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Research Article
Phylogenomics and taxonomic revision of Stenarella Szépligeti (Hymenoptera, Ichneumonidae, Cryptinae)
expand article infoBernardo F. Santos, Santiago Bordera§
‡ Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity Science, Berlin, Germany
§ University of Alicante, Alicante, Spain
Open Access

Abstract

The phylogeny of the Old World genus Stenarella Szépligeti is investigated using data from genomic ultraconserved elements. Thirteen species from the four biogeographic regions where the genus occurs were included in the analyses. The resulting trees corroborate the monophyly of Stenarella and show substantial biogeographic structure within the genus, with three major clades corresponding to species occurring in the Afrotropical, Palearctic and Oriental–Australasian regions. The Afrotropical component of the genus is represented by fifteen species, while outside of the Afrotropics there seem to be only three species, each with very wide geographic distribution. The taxonomic limits of the genus are reviewed, and an updated diagnosis is provided. A total of eighteen species are recognized as valid, of which twelve are described as new: S. aurea sp. nov., S. bugalana sp. nov., S. delicata sp. nov., S. favilla sp. nov., S. fenestralis sp. nov., S. hopkinsi sp. nov., S. katanga sp. nov., S. natalina sp. nov., S. nigriscuta sp. nov., S. nigromaculata sp. nov., S. saaksjarvi sp. nov., and S. vannoorti sp. nov. An updated identification key to the species of the genus is provided, as well as diagnoses, redescriptions, and new distribution records for the previously known species.

Keywords

Darwin wasp, Cryptini, Osprynchotina, mud-nesting, biogeography

1. Introduction

Stenarella Szépligeti, 1916 is a small genus of cryptine wasps (Hymenoptera, Ichneumonidae, Cryptinae), originally described to accommodate a large number of species of somewhat heterogeneous morphology but mostly defined by the small and square areolet; postpetiole longer than wide and distinctly wider than petiole; and T2 longer than wide. Roman (1943) designated Ichneumon gladiator Scopoli 1763 (currently a synonym of Ichneumon domator Poda 1761) as the type species of the genus. Townes (1957), while revising the generic names for Old World ichneumonids, noted the similarity between S. gladiator and the two species of the Indo–Australian Orientostenaraeus Uchida, and later on, synonymized the two genera (Townes in Townes et al. 1961). In the course of several works (Townes et al. 1961; Townes and Townes, 1966, 1973), Townes transferred most of the species previously assigned to Stenarella to other genera, arriving at the modern concept of the genus that included only six valid species: the type species S. domator (Poda), widely distributed in the Western Palearctic region and recently recorded an introduced species in Ohio, United States (Wahl and Green 2020); S. insidiator (Smith) from the Oriental region; S. victoriae (Cameron) from Australia; and three Afrotropical species: S. brykella Roman, S. lissonota (Cameron) and S. tripartita (Brullé).

All of these species share the following characters that have been used in the keys and descriptions of Townes’ revisionary works (Townes et al. 1961; Townes 1970; Townes and Townes 1973): mandible long with the dorsal tooth much longer than ventral one; areolet very small; first metasomal tergite slender, not widened posteriorly, its spiracle placed on posterior 0.42, first sternite ending opposite or posterior to spiracle; and ovipositor very long (>2.8× as long as hind tibia) with distinct hooked teeth on the tip of its dorsal valve.

Species of the genus are parasitoids of mud-nesting aculeate wasps and bees (Andrenidae, Megachilidae, Pompilidae, Sphecidae, Vespidae). Host records are available for four of the six species. According to the compilation of Yu et al. (2016), Stenarella domator is also recorded as a parasitoid of non-aculeate insects such as Coleoptera (Buprestidae, Cerambycidae, Curculionidae), Hymenoptera (Siricidae), and Lepidoptera (Noctuidae, Notodontidae, Tortricidae), but these records are likely spurious (see Wahl and Green 2020).

Stenarella was placed by Townes (1970) in the subtribe Nematopodiina (=Osprynchotina per the International Code of Zoological Nomenclature), which were characterized by the unusually long mandible, slender first metasomal tergite and the use of mud-nesting wasps as hosts. In a large-scale phylogenetic analysis of Cryptini (Santos 2017), Osprynchotina were shown to be polyphyletic. Part of its genera were recovered as a monophyletic group (Messatoporus Cushman, Photocryptus Viereck, Osprynchotus Spinola and Acroricnus Ratzeburg) but other five were scattered across a large, morphologically heterogeneous clade (Mesostenus group in Santos 2017). Due to the non-monophyletic status of most of Townes’ subtribes, Santos (2017) recommended the abandonment of this classification scheme in favor of informal genus groups. Subsequent works using genomic data from ultraconserved elements, however, recovered three other genera from the former OsprynchotinaStenarella, Picardiella and Nematopodius—as a clade, with each genus monophyletic (Supeleto et al. 2020; Santos and Brady 2024). Stenarella, represented by two undetermined species, had also been recovered as monophyletic in earlier molecular phylogenetic studies (Laurenne et al. 2006; Quicke et al. 2008)

In spite of their broad distribution and distinctive morphology, no new species of Stenarella have been described since 1943, even though Townes (1970) indicated the presence of a number of undescribed species from the Afrotropical region. Furthermore, the internal phylogeny of the genus has never been investigated in detail, with only two species included in previous phylogenetic treatments. The broad distribution of the genus prompts the question of whether the relationships among its species are structured according to zoogeographic regions. In this context, this work aims investigate the internal phylogeny of Stenarella and to provide a taxonomic review of the species of the genus, to describe new species and provide new distribution records.

2. Material and Methods

2.1. Material examined

In this work we included 328 specimens assigned to Stenarella from the following institutions:

CEUA: Colección Entomológica de la Universidad de Alicante (Alicante, Spain); FSCA: Florida State Collection of Arthropods (Gainesville, USA); IRSNB: Institut Royal des Sciences Naturelles Belgique (Bruxelles, Belgium); MFNB: Museum für Naturkunde (Berlin, Germany); MNHN: Muséum National d’Histoire Naturelle (Paris, France); NHMUK: The Natural History Museum (London, UK); NHRS: Naturhistoriska Riksmuseet (Stockholm, Sweden); OXUM: Oxford University Museum of Natural History (Oxford, UK) — ROM: Royal Ontario Museum (Toronto, Canada); SAMC: Iziko South African Museum (Cape Town, South Africa); TMA: Természettudományi Múzeum Allattára (Budapest, Hungary); UKICU: University of Kentucky Insect Collection (Kentucky, USA); USNM: National Museum of Natural History (Washington, DC, USA); USUC: Utah State University (Logan, USA); ZIU: Zoologiska Institutionen (Uppsala, Sweden); ZMUT: Insecta collections of Turku University (Turku, Finland); ZSMC: Zoologische Staatssammlung (Munich, Germany).

2.2. Taxonomy

Morphological terminology follows Broad et al. (2018). The first and subsequent flagellomeres are referred to as f1, f2, etc. The first and subsequent tarsomeres are referred to as t1, t2, t3, etc, while first and subsequent metasomal tergites are referred to as T1, T2, T3, etc. Biometric ratios used in descriptions are as follows: MLW, mandible maximum length/maximum width; MSM, malar space maximum width/basal width of mandible; CWL, clypeus maximum width/maximum length; SLW, propodeal spiracle maximum length /maximum width; APH, forewing cell 1+2Rs (areolet) maximum high/pterostigma maximum high; AWH, 1+2Rs maximum width/maximum high; HW1C, hind wing vein Cua/cu-a length; T1LW, first metasomal tergite maximum length/maximum width (dorsal view); T1WW, first metasomal tergite maximum width/minimum width (dorsal view); T2LW, second metasomal tergite maximum length/maximum width (dorsal view); T2WW, second metasomal tergite maximum width/minimum width (dorsal view); OST, ovipositor sheath length/ hind tibia length.

All measurements were rounded to the nearest 0.05; ratios expressed to the nearest tenth reflect estimated proportions. For description of the propodeum microsculpture we consider two areas: anterior area, between anterior margin and anterior transverse carina, and posterior area, between anterior transverse carina and posterior rim.

Images were captured using a custom photo system composed of a Sony alpha 7R III camera and Canon 65 mm and 90 mm objective lenses attached to a Novoflex CASTEL–MICRO Auto Step Motor Stack Rail System and using an Alladin 70w lighting setup. The captured stacks were exported and combined in a single in–focus image using Helicon Focus 6 and further processed with Adobe Photoshop 2024. Photos of the holotype of S. lissonota, however, were made with a simple Canon EOS Rebel T6 camera attached to a standard macro lens.

Distribution maps were made with the online open tool SimpleMappr (Shorthouse 2010). Coordinates are mostly according to the GEOnet Names Server (http://geonames.nga.mil/gns/html/index.html), but several other sources were used where necessary. Specimens with a NR note indicate new country records for that species relative to the latest catalog of Yu et al. (2016). New records obtained from GBIF (http://gbif.org) are indicated as NR-G, and records from iNaturalist (www.inaturalist.org) as NR-I; records from such public databases are only included when the identity of the respective species could be confirmed by the authors via reliable photo information.

2.3. Molecular phylogenetics

To assess the monophyly of Stenarella and the relationships among its species, we generated molecular data from genomic ultraconserved elements (UCEs; Faircloth et al. 2012) for all species from which we could extract viable DNA samples—many species were singletons or represented only by very old museum specimens that yielded low DNA recovery success (File S1). This resulted in an ingroup assemblage of 14 species, including two specimens of S. insidiator corresponding to its larger and smaller forms, and three specimens of S. domator corresponding to its recognized subspecies. The outgroup included a selection of 32 Cryptini taxa, mostly from the “Mesostenus clade” (sensu Santos 2017), drawn from the extensive treatment of the tribe in Santos and Brady (2024). The phygadeuontine Hemigaster taiwanus (Sonan) was used to root the tree. Laboratory protocols, including extraction, library preparation and targeted enrichment, followed the methods described in detail in Santos et al. (2021), with the following adjustment: library preparation was conducted either using the Kapa Hyper Prep Kit (Kapa Biosystems, Wilmington, MA, U.S.A.), with custom dual-indexing adapter-primers (Glenn et al., 2019), or with the Qiagen QIAseq Ultraow Input Library kit (Qiagen, Hilden, Germany), with end-polishing and ligation reactions conducted at quarter volume. Size-selected and enriched pools were sequenced as partial lanes in Illumina NovaSeq 4000 or NovaSeq X Plus Series platforms (2×150; Illumina Inc., San Diego, CA, U.S.A.). Raw sequence reads for all samples are available from the NCBI Sequence Read Archive under BioProject accession PRJNA632862.

All sequence data processing and analysis was conducted using the Smithsonian’s High-Performance Computing cluster (Smithsonian Institution, 2020). We used the Phyluce v1.5 pipeline (Faircloth, 2016) to process the data from raw sequence reads to final phylogenetic matrices, incorporating either Trinity v. r2013-02-25 (Grabherr et al., 2011) or SPAdes (Bankevich et al., 2012) for assembly, MAFFT v. 7.130b (Katoh et al., 2002) for alignment and GBLOCKS v. 0.91b (Castresana, 2000; Talavera and Castresana, 2007) for alignment trimming. Alignments were filtered to include only loci available for at least 50% of the taxa, resulting in a final alignment including 1,560 loci. The concatenated dataset (File S2) was partitioned using PartitionFinder2 (Lanfear et al., 2016) aided by the SWSC-EN algorithm (Tagliacollo and Lanfear, 2018). Phylogenetic analyses were run using IQTREE v1.6.12 (Nguyen et al., 2015), with 10 000 rounds of ultra-fast bootstrapping (Hoang et al., 2018) to assess clade support and ModelFinder (Kalyaanamoorthy et al., 2017) to choose the best model for each partition using the argument ‘-MFP’.

3. Results

3.1. Taxonomy

Stenarella Szépligeti, 1916

Stenarella Szépligeti, 1916: 307. Description. Type species: Ichneumon gladiator Scopoli, 1763 (= Ichneumon domatorPoda 1761), by subsequent designation by Roman (1943).

Orientostenaraeus Uchida, 1930: 321. Type species: Orientostenaraeus chinensis Uchida, 1930 (= Mesostenus insidiatorSmith 1859), by original designation.

Parasilsila Cheesman, 1936: 368. Type species: Parasilsila trilineata Cheesman, 1936 (= Mesostenus victoriae Cameron), by original designation.

Diagnosis.

Ovipositor very long, OST >2.7 (Fig. 1), its apex distinctly sinuate, dorsal valve with distinct teeth or hooks; mandible long, MLW 1.85–2.6, dorsal tooth much longer than ventral one; areolet very small; T1 0.85–1.25 as long as T2, slender and not widened posteriorly, its spiracle placed on posterior 0.42.

Figure 1. 

Propodeum of Stenarella spp., females. A Stenarella brykella (lectotype); B Stenarella victoriae; C Stenarella insidiator; D Stenarella domator; E Stenarella tripartita; F Stenarella delicata (holotype); G Stenarella nigromaculata (holotype); H Stenarella aurea (holotype); I Stenarella saaksjarvi (paratype); J Stenarella hopkinsi (paratype); K Stenarella nigriscuta (paratype); L Stenarella vannoorti (holotype).

Description.

