (INPA; curator: Marcio L. Oliveira).

Morphology and distribution
General morphological terminology follows that of Gauld (1991). New distributional records are indicated with an asterisk (*).

Photographs
Digital images were taken using a Leica DMC4500 digital camera attached to a Leica M205A stereomicroscope and combined using the software Leica Application Suite V4.10.0. The final images were edited in Adobe Photoshop.

Distribution maps
The distribution maps were created using SimpleMappr online software (Shorthouse 2010). Mesosoma and metasoma with a metallic blue (Fig. 2B) 10B); metasoma reddish brown, with a pair of large yellow blotches laterally on tergites I-IV (Fig. 10B)  Diagnosis. Pimpla can be distinguished from other genera of Pimplini (sensu Porter 1970 as Coccygomimus) by the combination of the following character states: 1) inner margin of eye weakly to rather strongly concave above antennal socket; 2) clypeus not divided by a transverse suture; 3) malar space 0.35-1.4 times as long as basal width of mandible; 4) mandible broad and with upper tooth approximately as long as the lower tooth; 5) notaulus weak or absent, without a distinct frontal crest; 6) propodeum with median longitudinal carinae varying from absent to sometimes weakly traceable throughout; 7) pleural carina usually present but sometimes absent; 8) length of fore wing 2.7-18.0 mm; 9) hind femur without a ventral tooth; 10) tarsal claws large and simple, without a basal lobe or an enlarged hair with a flattened tip; 11) metasoma varying from closely punctured to sometimes almost impunctate; 12) females with ovipositor approximately straight, ovipositor tip never sharply decurved. Gauld et al. (2002) found a single autapomorphy for the genus: mid tarsomere IV medioventrally with a longitudinal band of fine hairs. Figure 1A-F Pimpla albo-marginata Cameron, 1886: 267. Holotype ♀, Mexico (BMNH). Coccygomimus albomarginatus; Townes and Townes 1966: 24. Diagnosis. This species can be distinguished from the other Uruguayan species of the genus by the combination of the following character states: 1) wings hyaline; 2) clypeus with apex deeply bilobed; 3) malar space wide, longer than basal mandibular width, that males less than 0.6 times basal mandibular width; 4) mesoscutum entirely black; 5) postscutellum black; 6) mesopleural suture weakly faveolated; 7) propodeum with conspicuous posterolateral tubercles; 8) fore wing Rs more or less straight and cu-a slightly distal to the base of Rs&M; 9) coxae without black markings and fore coxa with markings; 10) metasoma black and white banded; 11) laterotergites V broad, more than 0.5 times as broad as long; 12) tergite I of female short and broad, in profile strongly convex, in profile with moderately high blunt hump; 13) sternite I with strongly produced swelling; 14) apex of ovipositor with dorsal valve of ovipositor apically without teeth.