FEMALE. Fore wing 5–16.5 mm long. — Head: Mandible long to very long, MLW 1.85–2.6, distinctly tapered towards apex; dorsal tooth distinctly longer than ventral one. Malar space moderately long, MSM 0.3–0.7. Clypeus wide, CWL 2.45–3.3 trapezoidal in front view, almost flat in lateral view; apical margin sharp, distinctly raised, inflexed, medially concave, without median teeth. Flagellum of uniform width, not distinctly enlarged towards apex; apical flagellomere uniformly tapered. Supra-antennal area ventrally slightly to distinctly concave, with distinct longitudinal carina. Occipital carina complete, meeting hypostomal carina far from mandible base. — Thorax. Dorsal margin of pronotum regular, not swollen; epomia indistinct to strong and long. Mesoscutum moderately convex; notaulus long, reaching 0.55–0.9 of mesoscutum length, moderately impressed, inside with weak to distinct striae. Epicnemial carina weak, sometimes almost indistinct from mesopleuron sculpturing, reaching 0.4–1.0 of distance to subtegular ridge. Sternaulus sinuous or evenly up curved, deep or shallow, reaching or not the posterior rim of mesopleuron, anteriorly placed somewhat ventrally on mesothorax, facing downwards, inside with very short to distinct vertical striae. Median portion of postpectal carina short, straight. Posterior margin of metanotum without lateral teeth–like projections. Propodeal furrow moderately deep, medially smooth. Juxtacoxal carina as short ridge or absent. Pleural carina absent. Fore tibia not distinctly swollen. Fourth tarsomeres not distinctly bilobed. — Propodeum. Propodeum moderately long, dorsal profile in lateral view evenly convex and sloped, in S. domator propodeum short with dorsal profile broken into a short anterodorsal face and a longer sloped posterodorsal face. Anterior margin medially slightly concave. Spiracle elongate, SLW 1.4–5.25. Longitudinal carinae absent, except sometimes anterior section of median longitudinal carina (area basalis defined). Anterior transverse carina complete, sharp, medially distinctly arched. Posterior transverse carina absent to complete, never forming distinct sublateral crests. Longitudinal carinae absent. — Wings: Nearly hyaline to strongly infuscate, dark brown. Forewing vein 1-Rs+M with bulla placed centrally to apically; ramellus absent; crossvein 1cu-a arising far from 1M+Rs, basad by 0.2–0.4 of its own length; vein 2Cua 1.05–1.80× as long as crossvein 2cu-a; crossvein 2m-cu uniformly arched, bulla moderately short, placed medially to anteriorly; cell 1+2Rs large, APH 0.25–0.55, about as long as wide or slightly longer, AWH 0.80–1.20, pentagonal; crossveins 2r-m and 3r-m subparallel, to distinctly convergent, both veins usually about the same length. Hind wing vein 1-M+Cu apically almost straight to slightly convex; vein Cua slightly to much longer than crossvein cu-a, HW1C 1.74–3.0; vein Cub distinct, its apical 0.5 almost straight to distinctly convex; vein 2-1A reaching at least 0.9 of distance to wing margin. — Metasoma: T1 moderately long, about 0.51–0.58× as long as mesosoma (from base of T1 to apex of mesoscutum, measured laterally), slender, T1LW 2.15–3.55, apex distinctly wider than base, T1WW 1.70–2.85, approximately cylindrical, without sharp anterolateral tooth, but sometimes with weak anterolateral flange; spiracle of T1 placed distinctly posteriorly to midlength, not distinctly prominent; median dorsal and dorsolateral carina absent or vestigial; ventrolateral carina usually distinct. Setae on T2–8 always associated with small punctures. T2 slender, T2LW 0.95–1.6, T2WW 1.75–2.7; thyridium subcircular or longer than wide. T7–8 about as long as T5–6. Ovipositor very long, OST 2.8–5.4, slender, distinctly upcurved (often contorted), scarcely punctate, basally cylindric, apically distinctly depressed; apex of ovipositor blunt, without nodus or notch; dorsal valve with 4–9 small teeth; ventral valve apically dilated and overlapping dorsal valve as a sheath, with 7–11 distinct teeth. — MALE. Known only for S. domator, S. lissonota, S. insidiator, S. tripartita, S. victoriae and S vannoorti sp. nov. Generally similar to the respective females, but not as readily recognizable as members of the genus due to the lack of the most conspicuous diagnostic trait, the long ovipositor. Morphological secondary sexual differences are usually more or less uniform within Cryptini, as noted by Santos and Aguiar (2013), and apply to the males of Stenarella as follows. General body size usually smaller than respective females. Head with extensive whitish marks on mandible, clypeus, supra–clypeal area and scape, even when the female is almost entirely black. Flagellomeres usually shorter and wider than in females; white band on flagellum starting more apically than in females of the same species. Propodeal furrow usually slightly longer than in female. Propodeum smaller, less strongly convex. Spiracle of propodeum from short and elliptic to distinctly elongate, SLW 1.45–6.0. First metasomal segment slenderer, with T1LW 2.9–5.3, and less widened apically, with T1WW 1.4–2.35. T2–7 much slenderer than in females.

Comments.

Females of Stenarella are readily distinguishable from all other cryptine taxa by the very long ovipositor, at least 2.8× as long as the hind tibia, and the characteristic shape of the ovipositor tip, distinctly twisted and with the dorsal valve showing distinct teeth or hooks. Other Old World taxa with very long ovipositors are the Malagasy genus Rambites and some species of Tolonus, but these can be differentiated from Stenarella by the shape of the ovipositor tip and the other characters in the diagnosis.

Males of Stenarella, however, are harder to recognize due to the absence of the ovipositor. By following the remaining characters in the diagnosis, males would be most similar to males of Picardiella, which also have a long mandible with the dorsal tooth much longer than ventral one, and a rather small areolet. Males of Stenarella can be separated from those of Picardiella by having the spiracle placed approximately at posterior 0.42 of its length (versus 0.47) and the areolet smaller (APH 0.25–0.55, versus 0.8–1.25 in Picardiella). In addition, many species of Stenarella have the posterior transverse carina of propodeum absent or almost indistinct from propodeal striation, while in Picardiella this carina is always complete and distinct.

Distribution.

Widely distributed in the Old World, Australia, and introduced into North America. The known distribution records are expanded from 47 countries (Yu et al. 2016 and additions by Wahl & Green 2020 and Pham et al. 2024) to 75 after this study, in all continents except South America and Antarctica.

Key to the species of Stenarella

Females

1 Mesoscutum densely punctate or granulate (e.g., Figs 4E, 6E, 7E, 21E) 2
1 Mesoscutum mostly smooth and shiny, at most anterior part of median lobe more or less finely punctate, rarely lateral lobes with sparse punctures anteriorly (e.g., Figs 3D, 9E, 11D, 14E, 15F) 7
2 Mesoscutum mostly granulate, lateral lobes without punctures (Fig. 4E) 2. S. brykella Roman
2 Mesoscutum densely punctate or punctate reticulate (e.g., Figs 7E, 12D, 21E) 3
3 Mesosoma black, with well defined whitish marks (Figs 12D, E, 21D, E). Areolet usually open, rarely with trace of vein 3rs-m, vein 2rs-m obliterated (Figs 12C, 21C) 4
3 Mesosoma entirely black, black and orange, or entirely orange except sometimes whitish or yellow marks on dorsal lateral part of pronotum and subtegular ridge (Figs 6D, E, 7D, E, 19B, D). Areolet closed, sometimes vein 3rs-m unpigmented but present, vein 2rs-m well defined 5
4 Propodeum with one medial posterior large whitish mark; medially and posteriorly to anterior transverse carina coarsely transversely strigose on a smooth and shiny background (Fig. 1B) 18. S. victoriae (Cameron)
4 Propodeum with two large posterolateral whitish marks; medially and posteriorly to anterior transverse carina punctate reticulate or reticulate rugose (Fig. 1C) 9. S. insidiator (Smith)
5 Propodeum short, dorsal profile in lateral view broken into a short anterodorsal face and a longer sloped posterodorsal face (Fig. 7A, D); area basalis usually defined (Fig. 1D); wing with a subdistal dark hue at pterostigma level (Fig. 7A); vein 2m-cu interstital or postfurcal to vin 3rs-m (Fig. 7C) 5. S. domator (Poda)
5 Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped (Figs 6A, D, 19A, B); area basalis not defined (Fig. 1E, F); wings hyaline (Figs 6A, 19A); vein 2m-cu antefurcal to vein 3rs-m (Figs 6C, 19E) 6
6 Vertex finely granulate with very sparse setiferous punctures (Fig. 6E). Genal orbit only dorsally yellow (Fig. 6D). Subtegular ridge orange (Fig. 6D) 4. S. delicata sp. nov.
6 Vertex densely punctate on a smooth or finely granulate background (Fig. 19D). Genal orbit completely yellow (Fig. 19A, B). Subtegular ridge yellow (Fig. 19B) 16. S. tripartita (Brullé)
7 Mesosoma entirely black (Fig. 17A, D, E). Wing hyaline with distal half dark brown (Fig. 17A, C) 14. S. nigromaculata sp. nov.
7 Mesosoma black and orange or entirely orange (e.g., Figs 3C, D, 18B–D). Wing hyaline or evenly infuscate (e.g., Figs 3E, 11A) 8
8 Posterior area of propodeum transversely striate, striae regularly separated and reaching pleural sulcus (Fig. 1H–K) 9
8 Posterior area of propodeum mostly rugose, rugulose or irregularly strigose 12
9 Mesosoma light orange without black marks (Fig. 3A, C, D). Face and genal orbit whitish (Fig. 3B, C). Antenna with white band on f5–f14(15) 1. S. aurea sp. nov.
9 Mesosoma mostly black and orange (e.g., Figs 11B, 16D, 18B–D). Face (except orbits) and ventral part of genal orbit black (e.g., Figs 11B, C, 16B, D). Antenna with white band on f6–f11(12) or f6–f13 (e.g., Figs 11A, 16A) 10
10 Pronotum coarsely and longitudinally striated (Fig. 18B–D). Mesoscutum mostly orange or dark orange (Fig. 18E). Coxae dark brown to black (Fig. 18A). Fore coxa mostly dark brown or black with white dorand sal distal longitudinal stripe (Fig. 18B–D) 15. S. saaksjarvi sp. nov.
10 Pronotum mostly smooth or shallowly striated at pronotal depression (Figs 11B, 16D). Mesoscutum dark brown to black (Figs 11D, 16E). Fore coxa without white narrow distal stripe, sometimes distally widely whitish (Figs 11B, 16D) 11
11 Mesopleuron, metapleuron and propodeum dark orange to brownish orange (Fig. 11B). Hind coxa black (Fig. 11A). Antenna with white band on f6–f11(12) (Fig. 11A) 8. S. hopkinsi sp. nov.
11 Mesopleuron, metapleuron and propodeum light orange to orange (Fig. 16D). Hind coxa orange (Fig. 16A). Antenna with white band on f6–f13 (Fig. 16A) 13. S. nigriscuta sp. nov.
12 Mandible striated at basal half (Fig. 20C) 13
12 Mandible smooth or shallowly punctate at basal half (Fig. 10C) 15
13 Notaulus conspicuously wide with coarse spaced transverse keels (Fig. 20F). Clypeus and face in lateral view with concave profile. Areolet open (Fig. 20D). Propodeal spiracle short, ellipsoidal, SLW about 1.4 (Fig. 20E) 17. S. vannoorti sp. nov.
13 Notaulus narrow or moderately wide with very fine dense transverse keels (Figs 9E, 15F). Clypeus and face in lateral view with flat profile. Areolet closed by an unpigmented vein 3rs-m (Figs 9D, 15E). Propodeal spiracle elongated, SLW 2.6–3.0 (Figs 9C, 15C) 14
14 Posterior end of S1 placed approximately opposite to spiracle of T1 (Fig. 9C). Thyridium at most 1.5 times as long as wide (Fig. 9C) 6. S. favilla sp. nov.
14 Posterior end of S1 placed posteriorly to spiracle of T1 (Fig. 15D). Thyridium 2.25–2.4 times as long as wide (Fig. 15D) 12. S. natalina sp. nov.
15 Pronotum at most with short fine striae at pronotal depression, elsewhere smooth (Fig. 14D). Thyridium usually ≤2 times as long as wide 11. S. lissonota (Cameron)
15 Pronotum mostly with distinct and coarse longitudinal striae, only dorsal posterior part smooth (Figs 5C, 10E, 13D). Thyridium usually ≥2.3 times as long as wide (Fig. 10G) 16
16 Posterior end of S1 placed distinctively posterior to spiracle of T1 (Fig. 5A, F). Base of mandible dark orange to dark brown (Fig. 5B). T1 dark brown (Fig. 5A) 3. S. bugalana sp. nov.
16 Posterior end of S1 placed approximately opposite to or slightly posterior to spiracle of T1 (Figs 10A, 13A). Base of mandible with whitish spot (Figs 10B, C, 13B). T1 vividly orange (Figs 10A, 13A) 17
17 Posterior area of propodeum mainly strigose (Fig. 2E). Sternaulus shallow and sinuous, reaching posterior rim of mesopleuron (Fig. 13D). Tergites II–III mostly dark brown to black (Fig. 13A, F). Longitudinal carina of frons incomplete 10. S. katanga sp. nov.
17 Posterior area of propodeum mainly rugulose (Fig. 2F). Sternaulus deep evenly upcurved, reaching anterior 0.6 of mesopleuron (Fig. 10E). Tergites II–III mostly orange (Fig. 10A). Longitudinal carina of frons complete 7. S. fenestralis sp. nov.

Males

(Known only for S. lissonota, S. domator, S. insidiator, S. tripartita, S. victoriae and S. vannoorti)

1 Mesoscutum smooth and shiny (As in Figs 14E, 20E) 2
1 Mesoscutum densely punctate (As in Figs 7E, 12D, 19D, 21E) 3
2 Head with face, clypeus, basal half of mandible, and ocular orbits except at vertex, whitish (Fig. 22C). Propodeum mostly orange (Fig. 22C). Front and mid coxae white (Fig. 22C). Mandible shallowly punctate at basal half. Notaulus narrow with shallow, dense, and minute transverse keels. SLW about 2.3 11. S. lissonota (Cameron)
2 Head dark brown except white facial orbits narrowly (Fig. 22E). Propodeum black (Fig. 22E). All coxae dark brown to black (Fig. 22E). Mandible striated at basal half. Notaulus conspicuously wide with coarse relatively long spaced transverse keels. SLW about 1.45 17. S. vannoorti sp. nov.
3 Mesosoma black, with numerous well defined whitish marks (Fig. 22B, F). Areolet open (Fig. 22F) 4
3 Mesosoma entirely black, black and orange, or entirely orange except sometimes whitish marks on dorsal lateral part of pronotum and subtegular ridge (Fig. 22A, D). Areolet closed, sometimes vein 3rs-m unpigmented (As in Figs 7C, 19E) 5
4 Propodeum with one medial posterior large whitish mark (Fig. 22F), medially and posteriorly transversely striated, laterally strongly rugose punctate (As in Fig. 1B) 18. S. victoriae (Cameron)
4 Propodeum with two lateral posterior whitish marks (Fig. 22B), mainly strongly rugulose medially, reticulate rugose laterally (As in Fig. 1C) 9. S. insidiator (Smith)
5 Propodeum short, dorsal profile in lateral view broken into a short anterodorsal face and a longer sloped posterodorsal face (Fig. 22A); area basalis usually defined (As in Fig. 1D). Vein 2m-cu interstitial or postfurcal to vein 3rs-m (As in Fig. 7C) 5. S. domator (Poda)
5 Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped (Fig. 22D); area basalis not defined (As in Fig. 1E). Vein 2m-cu antefurcal to vein 3rs-m (As in Fig. 19E) 16. S. tripartita (Brullé)

Stenarella aurea sp. nov.

Figures 1H, 3, 23A, 28

Etymology.

From the Latin adjective aureus, -a, -um, meaning “golden”, in reference to the light orange, yellowish body color of this species.

Diagnosis.

Stenarella aurea sp. nov. can be distinguished from all other Stenarella species by the combination of the following characters: mesoscutum mostly smooth and shiny (Fig. 3D); posterior area of propodeum transversely striate, striae regularly separated and reaching pleural sulcus (Fig. 1H); face and eye orbits whitish (Fig. 3B, C); antenna with white band on f5–f14(15); mesosoma mostly light orange.

Description.

FEMALE. Fore wing 5.1–6.2 mm long. — Head (Fig. 3A–C): Mandible, clypeus and face moderately covered with very long setae. Mandible punctate, MLW 2.25–2.30. Malar space short, MSM 0.4–0.5. Clypeus smooth, CWL 2.8–3.0. Clypeus and face centrally mostly flat. Face with a small tubercle between antennal sockets, centrally weakly wrinkled, elsewhere smooth. Frons smooth and shiny, with incomplete faint longitudinal carina. Antenna with 28–34 flagellomeres. — Mesosoma (Figs 1H, 3A, C–E): Pronotum with distinct striae along central concave part, elsewhere smooth. Epomia moderately strong, reaching pronotal depression. Mesoscutum almost entirely smooth and shiny. Notaulus reaching 0.75 of mesoscutum with short transverse striae. Mesopleuron and metapleuron moderately setose. Mesopleuron dorsally longitudinally striate, ventrally finely rugulose. Epicnemial carina distinct, reaching 0.7 of distance to subtegular ridge. Sternaulus deep evenly upcurved, reaching anterior 0.6 of mesopleuron. Metapleuron strigose punctate. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum rugulose punctate. Propodeal spiracle elliptic, SLW 2.0–2.25. Area posterior of propodeum covered by fine, more or less uniformly spaced transverse striae. Posterior transverse carina of propodeum present but weakly discernible from other transverse striae, distance to anterior transverse carina 1.8 times the distance from anterior carina to anterior margin of propodeum. Areolet small, closed distally by an unpigmented vein 3rs-m, APH 0.4–0.45. Vein 2m-cu antefurcal to vein 3rs-m. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 2.2–2.3. — Metasoma (Fig. 3A, F): Posterior end of S1 placed distinctly posterior to spiracle of T1. T1LW 2.85–3.1; T1WW 2.25–2.3; T2LW 1.55–1.6; T2WW 2.15–2.2. Thyridium about 2.5 times as long as wide. T2 anteriorly granulate, gradually transitioning to punctate reticulate on a granulate background. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 10 teeth. OST 4.30–4.75. — Color (Figs 1H, 3): Head black: labrum, clypeus, mandibles except apex, mouthparts, face and eye orbits whitish; facial tubercle and scapus and pedicel ventrally orange. Antenna dark brown, f6–14 dorsally and f5 and f15 partially, white. Mesosoma mostly light orange; dorsal lateral part of pronotum, subtegular ridge, lateral parts of scutellum, and slightly posterior part of propodeum, yellowish; propodeum with a medial anterior dark brown spot. Legs mostly orange; hind trochantellus, proximal part of t1 and t5 entirely dark brown; distal part of t1 and t2–t4 whitish. Hind tibia something infuscate. Metasoma mostly orange: anterior part of T2–T3, T6–T8 dark brown. Posterior part of T6–T7 white; rarely fine posterior band on T1–T2 yellowish. Wing hyaline. — MALE. Unknown

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from: Angola, Cameroon, Central African Republic, Gabon, Ivory Coast and Sierra Leone (Fig. 23A).

Material examined.

11 ♀♀. Holotype. SIERRA LEONE • ♀; Freetown Peninsula, Freetown; III.1969; Denis Owen (USUC). Dry pinned, complete. — Paratypes. ANGOLA • 1 ♀; Salazar; Instituto de Investigação Agronomica de Angola; 9–15.III.1972; Southern African Exp. (NHMUK). CAMEROON • 1 ♀; Nkoemvom; XII.1979–I.1980; Ms. D. Jackson (NHMUK) • 1 ♀; same data except 16.III–4.V.1980 (NHMUK) • 2 ♀♀; same data except 30.III–19.IV.1980 (NHMUK) • 1 ♀; same data except X–XI.1980; Ms. D. Jackson (NHMUK). CENTRAL AFRICAN REPUBLIC • 1 ♀; Sangha–Mbaéré, Dzanga–Ndoki National Park, 38.6 km 173° S Lidjombo; 350m; 2º21.60’N, 16º09.20’E; 22–23.V.2001; S. van Noort; Malaise; lowland rainforest / SAM–HYM–P053035 / CAR01–M186 (SAMC). GABON • 1 ♀; 1919, Coll. J. De Gaulle (MNHN)• 1 ♀; Haut–Ogooué, N’Kogo, 1901; J. Bouyssou (MNHN). IVORY COAST • 1 ♀; Odienné, Ranch de Sipilou; 22.V.1973; V. Viltard Rec. (MNHN).

Stenarella brykella Roman, 1943

Figures 1A, 4, 23B, 28

Stenarella brykella Roman, 1943: 20. Lectotype ♀ (NHRS, photos examined).

Diagnosis.

Stenarella brykella can be easily distinguished from all other species of Stenarella by the mesoscutum mostly granulate without punctures on lateral lobes (Fig. 4E).

Description.

FEMALE. Fore wing 8.5–10.6 mm long. Head (Fig. 4A, B, D): Mandible, clypeus and face moderately covered with very long setae. Mandible punctate, MLW 1.95–2.15. Malar space moderately long, MSM 0.5–0.55. Clypeus sparsely punctate, CWL 2.85–2.9. Clypeus and face centrally mostly flat. Face centrally distinctly wrinkled, elsewhere smooth. Frons shiny, granulate and sparsely punctate, rugulose near vertex, with complete but anteriorly faint longitudinal carina. Antenna with 32 flagellomeres. — Mesosoma (Figs 1A, 4A, C–E): Pronotum entirely covered by uniform longitudinal striae. Epomia strong, reaching pronotal depression. Mesoscutum mostly granulate, anterior part with shallow dense punctures. Notaulus reaching 0.75–0.8 of mesoscutum with long and deep transverse striae. Scutellum rugulose punctate. Mesopleuron and metapleuron moderately setose. Mesopleuron mostly finely rugulose granulate, shortly longitudinally strigose along epicnemial carina. Epicnemial carina distinct, reaching 0.6 of distance to subtegular ridge. Sternaulus shallow, evenly upcurved, reaching anterior 0.6 of mesopleuron. Metapleuron strigose–granulate. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum rugulose punctate. Propodeal spiracle elliptic, SLW 1.85–2.0. Posterior area of propodeum finely striate medially, rugulose laterally. Posterior transverse carina of propodeum absent. Areolet small, closed distally by an unpigmented vein 3rs-m, APH 0.45–0.5. Vein 2m-cu antefurcal to vein 3rs-m. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 1.74–2.05. — Metasoma (Fig. 4A, F): Posterior end of S1 placed posterior to spiracle of T1. T1LW 2.65–2.7; T1WW 1.8–2.2; T2LW 1.1–1.55; T2WW 2.15–2.55. Thyridium about 2.0 times as long as wide. T2 mostly finely punctate reticulate, anteriorly granulate. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 9 teeth. OST 4.30–4.45. — Color (Figs 1A, 4): Head mostly black; labrum, mandible basally, clypeus laterally facial, frontal and genal orbits (sometimes medially interrupted), f6–f10 dorsally, and f5 and f11 partially, whitish. Mesosoma mostly orange: propleuron, pronotum anteriorly, mesopleuron ventrally 0.2, mesosternum and metasternum black; subtegular ridge and dorsal narrow band on pronotum whitish. Legs mostly dark brown; fore coxa dorsally and posteriorly with a white stripe; hind femur mostly orange; hind t2–t3 sometimes white. Wing hyaline. Metasoma dark brown to black. T5–T7 posteriorly whitish banded. — MALE. Unknown.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from Kenya (Fig. 23B).

Material examined.

3 ♀♀. Lectotype. [KENYA] • ♀; “Elgon, Bryk / M.E.H. / 10/8(?) / typus; Stenarella brykella n. sp., A. Roman det.; Lectotype Stenarella brykella Roman, Townes, 1972 / NHRS–HEVA, 000019821” (NHRS). —

Non-type material.

2 ♀♀. KENYA • 1 ♀; Karen, Nairobi, Keya 6000ft; 25–31.VIII.1971; C. van Someren (USUC) • 1 ♀; same data except 2000±m; III.1973 (USUC).

Stenarella bugalana sp. nov.

Figures 2D, 5, 23C

Etymology.

Named after Bugala Island, where the paratype was collected. From the Latinized adjective bugalanus, -a, -um.

Diagnosis.

Stenarella bugalana sp. nov. can be distinguished from all other Stenarella species by the combination of the following characters: mandible punctate at basal half; pronotum mostly with distinct and coarse longitudinal striae (Fig. 5C); mesoscutum mostly smooth and shiny (Fig. 5D); posterior area of propodeum transversally strigose (Fig. 2D); posterior end of S1 placed distinctively posterior to spiracle of T1 (Fig. 5A); thyridium about 2.5 times as long as wide; base of mandible dark orange to dark brown (Fig. 5B); T1 dark brown (Fig. 5A); wing hyaline or evenly infuscate (Fig. 5E).

Figure 2. 

Propodeum of Stenarella spp., females. A Stenarella natalina (paratype); B Stenarella favilla (paratype); C Stenarella lissonota; D Stenarella bugalana (holotype); E Stenarella katanga (paratype); F Stenarella fenestralis (paratype).

Figure 3. 

Morphology of Stenarella aurea sp. nov. female (holotype). A Habitus, lateral view; B head, frontal view; C head and mesosoma, lateral view; D mesosoma, dorsal view; E fore wing; F metasoma and ovipositor, lateral view.

Figure 4. 

Morphology of Stenarella brykella, female. A Habitus, lateral view (lectotype); B head, frontal view (lectotype); C fore wing; D head and mesosoma, lateral view; E mesoscutum and scutellum, dorsal view; F metasoma and ovipositor, lateral view.

Figure 5. 

Morphology of Stenarella bugalana sp. nov., female (holotype). A Habitus, lateral view; B head, frontal view; C head and mesosoma, lateral view; D head, mesoscutum and scutellum, dorsal view; E fore wing; F metasoma and ovipositor, lateral view.

Description.

FEMALE. Fore wing 8.0–8.7 mm long. — Head (Fig. 5A–D): Mandible, clypeus and face moderately setose. Mandible punctate, MLW 2.05–2.2. Malar space short, MSM 0.4–0.45. Clypeus sparsely punctate, CWL 2.6–2.75. Clypeus and face centrally mostly flat. Face centrally distinctly wrinkled, elsewhere sparsely punctate on a granulate background. Frons shiny, granulate and sparsely punctate, with interrupted longitudinal carina. Antenna with 32 flagellomeres. — Mesosoma (Figs 2D, 5A, C–E): Pronotum with distinct longitudinal striae, median dorsal part smooth. Epomia strong, reaching pronotal depression. Mesoscutum mostly smooth and shiny, except for anterior part of mid lobe with shallow dense punctures. Notaulus reaching 0.75 of mesoscutum, with short transverse striae. Scutellum rugulose. Mesopleuron and metapleuron moderately setose. Mesopleuron dorsally rugulose striate, ventrally finely rugulose. Epicnemial carina distinct, reaching 0.6 of distance to subtegular ridge. Sternaulus deep evenly upcurved, reaching anterior 0.7 of mesopleuron. Metapleuron rugulose strigose. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum finely rugulose punctate. Propodeal spiracle elliptic, SLW 2.2–2.5. Posterior area of propodeum transversally strigose. Posterior transverse carina of propodeum absent. Areolet small, closed distally by an unpigmented vein 3rs-m, APH 0.50–0.55. Vein 2m-cu antefurcal to vein 3rs-m. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 1.85–2.15. — Metasoma (Fig. 5A, F): Posterior end of S1 placed distinctively posterior to spiracle of T1. T1LW 3.0–3.1; T1WW 2.25–2.30; T2LW 1.4–1.45; T2WW 2.55–2.60. Thyridium about 2.5 times as long as wide. T2 mostly minutely punctate reticulate on a granulate background, small anterior dorsal portion granulate. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 7 teeth. OST 4.4–4.6. — Color (Figs 2D, 5): Head dark brown to black: labrum, facial and frontal orbits and genal orbit dorsally, whitish. Antenna with f6–f11(12 proximally) dorsally white. Mesosoma mostly dark orange: pronotal collar dark brown; propleuron, mesosternum and metasternum black. Fore and mid legs mostly dark orange with coxae, trochanter and trochantellus, dark brown, tarsi darkened or entirely dark brown. Hind leg dark brown. Wing hyaline or evenly infuscate. Metasoma mostly black; T1 mostly orange-brown; T1–T2 narrowly and T5–T7 extensively, posteriorly whitish. — Male. Unknown.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from Uganda (Fig. 23C).