Pimpla albomarginata Cameron, 1886
Biological notes. Nothing is known about the host preferences of this species. Distribution. Colombia, Costa Rica, Mexico, Panama, Venezuela, and Uruguay* (Fig. 11A).  Diagnosis. This species can be distinguished from the other Uruguayan species of the genus by the combination of the following character states: 1) wings more or less blackish; 2) body metallic blue (male with fore coxae white marked); 3) laterotergite V narrow, less than 0.3 times as long as wide.
Biological notes. Parasitoid of Alabama argillacea (Hübner, 1818) (Noctuidae) (Porter 1970 Diagnosis. This species can be distinguished from the other Uruguayan species of the genus by the combination of the following character states: 1) wings darkened; 2) mesosoma and metasoma black; 3) laterotergite V 1.6-1.7 times as long as wide; 4) legs orange, except coxa, trochanter, trochantellus, apex of hind tibia and tarsus black; 5) tergite II silky shining, slightly coriaceous and mostly (except of apical rim), with almost uniformly distributed, large, strong, from more or less adjacent to confluent punctures; 6) malar space 1.0-1.2 times as long as basal width of mandibles; 7) ovipositor approx. 1.75 times as long as hind tibia; 8) ovipositor cylindric, with apex of dorsal valve without teeth and ventral valve with gently convex teeth on tip.
Biological notes. Nothing is known about the host preferences of this species. Distribution. Argentina and Uruguay (Fig. 11C). Material examined. Syntype, Chile (♀, EY9374), examined by photo ( Fig. 3A-C). Syntype, Chile (sex undetermined, EY9375), examined by photo (Fig. 4A-C). Remarks. Brullé (1846) described P. cyanipennis based on specimens from Montevideo (Uruguay; C. Gaudichaud collector). Later, Porter (1970) expanded the distribution of the species to Argentina. However, Porter did not study the type specimens of this species, deposited at MNHN. We analyzed the syntypes (EY9374 and EY9375), and verified that the type locality on the label is in Chile (C. Gay collector). The French botanist and naturalist Claude Gay carried out several expeditions in the Andes, es- pecially in Chile and Peru. A large part of the material deposited by him in MNHN originated from these countries. Furthermore, Gaudichaud, who was appointed by Brullé as a collector of types, made several expeditions in Uruguay and Brazil (materials also deposited in MNHN). Thus, we hypothesize that: 1) the labels may have been unintentionally replaced in specimens, 2) the photos of the labels may have been added to the specimens in a wrong way in the MNHN database, or 3) Brullé may have confused the type locality when describing this species. Townes (1961) corrected inconsistencies in type localities in some species described by Brullé in MNHN, but he did not men-tion this species. In fact, we have studied the type specimens by using only photos, and we believe that only an in situ specimen examination can solve this inconsistency. Thus, we have decided to report this species only from Argentina and Uruguay.
Diagnosis. This species can be distinguished from the other Uruguayan species of the genus by the combination of the following character states: 1) wings hyaline; 2) mesosoma black with hind corners of meso-and metapleuron brown and tegula white; 3) metasoma reddish; 4) laterotergite V 1.3 times as long as wide; 5) legs reddish, except of fore coxa often more or less broadly blackish basally, hind tibia sometimes slightly dusky, especially near apex, and tarsi usually duller often slightly dusky on apical segment; 6) tergite II shiny and with almost uniformly distributed large, deep, adjacent to reticulately confluent punctures, except narrowly smooth on apex; 7) malar space 0.8-1.0 (0.6-0.9 in male) times as long as basal width of mandibles; 8) ovipositor approx. 1.45-1.7 times as long as hind tibia; 9) ovipositor cylindric, dorsal valve with apex without teeth and ventral valve with gently convex teeth on tip.
Biological notes. Parasitoid of Gelechiidae: Pectinophora gossypiella (Saunders, 1844); Noctuidae: Alabama argillacea (Hübner, 1818) (Porter 1970 (Porter 1970). Based on the material collected in our samples in Uruguay, we verified that the peak of occurrence of this species in the sampled locations was between November and January.
Distribution. Argentina, Bolivia, Brazil, Colombia, Paraguay, and Uruguay (Fig. 11D). Remarks. Townes and Townes (1966) reported a new species of Coccygomimus as "Coccygomimus n. sp." from Argentina and considered Ephialtes kreibohmi Blanchard, 1942 to be nomen nudum of it. Later, Porter (1970) described the species mentioned by Townes and Townes (1966) Townes and Townes 1966: 27. Diagnosis. This species can be distinguished from the other Uruguayan species of the genus by the combination of the following character states: 1) wings hyaline with weak brownish staining; 2) mesosoma shining black; 3) metasoma black with more or less brown staining on apical rims; 4) laterotergite V 1.4-1.6 times as long as wide; 5) legs orange with fore coxae orange or black, fore and mid tarsi slightly duller orange to slightly dusky, hind tibia duller orange with rather weak blackish staining on apex, hind tarsus extensively blackish to black; 6) tergite II with larger and stronger punctures; 7) malar space 1.0-1.1 (0.85-1.0 in male) times as long as basal width of mandibles; 8) ovipositor 1.3-1.6 times as long as hind tibia; 9) ovipositor cylindric, apex of dorsal valve without teeth and ventral valve with gently convex teeth on tip.
Distribution. Argentina, Brazil, and Uruguay (Fig. 11F). Material examined. Lectotype, Uruguay, Montevideo (♀, EY9414), examined by photo (Fig. 8A-C). URUGUAY, Rocha, Branaa, Agricultura, 34°02'33.7"S, 53°50'03.1"W, 11.II.2015, Malaise trap II (E. Castiglioni and team leg.), 1♂, INPA; idem, but 12.I.2015, Malaise trap I, 3♂♂, INPA; idem, but Malaise trap II, 1♀ and  Diagnosis. This species can be distinguished from the other Uruguayan species of the genus by the combination of the following character states: 1) wings yellowish with distal margin of the fore wing blackish; 2) mesosoma yellow with black marks on mesoscutum (three stripes), hind part of tegula, hind margin of scutellum, anterior margin of mesopleuron, 7-shaped mark on upper hind part of mesopleuron, a continuous anterior band along the anterior margin of metapleura, and propodeum and hind rim of propodeum; 3) metasoma yellow with tergites I-IV anteriorly broadly and posteriorly narrowly black and with posterior tergites anteriorly black; 4) laterotergite V 2.7-3.4 times as long as wide; 5) legs yellow with dorsal longitudinal black band on med and hind coxa, fore, mid and hind (except the first tarsomere) tarsi strongly infuscate, femur darkened dorsally and ventrally, and tibia infuscate proximally, tibia with close and dark pubescence, giving them a dirty yellow appearance; 6) tergite II highly polished, with very fine sparse punctures, and with anterolateral corners separated by deep oblique grooves; 7) malar space 0.3-0.4 times as long as basal width of mandi- bles; 8) ovipositor 1.25-1.3 times as long as hind tibia; 9) apex of ovipositor slightly compressed, with weak denticles arranged in a median row on dorsal valve, and with ventral valve not expanded laterally, with a few weak teeth.
Biological notes. Nothing is known about the host preferences of this species. Distribution. Brazil, Costa Rica, Mexico, and Uruguay* (Fig. 11E). Diagnosis. This species can be distinguished from the other Uruguayan species of the genus by the combination of the following character states: 1) wings hyaline; 2) mesosoma shining black, lower hind corner of mesopleuron slightly brown, metapleuron red with a little black staining along front margin and sometimes also along dorsal margin and propodeum red with slight to extensive black staining basad and in spiracular area; 3) metasoma reddish with tergite V often with a little blackish staining apico-laterally, tergite VI broadly to almost wholly black and tergites VII+ completely black; 4) laterotergite V 1.6 times as long as wide; 5) legs red, fore coxa black except becoming more or less broadly reddish toward apex below, fore and mid tarsi little duller reddish with slight dusky staining on apical segment, hind femur often with slight dusky tinge above on apex, hind tibia dull red basad and blackish on about apical half, hind tarsus dull red with much dusky staining; 6) tergite II shining with abundant, large, strong, mostly adjacent to confluent punctures, except on the narrow smooth apical rim; 7) malar space 0.8-1.0 times as long as basal width of mandibles; 8) ovipositor 1.3-1.6 times as long as hind tibia; 9) ovipositor cylindric, apex of dorsal valve without teeth and ventral valve with gently convex teeth on tip.
Biological notes. Nothing is known about the host preferences of this species. Distribution. Argentina, Brazil, and Uruguay* (Fig. 11G) Townes and Townes 1966: 28. Diagnosis. This species can be distinguished from the other Uruguayan species of the genus by the combination of the following character states: 1) wings hyaline with pale yellow staining; 2) mesosoma shining black with variable yellow markings on pronotum, tegula, scutellum, postscutellum, and propodeum (a pair of elliptic blotches); 3) metasoma reddish brown with a pair of large yellow blotches laterally in tergites I-II (tergites I-IV in males); 4) laterotergite V 2.1-2.2 times as long as wide; 5) legs yellow, except for fore and mid coxa black (sometimes) and hind coxa with a black mark, femur and basal half of tibia reddish brown and last tarsus blackish; 6) tergite II rather dully to brightly shining with moderately strong to fine or very fine micro-reticulation and mostly sparse, irregularly spaced, small to large, obscure to well-defined punctures; 7) malar space 0.6-1.0 (0.4-0.7 in male) times as long as basal width of mandibles; 8) ovipositor 1.45 times as long as hind tibia; 9) ovipositor moderately depressed, apex of dorsal and ventral valves apically with teeth, the apical ridge-bearing portion not unusually flattened and in profile slightly convex.