Material examined.

2 ♀♀. Holotype. UGANDA • ♀; Kampala; 24.VII.1933; H. Hargreaves (NHMUK). Dry pinned. Distal flagellomeres of left antenna, tarsi of fore leg, hind left leg, and t5 of hind right leg missing. — Paratype. UGANDA • 1 ♀; Kalanga district, Bugala Island, Lake Victoria; 25.III.1912; G.D.H. Carpenter; Running over wooden table in hut (NHMUK).

Stenarella delicata sp. nov.

Figures 1F, 6, 23D, 28

Etymology.

From the Latin adjective delicatus, -a, -um, meaning “delicate”, referencing the fact that this is the smallest of the Stenarella species.

Diagnosis.

Stenarella delicata sp. nov. can be distinguished from all other Stenarella species by the combination of the following characters: vertex finely granulate with very sparse setiferous punctures (Fig. 6E); mesoscutum densely and shallowly punctate (Fig. 6E); propodeum relatively long, dorsal profile in lateral view evenly convex and sloped (Fig. 6D); areolet closed (Fig. 6C); vein 2m-cu antefurcal to vein 3rs-m (Fig. 6C); genal orbit only dorsally yellow cream (Fig. 6D); mesosoma mostly dark orange (Fig. 6D, E); wings hyaline (Fig. 6A, C).

Figure 6. 

Morphology of Stenarella delicata sp. nov., female (holotype). A Habitus, lateral view; B head, frontal view; C fore wing; D head and mesosoma, lateral view; E head, mesoscutum and scutellum, dorsal view; F metasoma and ovipositor, lateral view.

Description.

FEMALE. Fore wing 6.3–7.8 mm long. — Head (Fig. 6A, B, D, E): Mandible, clypeus and face moderately setose. Mandible punctate, MLW 2.3–2.5. Malar space short, MSM 0.50–0.55. Clypeus sparsely punctate, CWL 2.8–3.0. Clypeus and face centrally mostly flat. Face centrally weakly rugulose, elsewhere sparsely punctate on finely granulate background. Frons shiny, shallowly granulate and sparsely punctate, with complete but anteriorly faint longitudinal carina Vertex finely granulate with very sparse setiferous punctures. Antenna with 29–30 flagellomeres. — Mesosoma (Figs 1F, 6A, C–E): Pronotum with distinct longitudinal striae, anterodorsal corner smooth. Epomia strong, reaching pronotal depression. Mesoscutum densely and shallowly punctate. Notaulus reaching 0.65 of mesoscutum with minute transverse striae. Scutellum with sparse, large shallow punctures. Mesopleuron and metapleuron moderately setose. Mesopleuron dorsally rugulose–strigose, ventrally finely rugulose. Epicnemial carina weak but complete through subtegular ridge. Sternaulus shallow and sinuous, reaching posterior rim of mesopleuron. Metapleuron rugulose. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum rugulose. Propodeal spiracle elliptic, SLW 2.5–2.65. Posterior area of propodeum strongly rugulose. Posterior transverse carina of propodeum medially distinct as an M–shaped ridge, laterally absent, distance to anterior transverse carina 2 times the distance from anterior carina to anterior margin of propodeum. Areolet closed distally by an unpigmented vein 3rs-m, APH 0.45–0.50. Vein 2m-cu antefurcal to vein 3rs-m. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 2.0. — Metasoma (Fig. 6A, F): Posterior end of S1 placed approximately opposite to spiracle of T1. T1LW 2.3–2.4; T1WW 2.05–2.2; T2LW 0.95–1.15; T2WW 2.3–2.35. Thyridium about 2 times as long as wide. T2 mostly minutely punctate reticulate on a granulate background, small anterior dorsal portion granulate. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 9 teeth. OST 2.85–3.0. — Color (Figs 1F, 6): Head black: labrum, mandible except apex, mouthparts, facial and frontal orbits, genal orbit except ventral part and lateral spots on clypeus whitish; scape and pedicel yellowish ventrally. Antenna dark brown, f7–f10 dorsally white, f6 and f11 only partially white. Mesosoma mostly dark orange: propleuron and ventral part of pronotum dark brown; dorsal lateral rim of pronotum yellow. Legs mostly dark orange; fore femur and tibia clearer. Wing hyaline. Metasoma black. T1 brown; posterior bands on T1–T2 and posterior part of T5–T7 extensively, whitish. — MALE. Unknown.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from Nigeria and Uganda (Fig. 23D).

Material examined.

3 ♀♀. Holotype. UGANDA • ♀; Kampala; X.1965; Denis F. Owen (USUC). Dry pinned. Complete. — Paratypes. NIGERIA • 1 ♀, Oyo state, Ibadan; Oyo International Institute of Tropical Agriculture, 23–30.I.2007; bamboo thicket rainforest 230m; 48666, 3.8901 (USNM). UGANDA • 1 ♀; Wakison district, Entebbe; 11.IX.1909; C.A. Wiggins; No. 475 (OXUM).

Stenarella domator (Poda, 1761)

Figures 1D, 7, 8, 26, 28

Ichneumon domator Poda, 1761: 105. Type lost.

Ichneumon gladiator Scopoli, 1763: 283. Type lost. Synonymized by Gravenhorst (1829).

Ichneumon tarsosus Geoffroy in Fourcroy, 1785: 408. Type lost. Synonymized by Gravenhorst (1829).

Ichneumon insignitor Villers, 1789: 164. Type lost. Synonymized by Olivier (1792).

Ichneumon macrourus Gmelin, 1790: 2687. Type lost. Synonymized by Gravenhorst (1829).

Listrognathus transsylvanicus Kiss, 1924: 60. Holotype ♀ (TMA; not examined). Synonymized by Roman (1939).

Stenarella gladiator corsicator Aubert, 1969: 55. Holotype ♀ (MNHN; not examined)

Mesostenus cruentator Klug in Waltl, 1835: 80. Lectotype ♀ (MFNB, designated and synonymized by Horstmann (1989), examined).

Ichneumon ensator Thumberg, 1822: 259. Holotype ♀ (ZIU, not examined). Synonymized by Roman (1912).

Ichneumon domator var. ensator Roman, 1912.

Mesostenus juvenilis Tosquinet, 1896: 214. Holotype ♀ (IRSNB, not examined). Synonymized by Roman (1912).

Mesostenus ruficollis Rudow, 1882: 33. Type lost. Synonymized by Horstmann (1993).

Stenarella domator: Horstmann (1989).

Diagnosis.

Stenarella domator can be distinguished from all other Stenarella species by the combination of the following characters: mesoscutum densely and coarsely punctate (Fig. 7E); propodeum short, dorsal profile in lateral view broken into a short anterodorsal face and a longer sloped posterodorsal face (Figs 7D, 8); area basalis usually defined (Fig. 1D); areolet closed (Fig. 7C); vein 2m-cu interstital or postfurcal to vein 3rs-m (Fig. 7C); mesosoma entirely black, black and orange, or entirely orange except some whitish marks on dorsal lateral part of pronotum and subtegular ridge (Fig. 8).

Figure 7. 

Morphology of Stenarella domator domator female. A Habitus, lateral view; B head, frontal view; C fore wing; D head and mesosoma, lateral view; E head, mesoscutum and scutellum, dorsal view; F metasoma and ovipositor, lateral view.

Figure 8. 

Morphology of Stenarella domator subspecies, habitus of females, lateral view. A S. domator domator; B S. domator corsicator; C S. domator cruentator; D S. domator ensator.

Description.

FEMALE. Fore wing 5.8–10.4 mm long. — Head (Figs 7A, B, D, E, 8): Mandible, clypeus and face moderately covered with very long setae. Mandible distinctly striate on basal 0.6, MLW 2.0–2.2. Malar space moderately long, MSM 0.6–0.7. Clypeus sparsely punctate, CWL 3.2–3.3. Clypeus and face centrally mostly flat. Face, frons and vertex densely rugulose punctate on a granulate background, frons with only traces of longitudinal carina. Antenna with 30–35 flagellomeres. — Mesosoma (Figs 1D, 7A, C–E, 8): Pronotum dorsally densely rugulose punctate on a granulate background, ventrally longitudinally striate. Epomia long and strong reaching anterior margin of pronotum. Mesoscutum densely and coarsely punctate, punctures mostly coalescent. Notaulus relatively wide and deep, with strong transverse striae, reaching 0.75 of mesoscutum. Scutellum densely punctate, rugulose punctate laterally. Mesopleuron and metapleuron moderately setose. Mesopleuron mostly densely rugulose with sparse punctures and striae. Epicnemial carina weak, reaching 0.5 of distance to subtegular ridge. Sternaulus deep and sinuous, with strong transverse keels, reaching posterior rim of mesopleuron. Metapleuron coarsely rugulose–reticulate on a granulate background. Propodeum short, dorsal profile in lateral view broken into a short anterodorsal face and a longer sloped posterodorsal face. Anterior area of propodeum finely rugulose on a granulate background, with median longitudinal carina distinct. Propodeal spiracle elliptic, SLW 2.0–2.15. Posterior area of propodeum coarsely rugulose. Posterior transverse carina of propodeum distinct but sometimes medially interrupted, confluent with traces of median longitudinal carina, distance to anterior transverse carina 1.0 times the distance from anterior carina to anterior margin of propodeum. Areolet small, closed distally by an unpigmented vein 3rs-m, APH 0.4–0.45. Vein 2m-cu interstital or postfurcal to vein 3rs-m. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 2.2–2.5. — Metasoma (Figs 7A, F, 8): Posterior end of S1 placed anteriorly or approximately opposite to spiracle of T1. T1LW 2.15–2.3; T1WW 2.25–2.5; T2LW 0.95–1.1; T2WW 1.75–1.95. Thyridium about 1.25 times as long as wide. T2 punctate–reticulate on a finely granulate background, anteriorly only granulate. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 9 teeth. OST 4.6–5.65. — Color (Figs 1D, 7, 8): Variable in different subspecies as follows. • S. domator domator (Figs 1D, 7, 8A): Head mostly black: clypeus, mandibles partially, ocellus, usually central mark at face, facial and frontal orbits narrowly and median spot on genal orbit, orange. f7–f9 dorsally white; f6 and f10 partially white. Mesosoma black. All coxae, trochanters, and trochantelli partially, black; femora and tibiae orange; mid and hind tibiae distally and tarsi dark brown; hind t2–t4 white. Wing hyaline, with a darkened hue at pterostigma level. Metasoma black. • S. domator corsicator (Fig. 8B): As in domator domator except all legs black. • S. domator cruentator (Fig. 8C): As in domator domator but face and gena much more extensively orange, pronotum mostly, mesoscutum except a longitudinal short black stripe on mid lobe, scutellum laterally and dorsal half of mesopleuron, orange. Legs entirely dark brown to black, hind t2–t4 white. • S. domator ensator (Fig. 8D): Head orange; mandible teeth and vertex black. Mesosoma orange; propleuron, axillae, mesepisternum, metasternum, dorsal part of metapleuron and anterior part of propodeum, black, usually only the area spiracularis and area basalis black. Metasoma dark brown. Legs mostly dark brown, femora and tibiae tending to be dark orange. — MALE (Fig. 22A). Fore wing 6.2–8.0 mm long. — Head: MLW 1.9–2.1. Malar space moderately long, MSM 0.45–0.55. CWL 3.75–4.0. Antenna with 31–33 flagellomeres, tyloids on f12–f15. — Mesosoma: SLW 4.0–4.1. APH about 0.3.; HW1C 2.75–2.8. — Metasoma: T1LW 3.0–4.15; T1WW 1.55–2.0; T2LW 1.15–1.55; T2WW 1.9–2.1. — Color: Similar to female without white band in flagellum. Other characters as in female.

Comments.

This species is widely distributed across the Western Palearctic region, seeming to occur across all Europe as well as North Africa and in Central Asia as far as Iran and east Kazakhstan. At least in Western and Central Europe, it appears to be a common species, including in many anthropic areas such as wooden buildings where it likely searches for hosts that nest inside holes in the wood. The species counts thousands of records in public platforms such as GBIF, iNaturalist and Flickr (http://flickr.com), in stark contrast with the Afrotropical species of the genus for which the distribution is still poorly known. Wahl and Green (2020) showed that S. domator was also introduced in North America, and indeed, both iNaturalist and GBIF show reliable occurrences of the species in several localities (see “Distribution” below).

According to Horstmann (1989), there are four valid subspecies of S. domator, which mostly correspond to color morphs (see “Color” above) found at different latitudes. S. domator cruentator is found at the Iberian Peninsula (with the Pyrenees seeming to be the barrier between the populations of the subspecies and those of S. domator domator), and also in Iran (see photos in Heydari et al. 2021) and Armenia (https://www.inaturalist.org/observations/86050747), but not in the rest of Southern Europe (Italy, Greece), where S. domator domator is the resident subspecies. This leaves an open question of whether S. domator cruentator represents a distinct lineage or merely environmentally induced color variation. S. domator ensator is restricted to the Western part of North Africa, with records in Algeria and Morocco – Stenarella is apparently absent from the Eastern part of North Africa and the Eastern coast of the Mediterranean. Finally, S. domator corsicator is recorded only from Corsica, but its color differences relative to S. domator domator are so slight that it is dubious whether it can be considered a distinct population in any meaningful level.

As noted above, S. domator shows relevant color variation across its very broad geographical range, but we have only examined a small fraction of the very large number of specimens available in collections and analyzed molecular only from a few specimens. Hence, a more detailed investigation of genetic and phenotypic variation within S. domator, and its potential status as multiple specific entities, is beyond the scope of this work.

Hosts.

The electronic catalog Taxapad (Yu et al. 2016) records ten host species for S. domator; of these, four are aculeate Hymenoptera: Eumenes pomiformis (Fabricius) [Vespidae: Eumeninae], Sceliphron spirifex (Linnaeus) [Sphecidae], Diodontus tristris (Vander Linden) [Crabronidae: Pemphredoninae], and Andrena bisulcata (Morawitz) [Andrenidae]. Other wood–associated hosts are recorded as well, but Wahl and Green (2020) make the case that records of sawflies and beetles as hosts are probably due to observers overlooking aculeate nests that used pre-existing holes.

Distribution.

Palearctic, and indroduced in the Nearctic. Known records from Algeria, Austria, Belarus, Belgium, Bulgaria, Croatia, Cyprus, Czech Republic, DenmarkNR-G, EstoniaNR-G, Finland, France, GeorgiaNR-I, Germany, Greece, Hungary, Iran, Italy, KazakhstanNR-G-I, Latvia, Lithuania, LuxembourgNR-G, Morocco, Netherlands, Norway, Poland, Portugal, Romania, Russia, Slovakia, Spain, Sweden, Switzerland, Tunisia, Turkey, Ukraine and former Yugoslavia. Introduced in the United States (MinnesotaNR-I, New YorkNR-I, Ohio). The species has also been listed from the United Kingdom in Yu et al. (2016) based on a primary record for Staffordshire by Carr (1924), but his collection had specimens from southern Europe mixed with British material and hence these records are unreliable (Schwarz and Shaw 1998) (Fig. 26).

Material examined.

129 ♀♀; 10 ♂♂.

Stenarella domator corsicator (4 ♀♀): FRANCE • 1 ♀; Corsica; C. Villemant (MNHN) • 1 ♀; same locality; July 1994; Cl. Girard (MNHN) • 1 ♀; Corsica; Bonifacio; April 1999; Coll. Fcitoi (MNHN) • 1 ♀; Corsica; Porto Vecchio; May 1972; S. Kelner–Pillault (MNHN).

Stenarella domator cruentator (8 ♀♀): SPAIN • 1 ♀; Andalusien; Wal., S.; “Lectotypus Cryptus cruentator Klug, Horstmann, 1984” (MFNB) • 1 ♀; Banyeres de Mariola (Alicante); 8.VI.2014; leg. Bordera (CEUA) • 1 ♀; El Ventorrillo, Cercedilla (Madrid); 3.VIII.1962; Docavo and Llopis leg. (CEUA) • 1 ♀; Lusitania, Spanien; 10829; “Lectoparatypus; Cryptus cruentator Klug, Hortmann, 1984” (MFNB) • 1 ♀; Mallorca; Keifel, S.; 10829 (MFNB) • 1 ♀; P. N. de Cabañeros (Ciudad Real); Abierto–Raña del Pocico; 25.IV/8.V.2004; TM1 (CEUA) • 1 ♀; same locality; Fresneda–Gargantilla; 15.IV/7.V.2004; TM1 (CEUA) • 1 ♀; Villafranca del Cid (Castellón); 5.VIII.1989 (CEUA).

Stenarella domator domator (102 ♀♀, 9♂♂): FRANCE • 2 ♀♀; Allier, Brout–Vernet; August 1928; H. du Buysson (MNHN) • 2 ♀♀; Argentat–sur–Dordogne; June 1987; Coll. J. Vachal (MNHN) • 1 ♂; Argentat–sur–Dordogne; May 1989; Coll. J. Vachal (MNHN) • 1 ♀: Aube; July 1992; P. Viette Coll. (MNHN) • 1 ♀; Bethonvilliers; August 1984; C. Thirion (MNHN) • 1 ♀; Bézu–saint–Eloi; 1888; Ch. Brongniart (MNHN) • 1 ♀; Bormes–les–Mimosas; Coll. Lichtenstein (MNHN) • 1 ♂; Boulogne–sur–Mer; 1877; Coll. Giraud (MNHN) • 1 ♀; Brissac; 1902; R. du Buysson (MNHN) • 1 ♂; Brout–Vernet; June 1990; H. du Buysson (MNHN) • 1 ♀; Brout–Vernet; July 1989; H. du Buysson (MNHN) • 2 ♀♀; Champfleury; May 1966/67; Coll. O. Sichel (MNHN) • 1 ♀; Champfleury; May 1960; Coll. O. Sichel (MNHN) • 1 ♀; same locality; July 1966; Coll. O. Sichel (MNHN) • 1♀; same locality; 1867; Coll. O. Sichel (MNHN) • 1 ♀; Digne–les–Bains; 1923; A. Seyrig (MNHN) • 1 ♀; Dun–sur–Grandy; September 1970; Coll. Giraud (MNHN) • 2 ♀♀; Fontainebleau; 1897; A. Finot; (MNHN) • 1 ♂; same locality; July 1979; Finot (MNHN) • 1 ♀; Fouesnant; 1927; Alain Hémon (MNHN) • 1 ♀; Galié; 1887; Coll. O. Sichel (MNHN) • 1 ♀; Gavarnie–Gèdre; August 1916; F. Picard Coll. (MNHN) • 1 ♀; Guern; 1908; M. Pic (MNHN) • 3 ♀; Foret de la Hardt; June 1926; A. Seyrig (MNHN) • 1 ♀; La Celle–Dunoise; 1917; Alluaud (MNHN) • 4 ♀♀; La Môle; August 1984; Coll. A. Adamski (MNHN) • 1 ♀; Lardy; May 1995; F. Picard Coll. (MNHN) • 1 ♀; Le Mesnil; August 1989 (MNHN) • 1 ♀; Le Vésinet; 1867; Coll. O. Sichel (MNHN) • 4 ♀♀; Levant Island; August 1974; H. Le Grand (MNHN) • 1 ♀; Lozère; H. Maneval • 1 ♀; Maison–Lafitte; 1919; Coll. J. de Gaulle (MNHN) • 1 ♀; Maraye–en–Othe; 1927; A. Hemon Coll.; (MNHN) • 1 ♀; France méridionale; 1867; O. Sichel (MNHN) • 2 ♀♀; Montpellier; July 1918; Jean Lichtenstein (MNHN) • 1 ♀; same locality; August 1922; J. Lichtenstein (MNHN) • 1 ♀; same locality; 1962; O. Sichel Coll. (MNHN) • 1 ♀; [France] Montp. May–1911; ex Coll Pfankuch; Mesostenus gladiator Grav. (MFNB) • 1 ♀; Moutiers; 1867; O. Sichel Coll. (MNHN) • 1 ♀; Narbonnes; 1919; J.de Gaulle Coll. (MNHN) • 1 ♀; Orry–la–Ville; June 1938; J. Bourgogne (MNHN) • 1 ♀; Plouharnel; 1919; J. de Gaulle Coll. (MNHN) • 1 ♀; Pont–de–l’Arche; July 1903; J. de Gaule Coll. (MNHN) • 1 ♀; Pontarlier; June 1960; O. Sichel Coll. (MNHN) • 2 ♀♀; Rennes; René Oberthur (MNHN) • 1 ♀; Royan; 1915; J. Pérez Coll. (MNHN) • 1 ♀; Saint Mandé; June 1967; O. Sichel Coll. (MNHN) • 1 ♀; Saint–Sendoux; 1989; H. du Buysson (MNHN) • 2 ♀♀; Sainte Baume; L. Chopard (MNHN) • 1 ♀; Samoens; A. Villiers (MNHN) • 1 ♀; Sèvres; June 1922; A.Seyrig; (MNHN) • 2 ♀♀; Thann; May 1921; A. Seyrig (MNHN) • 1 ♀; Toulouse; H. du Buysson (MNHN) • 1 ♂; Tuchan; 1887; Coll. O. Sichel (MNHN) • 1 ♀; Vachères; July 1976; Casevitz–Weulersse rec. (MNHN) • 1 ♀; Varenne; 1867; O. Sichel Coll. (MNHN) • 1 ♂; Vaux; June 1926; Coll. J. de Gaulle (MNHN) • 1 ♀; Verdun; J. de Gaulle Coll. (MNHN). GERMANY • 4♀♀; Berlin; J.P.E. Fr. Stein S.; 24182 (MFNB) • 1 ♀; Berlin; Brieselang; Gerstaecker; S.; Mesostenus gladiator Grav. Ariesel (MFNB) • 2 ♀♀; Döl. Heide; 18.V.1918; Halle A. S.; Mesostenus gladiator Grav. det. Haupt; 1926 (MFNB) • 1 ♀; Furstenberg (MNHN) •1 ♀; Jena; Thür.; 19.VI.1900; Mesostenus gladiator Grav. Friese det 1900 (MFNB) • 4 ♀♀, 1 ♂; Schwerin (MFNB) • 1 ♀; Schwerin; 21.VI.1881 (MFNB) • 2 ♀♀; Stobben; Elbing; (Noctzendorf) (MFNB) • 2♀♀; Thüringen; O. Smiedecknecht; S.; 25927; Mesostenus gladiator (MFNB) • 1 ♂; Weissburg; Bern Oberl. Mesostenus gladiator Grav. Coll. Schmiedeknecht (MFNB). GREECE • 1 ♀; Crete; Chania; April 1971; J. F. Aubert (MNHN) • 1 ♀; Salonique; 1916; Dr Rivet (MNHN) • 2 ♀♀; Thalos; May 1924; A. Seyrig (MNHN). [ITALY] • 1 ♀; Bozen; 13.VII.1908 (MFNB) • 2 ♀♀; Ragusa; Rudolf; Mesostenus domator (Poda) Krieg. det. (MFNB). [LATVIA] • 1 ♀; Kurland, Wezkukkul; 12.VI.1916; 1741 (MFNB) • 1 ♀; Kurland, Wezkukkul; 17.VII.1916; Bischoff, S.G. (MFNB). RUSSIA • 1 ♀; [Russia] Rossitten; 23.VI.1912; Mesostenus gladiator Grav. (MFNB) • 1 ♀; Saint Petersburg; 1867; O. Sichel Coll. (MNHN). SWITZERLAND • 1 ♀; Bern; 1887; O. Sichel Coll. (MNHN) • 1 ♂; Burgdof (MNHN) • 1 ♀; Charmey; 1919; J. de Gaulle Coll. (MNHN) • 1 ♀; Jura; A. August 1924; Seyrig (MNHN) • 1 ♀; Lugano; May 1963; O. Sichel Coll. (MNHN) • 1 ♀; Yverdon–les–Bains; 1867; O. Sichel Coll. (MNHN).

Stenarella domator ensator (15 ♀♀, 1♂): ALGERIA • 11 ♀♀, 2 ♂; Annaba; 1963; O. Sichel (MNHN) • 1 ♀; Berrouaghia; Coll. F. Picard; 1939 (MNHN) • 1♀; Moyen Atlas; Tameghilt, 1600–2100 m; June 1929; F. Le Cerf (MNHN). MOROCCO • 1 ♀; Tangier; 1914; H. Marmottan (MNHN). TUNISIA • 1 ♀; Menzel Bourguiba; A. Jung Coll. (MNHN).

Stenarella favilla sp. nov.

Figures 2B, 9, 23E

Etymology.

From the Latin favilla, -ae, meaning “embers”, in reference to the irregular pattern of black and ferruginous resembling glowing embers. Noun in apposition.

Diagnosis.

Stenarella favilla sp. nov. can be distinguished from all other Stenarella species by the combination of the following characters: clypeus and face in lateral view with flat profile; mandible striated at basal half (Fig. 9B); mesoscutum mostly smooth and shiny (Fig. 9E); notaulus relatively narrow and deep with minute dense transverse striae (Fig. 9E); posterior area of propodeum mostly rugulose, medially strigose (Fig. 2B); propodeal spiracle elongated, SLW about 3.0 (Fig. 9C); areolet closed (Fig. 9D); posterior end of S1 placed approximately opposite to spiracle of T1 (Figs 9C); thyridium at most 1.5 times as long as wide (Fig. 9C); mesosoma black and orange (Fig. 9C, E)

Figure 9. 

Morphology of Stenarella favilla sp. nov., female. A Habitus, lateral view (holotype); B head, frontal view (holotype); C head, mesosoma and T1–T2, lateral view (holotype); D fore wing (paratype); E: head, mesoscutum and scutellum, lateral and dorsal view respectively (holotype); F metasoma and ovipositor, lateral view (holotype).

Description.