Discussion
During the last 30 years, the Darwin wasp fauna of some Neotropical countries (i.e. Brazil, Costa Rica, and Peru) have been sampled in more detail. These studies have revealed a very high species richness and a plethora of new taxa from many parts of the region (e.g. Gauld 1991;Sääksjärvi et al. 2004;Veijalainen et al. 2012). However, most parts of the Neotropical region have remained understudied. Uruguay's biodiversity knowledge is still very fragmentary (Aldabe et al. 2008) and this is also shown by the genus Pimpla. Before of our study, only four species of Pimpla were known from the country: P. cyanipennis Brullé, 1846;P. golbachi (Porter, 1970); P. patirrufa nom. nov.; and P. tomyris Schrottky, 1902(Yu et al. 2016. Here, we have doubled the species richness of Pimpla in Uruguay. All known Uruguayan species are also widely distributed in South America or the Neotropical region in general. Pimpla Figure 11. Distribution of Pimpla spp. in Uruguay A P. albomarginata Cameron, 1886 B P. caerulea Brullé, 1846 C P. cyanipennis Brullé, 1846 D P. golbachi (Porter, 1970) E P. perssoni Gauld, 1991 F P. patirrufa nom. nov. G P. semirufa Brullé, 1846H P. tomyris Schrottky, 1902. Gray area = Rocha Department. Red circle = previous record. Blue circle = new record. species are moderately large and strong-flying insects, which explains their wide distribution over vast regions.
We hope that this study draws more attention to Uruguay's apparently rich, but very little-known, Darwin wasp fauna.