FEMALE. Fore wing 10.7–11.3 mm long. — Head (Fig. 9A–C, E): Mandible moderately covered with very long setae. Clypeus and face sparsely setose. Mandible striate on basal 0.6, striation stronger at mid length MLW 2.0–2.2. Malar space short, MSM 0.35–045. Clypeus smooth and shiny, CWL 2.55–2.7. Clypeus and face with flat profile from lateral view. Face centrally slightly rugulose, elsewhere shallowly granulate–punctate. Frons mostly smooth and shiny sparsely punctate with weak but complete longitudinal carina. Vertex smooth and shiny with few punctures on ocellar area. Antenna incomplete. — Mesosoma (Figs 2B, 9A, C–E): Pronotum mostly smooth and shiny, with short and very shallow longitudinal striae along anterior depression. Epomia weak, reaching pronotal depression. Mesoscutum almost entirely smooth and shiny, except for distinct set of longitudinal striae between posterior end of notauli. Notaulus narrow and deep with minute dense transverse striae, reaching about 0.7 of mesoscutum. Scutellum densely rugulose punctate. Mesopleuron and metapleuron moderately setose. Mesopleuron mostly densely punctate reticulate, with sparse striae along epicnemial carina, dorsal central part tending to be smooth with more sparse punctures. Epicnemial carina distinct, reaching 0.8 of distance to subtegular ridge. Sternaulus deep and narrow, evenly upcurved, reaching anterior 0.7 of mesopleuron. Metapleuron finely rugulose striate. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum coarsely rugulose punctate. Propodeal spiracle elliptic, SLW 3.0. Posterior area of propodeum mostly rugulose, medially strigose. Posterior transverse carina of propodeum absent. Areolet closed distally by an unpigmented vein 3rs-m, APH 0.40–0.45. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 3.0. — Metasoma (Fig. 9A, C, F): Posterior end of S1 placed approximately opposite to spiracle of T1. Thyridium 1.3–1.4 times as long as wide. T2 minutely punctate reticulate on a granulate background except for small granulate portion on anterior part. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 9 teeth. OST 4.50–4.95.— Color (Figs 2B, 9): Head mostly black: base of mandible, clypeus, facial, frontal and genal orbits dorsally, yellow cream. Antenna dark brown with whitish dorsal band. Mesosoma mostly orange: propleuron, ventral part of pronotum, epicnemium and mesopleuron, mesosternum, metasternum and ventral part of metapleuron, dark brown to black; dorsal posterior part of propodeum lighter. Fore and mid legs mostly light orange, coxae dark brown, femora and tibiae ventrally and all tarsi somewhat darker, trochanters whitish. Hind leg mostly dark brown, trochanter whitish ventrally. Wing hyaline. Metasoma mostly black: posterior narrow bands on T1–T2, and dorsal posterior part of T6–T7 whitish. — Male. Unknown.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from South Africa (Fig. 23E).

Material examined.

2 ♀♀. Holotype. SOUTH AFRICA • ♀; KwalaZulu–Natal, Umhlanga; 22.I.1960; G. C. Clark (NHMUK). Dry pinned. Distal flagellomeres from f8 of left antenna, and from f13 of right antenna, hind left leg and mid right leg from coxae, t2–t5 of mid left leg, all tarsi of hind right leg and distal part of fore wing, missing. — Paratype. SOUTH AFRICA • 1 ♀; KwaZulu–Natal; 1875; W.W Saunders (OXUM).

Stenarella fenestralis sp. nov.

Figures 2F, 10, 23F, 28

Etymology.

From the Latinized adjective fenestralis, -e, meaning “window”, in reference to the amusing fact that one of the specimens was collected at a window.

Diagnosis.

Stenarella fenestralis sp. nov. can be distinguished from all other Stenarella species by the combination of the following characters: mandible punctate at basal half (Fig. 10C); longitudinal carina of frons complete; pronotum mostly with distinct longitudinal striae (Fig. 10E); mesoscutum mostly smooth and shiny (Fig. 10F); sternaulus deep and narrow, evenly upcurved, reaching anterior 0.6 of mesopleuron (Fig. 10E); posterior area of propodeum rugulose (Fig. 2F); thyridium 3.45–3.75 times as long as wide; posterior end of S1 placed approximately opposite to spiracle of T1 (Fig. 10A, G); base of mandible whitish (Fig. 10B, C); mesosoma mostly orange (Fig. 10E, F); tergites II–III mostly orange (Fig. 10A, G).

Figure 10. 

Morphology of Stenarella fenestralis sp. nov., female (holotype). A Habitus, lateral view; B head, frontal view; C mandible; D fore wing; E head and mesosoma, lateral view; F head, mesoscutum and scutellum, dorsal view; G metasoma and ovipositor, lateral view.

Description.

FEMALE. Fore wing 7.2–12 mm long. — Head (Fig. 10A–C, E, F): Mandible, clypeus and face densely covered with very long setae. Mandible punctate at base, MLW 2.4–2.6. Malar space short, MSM 0.35–0.45. Clypeus smooth, CWL 2.85–3.0. Clypeus and face centrally mostly flat. Face centrally weakly rugulose, elsewhere shallowly granulate. Frons shiny, mostly smooth, granulate laterally, sparsely punctate, with strong and complete longitudinal carina. Vertex shiny, shallowly granulate laterally with fine and dense punctures. Antenna with 31–33 flagellomeres. — Mesosoma (Figs 2F, 10A, D–F): Pronotum covered by uniform longitudinal strong and coarse striae, the striae fading dorsally leaving small dorsal smooth area. Epomia strong, reaching pronotal depression. Mesoscutum mostly smooth and shiny, anterior part of mid and lateral lobes with shallow dense punctures, area between posterior end of notauli with longitudinal striae Notaulus relatively wide and deep with transverse striae, reaching about 0.8 of mesoscutum. Scutellum densely and coarsely punctate. Mesopleuron and metapleuron densely setose. Mesopleuron almost entirely minutely rugulose with some striae dorsally. Epicnemial carina distinct, reaching 0.9 of distance to subtegular ridge. Sternaulus deep and narrow, evenly upcurved, reaching anterior 0.6 of mesopleuron. Metapleuron rugulose. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum coarsely rugulose reticulate. Propodeal spiracle elliptic, SLW 2.35–2.50. Posterior area of propodeum rugulose. Posterior transverse carina of propodeum absent. Areolet very small, closed distally by an unpigmented vein 3rs-m, APH 0.25–0.35. Vein 2m-cu antefurcal to vein 3rs-m. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 2.1–2.2. — Metasoma (Fig. 10A, G): Posterior end of S1 placed approximately opposite to spiracle of T1. T1LW 2.70–2.75; T1WW 2.0–2.35; T2LW 1.1–1.2; T2WW 2.3–2.45. Thyridium 3.45–3.75 times as long as wide. T2 minutely punctate reticulate on a granulate background, anterior dorsal part only granulate. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 6–7 teeth. OST 3.5–4.3. — Color (Figs 2F, 10): Head mostly black: base of mandibles, mouth parts, clypeus, facial, frontal and genal orbits, whitish. Antenna mostly dark brown, basally lighter ventrally, f7–f10 dorsally and f6, f11 partially, white. Mesosoma mostly orange: propleuron, ventral part of pronotum anteriorly, mesosternum, metasternum and metapleural carina, black; dorsal lateral rim of pronotum and subtegular ridge white cram. Fore and mid legs, mostly light orange; coxae and dorsal part of trochanters dark brown, ventral part of trochanters whitish. Hind leg dark brown. Wing hyaline. Metasoma mostly black: T1–T2 anterior part of T3 and rarely T4 mostly orange, posterior margin of T1–T2 yellow, posterior part of T5–T7 white. — Male. Unknown.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from Angola, Malawi, Zambia and Zimbabwe (Fig. 23F).

Material examined.

21 ♀♀. Holotype. ZIMBABWE (“Rhodesia”) • ♀; Harare (“Salisbury”), Chishawasha; III.1979; A. Watsham (NHMUK). Dry pinned. Complete. — Paratypes. ANGOLA • 1 ♀; Huíla province, Tundavala; 8–10 miles NW. Sa da Bandeira; 27–29.III.1972; Southern African Exp. B.M. 1972–1 (NHMUK). MALAWI(“Nyasaland”) • 1 ♀; Mangochi (“Fort Johnston”); 12 Km E of Ft. Johnston; 3280 ft; 1.I.1929; rather thick bush; W.A. Lamborn (OXUM). ZAMBIA • 1 ♀; 15km. E. Lusaka; III.1980; R.A. Beaver (NHMUK). ZIMBABWE (“South Rhodesia”) • 1 ♀; Harare (“Salisbury”), Chishawasha; II–III.1978; A. Watsham (NHMUK) • 1 ♀; same data except X.1978 • 5 ♀♀; same data except XII.1978 • 1 ♀; same data except III.1979 • 1 ♀; same date except III.1980 • 1 ♀; same data except IX.1980 • 1 ♀; same data except XI.1980 • 2 ♀♀; same data except XII.1980 • 1 ♀; same data except I.1981 • 1 ♀; same data except II.1981 • 1 ♀; same data except III.1981 • 1 ♀; same data except “At window Ent. Laboratory Salisbury Dist. Dept. Agric. S. Rhodesia”; 27.IX.1951; M.C. Mossop; “Pres by Com Inst Ent B M 1953–623” / Com. Inst. Ent. Coll. NO. 1318 / Parasilsila sp. G.J. Kerroch det. 1953 / Accession No. 6709 (all of them NHMUK).

Stenarella hopkinsi sp. nov.

Figures 1J, 11, 24A, 28

Etymology.

Named after hymenopterist Tapani Hopkins (ZMUT), who collected the type specimens of this species during many months of tireless fieldwork in Uganda. Noun in genitive case.

Diagnosis.

Stenarella hopkinsi sp. nov. can be distinguished from all other Stenarella species by the combination of the following characters: pronotum and mesoscutum smooth and shiny (Fig. 11B, D); anterior area of propodeum coarsely rugose punctate; posterior area of propodeum transversely striate, striae regularly separated and reaching pleural sulcus (Fig. 1J); face and facial orbits black (Fig. 11C); antenna with white band on f6–f11(12) (Fig. 11A); mesoscutum black (Fig. 11D); mesopleuron, metapleuron and propodeum dark orange to brownish orange (Fig. 11B); hind coxa black (Fig. 11A).

Figure 11. 

Morphology of Stenarella hopkinsi sp. nov., female (holotype). A Habitus, lateral view; B head and mesosoma, lateral view; C head, frontal view; D head, mesoscutum and scutellum, dorsal view; E fore wing; F metasoma and ovipositor, lateral view.

Description.

FEMALE. Fore wing about 13.7mm long. — Head (Fig. A–D): Mandible covered with very dense and long setae. Clypeus and face moderately setose. Mandible punctate or slightly striate at basal half, MLW 2.0–2.2. Malar space short, MSM 0.40–0.45. Clypeus smooth, CWL 2.75–3.0. Clypeus and face flat. Face centrally distinctly rugulose, elsewhere smooth or slightly granulated. Frons smooth and shiny, sometimes slightly rugulose behind antennal sockets, with weak interrupted longitudinal carina. Antenna with 36 flagellomeres. — Mesosoma (Figs 1J, 11A, B, D, E): Pronotum almost entirely smooth and shiny, punctate at dorsal medial part, weakly longitudinally striate at ventral part of pronotal depression. Epomia strong, reaching pronotal depression. Mesoscutum almost entirely smooth and shiny with coarse convergent rugae between posterior end of notauli. Notaulus reaching 0.85 of mesoscutum with fine transverse striae. Scutellum mostly punctate with central part smooth. Mesopleuron and metapleuron moderately setose. Mesopleuron very densely and finely punctate, dorsally with weak longitudinal rugae. Epicnemial carina distinct, reaching 0.8 of distance to subtegular ridge. Sternaulus moderately deep and sinuous, almost reaching posterior rim of mesopleuron. Metapleuron densely punctate rugulose. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum coarsely rugose punctate. Propodeal spiracle elliptic, SLW 2.65–2.8. Posterior area of propodeum covered by more or less uniformly spaced transverse striae. Posterior transverse carina of propodeum indiscernible from transverse striae. Areolet very small, closed distally by an unpigmented vein 3rs-m, APH 0.30–0.35. Vein 2m-cu antefurcal to vein 3rs-m. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 2.7–2.85. — Metasoma (Fig. A, F): Posterior end of S1 placed posteriorly to spiracle of T1. T1LW 2.7–3.1; T1WW 2.3–2.45; T2LW 1.20–1.4; T2WW 2.1–2.15. Thyridium about 1.8–2.2 times as long as wide. T2 mostly minutely densely punctate reticulate on a finely granulate background, anterior part only finely granulate. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 10 teeth. OST 3.4–3.60. — Color (Fig. 1J, 11): Head mostly black: labrum, clypeus, mandible except distal part, mouth parts, widely frontal orbits, narrow spot on dorsal part of genal orbit and f6–11(12) dorsally, whitish. Mesosoma mostly dark orange to brownish orange: propleuron, pronotum, mesoscutum, anterior part and widely median part of propodeum, dark brown to black; small spot on dorsal posterior part of pronotum, subtegular ridge, lateral parts of scutellum and posterior medial wide spot on propodeum, whitish. Legs mostly orange; fore and mid coxae and trochanters partially dark brown and whitish, tarsi dark brown; hind leg dark brown; t1 posteriorly, t2–t4, and distal part of t5, yellowish. Wing hyaline. Metasoma dark brown; T1–T2 narrowly and T6–T7 extensively, posteriorly whitish. — Male. Unknown.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from Uganda and Democratic Republic of the Congo (Fig. 24A).

Material examined.

2 ♀♀. Holotype. UGANDA • ♀; Kabarole district, Kibale National Park, Fort Portal; 6–21.XI.2014; T. Hopkins, http://mus.utu.fi/ZMUT.11366/HILLT1 141121 (ZMUT). Dry pinned. Right hind leg removed for DNA sequencing. — Paratype. DEMOCRATIC REPUBLIC OF THE CONGO • 1 ♀; North Kivu, Irangi, Station Rech. Scient., Fluß Luoho; 800–1000m; 3–6.III.1984; Burmeister, Fuchs, Kühbandner (ZSMC).

Stenarella insidiator (Smith, 1859)

Figures 1C, 12, 22B, 27, 28

Mesostenus insidiator Smith 1859: 60. Holotype ♀ (OXUM, examined).

Orientostenaraeus chinensis Uchida, 1930: 321. Holotype ♀ (Hokkaido University, not examined). Synonymized by Townes et al. (1961).

Stenarella insidiator: Townes et al. (1961).

Diagnosis.

Stenarella insidiator can be distinguished from all other Stenarella species by the combination of the following characters: mesoscutum densely and coarsely punctate on a granulate background (Fig. 12D); areolet usually open, rarely with trace of vein 3rs-m, vein 2rs-m obliterated (Fig. 12C); posterior area of propodeum medially punctate reticulate, laterally reticulate rugose (Fig. 1C); mesosoma black, with well-defined whitish marks (Figs 12A, D, E, 22B), propodeum with two large posterolateral whitish marks (Figs 1C, 12E, 22B).

Figure 12. 

Morphology of Stenarella insidiator, female. A Habitus, lateral view; B head, frontal view; C fore wing; D mesoscutum and scutellum, dorsal view; E head, mesosoma and T1, lateral view; F metasoma and ovipositor, lateral view.

Description.

FEMALE. Fore wing 7.5–16.3 mm long. — Head (Fig. 12A, B, E): Mandible, clypeus and face moderately setose. Mandible finely punctate granulate, MLW 1.85–2.0. Malar space short, MSM 0.45–0.5. Clypeus sparsely punctate, CWL 3.0–3.25. Clypeus and face centrally mostly flat. Face centrally weakly wrinkled, elsewhere smooth. Frons medially rugulose punctate, laterally granulate, with weak but complete longitudinal carina. Antenna with 33–36 flagellomeres. — Mesosoma (Figs 1C, 12A, C–E): Pronotum dorsal margin with shallow dense punctures, ventrally with distinct longitudinal striae, otherwise smooth and shiny. Epomia long but weak, reaching pronotal depression. Mesoscutum densely and coarsely punctate on a granulate background, punctures coalescent. Notaulus reaching 0.90 of mesoscutum, with small transverse striae. Scutellum densely punctate. Mesopleuron and metapleuron moderately setose. Mesopleuron dorsally rugulose striate, ventrally rugulose. Epicnemial carina weak, reaching 0.5 of distance to subtegular ridge. Sternaulus shallow and sinuous, reaching posterior rim of mesopleuron. Metapleuron dorsally coarsely and densely punctate reticulate, ventrally rugulose striate. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum densely punctate. Propodeal spiracle elliptic, SLW 3.0–3.2. Posterior area of propodeum medially punctate reticulate, laterally reticulate rugose. Posterior transverse carina of propodeum absent. Areolet small usually open distally, if close, vein 3rs-m unpigmented, APH 0.45–0.5. vein usually 2rs-m obliterated. Vein 2m-cu interstitial to vein 3rs-m. Hind wing vein Cua distinctly longer than cross vein cu-a, HW1C 1.85–2.3. — Metasoma (Fig. 12A, F): Posterior end of S1 placed approximately opposite to spiracle of T1. T1LW 2.9–3.1; T1WW 2.05–2.3; T2LW 0.95–1.4; T2WW 2.1–2.3. Thyridium about 1.2 times as long as wide. T2 punctate reticulate except for small granulate part on anterior part. Dorsal valve of ovipositor with 9 teeth. Ventral valve with 11 teeth. OST 2.6–5.2. — Color (Figs 1C, 12): Head mostly black: labrum, clypeus, face, mandibles except distal part and eye orbits, except dorsal part of genal orbit, whitish. Antenna with f5(6)–f15(17) white. Mesosoma mostly black; dorsal margin and usually a spot ventrally on pronotum, subtegular ridge, large central spot on mesopleuron, dorsal division of metapleuron, central spot on mesoscutum, tegula, scutellum and postscutellum and two large posterior lateral marks on propodeum, yellow cream. Legs mostly orange; fore and mid coxae and trochanters, hind coxa dorsally and ventrally, sometimes also femur and tibia mostly, and hind tarsi, yellow cream; hind coxa mostly, hind trochanter and trochantellus, distal part of femur and base and distal part of tibia, dark brown. Wing hyaline. Metasoma mostly dark brown T1–T8 posteriorly yellow cream banded. — MALE (Fig. 22B). Fore wing 7.5–9.5 mm long. — Head. MSM 0.35–0.45. Clypeus sparsely punctate, CWL 2.2–2.4. Antenna with 33–35 flagellomeres; tyloids on t12–t17(18). — Mesosoma. SLW 3.2–4.5. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 1.75–2.1. — Metasoma. Posterior end of S1 placed approximately opposite to spiracle of T1 or slightly posterior. T1LW 3.5–3.9; T1WW 1.9–2.0; T2LW 1.7–2.1; T2WW 1.8–1.9. — Color. As in female. Antenna with f6–f18 or f7–f17 white, first one and last one, partially. white. Legs lighter than in female. Other characters as in female.

Comments.

There are two well defined groups of specimens: specimens from Borneo have a smaller body size (fore wing 6.1–8.8 mm) and shorter ovipositor (OST 2.6–3.2), while populations elsewhere are larger (fore wing 9.1–13.0 mm) and have longer ovipositors (OST 4.0–5.2). However, no other structural differences were observed that could justify the split of S. insidiator into two species at the present. Further analyses using molecular data for a broader geographic sampling will help to determine whether the population from Borneo represents a distinct species.

Hosts.

Sceliphron madraspatanum Klug, 1801 (Hymenoptera, Sphecidae); Orancistrocerus aterrimus (de Saussure, 1852) (Hymenoptera; Vespidae; Eumeninae).

Distribution.

Oriental. Known records from BruneiNR, Cambodia, China (Liaoninh, Sichuan, Yunnan), IndiaNR, Indonesia, MalaysiaNR, MyanmarNR, Philippines, Taiwan, ThailandNR, Vietnam (Fig. 27).

Material examined.

18 ♀♀, 2 ♂♂. Holotype. INDONESIA • ♀; Sulawesi, Makasar (OXUM). — Non-type material. BRUNEI • 1 ♀; Kuala Belalong Field Studies Centre; 115°7′E 4°34′N; 18.V.1991; N. Mawdsley; Malaise; BMNH(E): 1991–173 (NHMUK) • 1 ♀, same data except V.1991 (NHMUK). CHINA • 2 ♀♀; Szechuen, Mt. Omei; 11,000 ft; 21.VII.1935; D.C. Graham (USNM). INDIA • 1 ♀; Bikham / Claude Morley coll B.M. 1952–159 / “B.M. 1967 under Silsila spionata”/ Silsila longicantus karl/ Co–type B.M. (NHMUK) • 1 ♀; Meghalaya; Khasia Hills; Assam Athimus K. / Mesostenus longicauda/Munich. INDONESIA • 1 ♀; S.B. Heppner coll., Royal Ent. Soc. London Poject Wallace Exped. 1985 (FSCA) • 1 ♀; East Kalimantan, Balikpapan. Mentawir river; 50m; X.1950; A.M.R. Wegner (USUC). MALAYSIA • 1 ♀; Negri Sembilan; Pasoh Forest Reserve, forest; 17.XII.1978; P. & M.Becker (USUC) • 1 ♂, same data except 6.X.1979 (USUC) • 1 ♀, 1 ♂; Sarawak; Gunung Mulu National Park; “4th div. Gn. Mulu”; VI–VII.1978; H. Vallack (NHMUK) • 1 ♀; same data except 58m; Deer Cave trail, nr. jct. with small forest trail; 4.025417, 114.824683; 17.II–19.III.2012; D.C.Darling, Malaise; ROM_OSU 308613; glydant; limestone/seq (ROM) • 1 ♀; same date except Kuching; 11.XI.1999; Dyak, coll. Pres. 1900 by R. Shelford (OXUM). MYANMAR (“Burma”) • 1 ♀; Kachin state, Kachin hills, Mali Hka valley; 2000 ft; 3.XII.1930; F. Kingdon Ward, B.M. 1936–91 (NHMUK). PHILIPPINES • 1 ♀; Island Samar, Baker coll. (USNM) • 1 ♀; Oriental Mindoro; S. Luis Calapan, 150 ft; 15.IV.1954, H.M. & D. Townes / Homotype Orientostenaraeus chinensis Uch. H.K. Townes ’54 (USUC). TAIWAN • 1 ♀; Taipei, Fu–Shan; 10.IX.2001; J. & L. Stange (FSCA). THAILAND • 1 ♀; Chiang Mai; Huai Nam Dang NP Helipad; 19˚18.33’N 98˚36.289’E; 20–27.III.2008; Malaise trap; Anuchart & Thawatchai leg; T5635 (UKICU).

Stenarella katanga sp. nov.

Figures 2E, 13, 24B

Etymology.

Named after the province of Haut–Katanga, where the holotype was collected. Noun in apposition.

Diagnosis.

Stenarella katanga sp. nov. can be distinguished from all other Stenarella species by the combination of the following characters: mandible shallowly punctate at basal half; longitudinal carina of frons incomplete; pronotum mostly with distinct longitudinal striae (Fig. 13D); mesoscutum mostly smooth and shiny (Fig. 13C); sternaulus shallow and sinuous, reaching posterior rim of mesopleuron (Fig. 13D); posterior area of propodeum mostly strigose (Fig. 2E); thyridium 2.3–2.5 times as long as wide (Fig. 13A); posterior end of S1 placed approximately opposite to or slightly posterior to spiracle of T1 (Fig. 13A); base of mandible whitish (Fig. 13B); mesosoma mostly orange (Fig. 13A, D); T1 vividly orange, T2-T3 mostly dark brown to black (Fig. 13A, F).

Figure 13. 

Morphology of Stenarella katanga sp. nov., female. A Habitus, lateral view (holotype); B head, frontal view (paratype); C head, mesoscutum and scutellum, dorsal view (holotype); D head and mesosoma, lateral view (holotype); E fore wing (paratype); F metasoma and ovipositor, lateral view (holotype).

Description.

FEMALE. Fore wing 7.7–10.8 mm long. — Head (Fig. 13A–D): Mandible, clypeus and face densely covered with very long setae. Mandible punctate at base, MLW 2.25–2.3. Malar space short, MSM 0.45–0.50. Clypeus smooth, CWL 3.0–3.15. Clypeus and face centrally mostly flat. Face centrally weakly rugulose, elsewhere shallowly granulate. Frons shiny, granulate and sparsely punctate, with interrupted longitudinal carina. Vertex shiny, shallowly granulate with very fine and dense punctures. Antenna with 35 flagellomeres. — Mesosoma (Figs 2E, 13A, C–E): Pronotum covered by uniform longitudinal strong striae, the striae fading dorsally leaving small dorsal smooth area. Epomia strong, reaching pronotal depression. Mesoscutum mostly smooth and shiny, anterior part of mid and lateral lobes with shallow dense punctures. Area between posterior end of notauli with longitudinal striae. Notaulus moderately wide and deep with dense transverse striae, reaching about 0.7 of mesoscutum. Scutellum densely punctate. Mesopleuron and metapleuron densely setose. Mesopleuron almost entirely minutely rugulose with some longitudinal striae dorsally. Epicnemial carina distinct, reaching 0.9 of distance to subtegular ridge. Sternaulus shallow and wide, sinuous, reaching posterior rim of mesopleuron. Metapleuron rugulose. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum coarsely rugulose reticulate. Propodeal spiracle elliptic, SLW 2.0–2.3. Posterior area of propodeum mostly strigose, laterally tending to be rugulose. Posterior transverse carina of propodeum indiscernible from transverse striae. Areolet small, closed distally by a pigmented vein 3rs-m, APH 0.40. Vein 2m-cu antefurcal to vein 3rs-m. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 2.0–2.3. — Metasoma (Fig. 13A, F): Posterior end of S1 placed slightly posterior to spiracle of T1. T1LW 2.65–3.15; T1WW 1.9–2.0; T2LW 1.05–1.3; T2WW 2.45–2.7. Thyridium 2.3–2.5 times as long as wide. T2 minutely punctate reticulate on a granulate background, anterior dorsal part only granulate. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 7 teeth. OST 4.55–4.65. — Color (Figs 2E, 13): Head mostly black; base of mandibles facial, frontal and genal orbits dorsally, whitish. Antenna mostly dark brown, f6–f10(11) dorsally white. Mesosoma mostly orange: propleuron, ventral part of pronotum anteriorly, mesosternum, metasternum and metapleural carina, black; dorsal lateral rim of pronotum yellow. Fore and mid legs, mostly light orange; coxae and dorsal part of trochanters dark brown, ventral part of trochanters whitish. Hind leg dark brown. Wing hyaline. Metasoma mostly black: T1 orange, posterior dorsal spot on TI, posterior band on T2 and T5–T7 widely, white. — Male. Unknown.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from Angola and Democratic Republic of the Congo (Fig. 24B).

Material examined.

2 ♀♀. Holotype. DEMOCRATIC REPUBLIC OF THE CONGO • ♀; Haut–Katanga, Kundelungu Mountains; Malaise; “Stenarella sp. 7 Tow. 1972” (AEIC). Dry pinned. Right antenna missing. — Paratype. ANGOLA • 1 ♀; Benguela, Ganda (ZSMC).

Stenarella lissonota (Cameron, 1906)

Figures 2C, 14, 22C, 24C, 28

Mesostenus lightfooti Cameron, 1906: 147. Lectotype ♀ (SAMC; designated by Townes and Townes, 1973; not examined). Synonymized by Townes and Townes (1973).

Mesostenus levifrons Cameron, 1906: 148. Holotype ♀ (SAMC; not examined). Synonymized by Townes and Townes (1973).

Mesostenus lissonotus Cameron, 1906: 149. Holotype ♀ (SAMC; not examined). Lectotype ♀ (SAMC; designated by Townes and Townes (1973); not examined).

Mesotenus albilabris Cameron, 1911: 177. Holotype ♀ (SAMC; not examined). Synonymized by Townes and Townes (1973).

Stenaraeus (Mesostenus) lissonotus var. nigripes Morley, 1916: 377. Holotype ♀ (SAMC; not examined). Synonymized by Townes and Townes (1973).

Stenaraeus lightfooti: Morley (1916).

Stenaraeus laevifrons (!): Morley (1926). Incorrect subsequent spelling of Mesostenus levifrons Cameron, 1906.

Stenarella lissonotus: Townes and Townes, 1973.

Diagnosis.

Stenarella lissonota can be distinguished from all other Stenarella species by the combination of the following characters: mandible smooth or shallowly punctate at basal half; pronotum only with short striae at pronotal depression, elsewhere smooth (Fig. 14E, 22C); mesoscutum mostly smooth and shiny (Fig. 14E); notaulus narrow with shallow, dense, and minute transverse keels (Fig. 14E); posterior area of propodeum transversely strigose, tending to be rugulose laterally (Fig. 2C); thyridium about 2.0 times as long as wide; mesosoma mostly orange (Figs 14D, E, 22C). Additionally, males can be distinguished by head mostly whitish and by front and mid coxae whitish (Fig. 22C).

Figure 14. 

Morphology of Stenarella lissonota, female. A Habitus, lateral view (paratype); B head, frontal view; C fore wing; D head and mesosoma, lateral view; E head, mesoscutum and scutellum, dorsal view; F metasoma and ovipositor, lateral view (paratype).

Description.

FEMALE. Fore wing 7.5–9.2 mm long. — Head (Fig. 14A, B, D, E): Mandible, clypeus and face moderately setose. Mandible mostly sparsely punctate, MLW 2.1–2.3. Malar space short, MSM 0.35–0.45. Clypeus smooth, CWL 2.9–3.0. Clypeus and face centrally mostly flat. Face centrally weakly wrinkled, elsewhere smooth. Frons shiny, granulate and wrinkled medially, with interrupted longitudinal carina. Antenna with 34 flagellomeres. — Mesosoma (Figs 2C, 14A, C–E): Pronotum with distinct striae along anterior depression, elsewhere smooth. Epomia moderately strong, reaching pronotal depression. Mesoscutum mostly smooth and shiny, except anterior part of mid lobe with shallow dense punctures. Notaulus reaching 0.70 of mesoscutum with short transverse striae. Scutellum with sparse, large shallow punctures. Mesopleuron and metapleuron moderately setose. Mesopleuron dorsally rugulose–striate, ventrally finely rugulose. Epicnemial carina distinct, reaching 0.7 of distance to subtegular ridge. Sternaulus shallow and sinuous, reaching posterior rim of mesopleuron. Metapleuron rugulose strigose. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum rugulose punctate. Propodeal spiracle elliptic, SLW 2.0. Posterior area of propodeum transversely strigose, tending to be rugulose laterally. Posterior transverse carina of propodeum present but weakly discernible from other transverse striae, distance to anterior transverse carina 1.5 times the distance from anterior carina to anterior margin of propodeum Areolet small, closed distally by an unpigmented vein 3rs-m, APH 0.45–0.5. Vein 2m-cu antefurcal to vein 3rs-m. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 2.0–2.4. — Metasoma (Fig. 14A, F): Posterior end of S1 placed posteriorly to spiracle of T1. T1LW 3.1–3.25; T1WW 1.7–2.0; T2LW 1.3–1.45; T2WW 2.25–2.4. Thyridium about 2 times as long as wide. T2 punctate reticulate on a granulate background, anterior part granulate. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 10 teeth. OST 4.1–5.1. — Color (Figs 2C, 14): Head mostly black: clypeus, mandibles except teeth, and facial orbits as a hue, orange; f6–f12 dorsally white f5, f13 partially. Mesosoma mostly orange; propleuron and anterior part of pronotum dark brown. Wing hyaline. Metasoma mostly black: T1 dark brown, T1–T2 and T5–T8 posteriorly whitish. — MALE (Fig. 22C). Fore wing about 6.9 mm long. — Head. Mandible moderately punctate, MLW 2.4. Malar space moderately long, MSM 0.5. CWL 2.6. Antenna with 31 flagellomeres; tyloids on f13–f16. — Mesosoma. Pronotum with very weak and short striae along anterior depression, elsewhere smooth. Anterior area of propodeum rugulose punctate laterally, shallowly punctate medially. SLW about 2.3. Posterior area of propodeum finely and shallowly transversely strigose granulate medially, tending to be rugulose laterally. Areolet very small, APH 0.35. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C about 1.8. — Metasoma. Posterior end of S1 placed far posteriorly to spiracle of T1. T1LW 4.3; T1WW 1.4; T2LW 2.8; T2WW 1.8. Thyridium about 2.7 times as long as wide. — Color. Head mostly whitish; mandible teeth, frons and vertex except eye orbits, posterior dorsal part of gena and occiput dark brown to black; f10–f15 dorsally white, f16 partially. Mesosoma mostly orange; dorsal part of pronotum, subtegular ridge, speculum and sternum, yellowish; metapleuron ventrally dark brown. Front and mid coxae and trochanters, hind trochanter and hind t2–t4, whitish. Hind leg mostly dark brown. Wing slightly infuscate. Metasoma mostly dark brown to black; T1–T3 and T6–T8 posteriorly whitish. Other characters as in female.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from South Africa (Fig. 24C).

Material examined.

4 ♀♀, 1 ♂. SOUTH AFRICA • 1 ♀; Eastern Cape, Grahamstown; III.1972; Fred Gess (USUC) • 1 ♂; same data except KwaZulu–Natal; Hluhluwe Game Reserve; 13.XI.1970; H. & M. Townes (USUC) • 1 ♀; same data except Karkloof; 15.III.1942; Nat. Museum S.R. Millar; “South African museum ex national museum Bulawayo 1981” / SAM–HYM–P005 147 (SAMC) • 1 ♀, same data except Mfongosi; W. E. Jones; SAM–HYM–P000733 (SAMC) • 1 ♀; same data except Mpumalanga (“Transvaal”); Pilgrims Rest, L. Schunke / SAM–HYM–P000732 (SAMC).

Stenarella natalina sp. nov.

Figures 2A, 15, 24D, 28

Etymology.

Named after the KwaZulu-Natal province, where one of the paratypes was collected. From the Latinized adjective natalinus, -a, -um.

Diagnosis.

Stenarella natalina sp. nov. can be distinguished from all other Stenarella species by the combination of the following characters: clypeus and face in lateral view with flat profile; mandible striated at basal half; mesoscutum smooth and shiny (Fig. 15F); notaulus narrow with very fine dense transverse keels (Fig. 15F); posterior area of propodeum finely rugulose strigose medially, strongly rugulose laterally (Fig. 2A); propodeal spiracle elongated, SLW 2.6–2.75 (Fig. 15C); areolet closed (Fig. 15E); posterior end of S1 placed posteriorly to spiracle of T1 (Fig. 15D); thyridium 2.25–2.4 times as long as wide (Fig. 15D); mesosoma mostly dark orange (Fig. 15C); wing hyaline or evenly infuscate (Fig. 15A).

Figure 15. 

Morphology of Stenarella natalina sp. nov., female. A Habitus, lateral view (holotype); B head, frontal view (paratype); C head and mesosoma, lateral view (holotype); D first and second metasomal segment, lateral view (holotype); E fore wing (paratype); F head, mesoscutum and scutellum, dorsal view (paratype); G metasoma and ovipositor, lateral view (holotype).

Description.

FEMALE. Fore wing 8.0–11.3 mm long. — Head (Fig. 15A–C, F): Mandible, clypeus and face moderately setose. Mandible striated, MLW 2.3–2.5. Malar space short, MSM 0.35–0.40. Clypeus smooth, CWL 2.6–2.9. Clypeus and face centrally mostly flat. Face centrally distinctly rugulose, elsewhere smooth and shiny. Frons sparsely punctate with interrupted longitudinal carina. Vertex smooth and shiny. Antenna with 32 flagellomeres (Missing). — Mesosoma (Figs 2A, 15A, C, E, F): Pronotum mostly smooth, with short longitudinal striae along pronotal depression. Epomia strong, reaching pronotal depression. Mesoscutum almost entirely smooth and shiny. Notaulus reaching 0.70 of mesoscutum, with minute transverse striae. Scutellum with sparse, large shallow lateral punctures. Mesopleuron and metapleuron moderately setose. Mesopleuron mostly rugulose punctate, with short irregular striae on anterodorsal corner and bordering posterior part of epicnemium. Epicnemial carina weak (better seen in ventrolateral view), reaching 0.6 of distance to subtegular ridge. Sternaulus shallow and sinuous, reaching posterior rim of mesopleuron. Metapleuron rugulose punctate. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum coarsely rugose punctate. Propodeal spiracle elliptic, SLW 2.6–2.75. Posterior area of propodeum finely rugulose strigose medially, strongly rugulose laterally. Posterior transverse carina of propodeum absent. Areolet small, closed distally by an unpigmented vein 3rs-m, APH 0.4–0.45. Vein 2m-cu antefurcal to vein 3rs-m. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 2.15–2.25. — Metasoma (Fig. 15A, D, G): Posterior end of S1 placed posteriorly to spiracle of T1. T1LW 2.55–2.75; T1WW 2.15–2.3; T2LW 1.25–1.45; T2WW 2.05–2.2. Thyridium 2.25–2.4 times as long as wide. T2 mostly punctate reticulate on a finely granulate background, anterior part finely granulate. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 9 teeth. OST 4.50–4.9. — Color (Fig. 2A, 15): Head black; labrum clypeus, mandibles except apex, mouthparts, spot on malar space, facial and frontal orbits and genal orbit dorsally, whitish; f6–f12 dorsally white. Mesosoma dark orange; propleuron ventrally, pronotum anteroventrally mesosternum and metasternum, black. Fore and mid legs mostly brown; coxae dark brown; trochanters and partially trochantelli, whitish. Hind leg mostly dark brown; t2–t3 whitish. Wing hyaline. Metasoma black. T1–T2 with whitish mark on about 0.2 of their length, slightly visible on T3; T6–T7 dorsally whitish. — MALE. Unknown.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from South Africa and Tanzania (Fig. 24D).

Material examined.

3 ♀♀. Holotype. SOUTH AFRICA • ♀ Bethel; W. Capland Beste; 26368; Mesostenus tripartitus var. det. Brulle (MFNB). Dry pinned. Both antennae broken at first third; right hind tibia and tarsus missing. — Paratypes. SOUTH AFRICA • 1 ♀; Natal (=KwaZulu–Natal); Purchased 1875; W.W. Saunders Coll. (OXUM). TANZANIA • 1 ♀; Lindi Region, Matumbi Highlands, Mwengei; 1050 ft; –8.3463889, 38.96; 25.XI.1989; R.B. Hynd collection / BMNH(E)1998–129 (NHMUK).

Stenarella nigriscuta sp. nov.

Figures 1K, 16, 24E, 28

Etymology.

From the Latin adjective niger, -gra, -grum, meaning “black”, and the noun scutum, -i, in reference to the entirely black mesoscutum in this species.

Diagnosis.

Stenarella nigriscuta sp. nov. can be distinguished from all other Stenarella species by the combination of the following characters: pronotum and mesoscutum mostly smooth (Fig. 16D, E); posterior area of propodeum transversely striate, striae regularly separated and reaching pleural sulcus (Fig. 1K); face black (Fig. 16B); antenna with white band on f6–f13 (Fig. 16A); mesoscutum dark brown to black (Fig. 16E); mesopleuron, metapleuron and propodeum light orange to orange (Figs 1K, 16A, D); hind coxa orange (Fig. 16A); wing hyaline or evenly infuscate (Fig. 16A).

Figure 16. 

Morphology of Stenarella nigriscuta sp. nov., female. A Habitus, lateral view (holotype); B head, frontal view (paratype); C fore wing (paratype); D head and mesosoma, lateral view (holotype); E head, mesoscutum and scutellum, dorsal view (holotype); F metasoma and ovipositor, lateral view (holotype).

Description.

FEMALE. Fore wing 11.3–12.3 mm long. — Head (Fig. 16 A, B, D, E): Mandible covered with very dense and long setae, clypeus and face moderately setose. Mandible punctate, MLW 2.1–2.55. Malar space short, MSM 0.35–0.45. Clypeus smooth, CWL 2.5–2.65. Clypeus and face mostly flat. Face centrally distinctly rugulose, or rugulose granulate, elsewhere smooth. Frons smooth and shiny, with weak interrupted longitudinal carina. Antenna with 34 flagellomeres. — Mesosoma (Figs 1K, 16A, C–E): Pronotum almost entirely smooth and shiny. Epomia strong, reaching pronotal depression. Mesoscutum almost entirely smooth and shiny. Notaulus reaching 0.55–0.75 of mesoscutum with fine transverse striae. Scutellum mostly smooth, with shallow lateral punctures. Mesopleuron and metapleuron moderately setose. Mesopleuron mostly moderately punctate, dorsally weakly longitudinally striate, slightly rugulose punctate along epicnemial carina. Epicnemial carina distinct, reaching 0.7 of distance to subtegular ridge. Sternaulus shallow and sinuous, reaching posterior rim of mesopleuron. Metapleuron densely punctate reticulate. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum rugose punctate sometimes medially weakly transversely strigose. Propodeal spiracle elliptic, SLW 2.35–2.6. Posterior area of propodeum covered by fine, more or less uniformly spaced transverse striae. Posterior transverse carina of propodeum indiscernible from transverse striae. Areolet small, closed distally by an unpigmented vein 3rs-m, APH 0.35–0.45. Vein 2m-cu antefurcal to vein 3rs-m. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 2.0–3.0. — Metasoma (Fig. 16A, F): Posterior end of S1 placed posteriorly to spiracle of T1. T1LW 3.0–3.55; T1WW 1.85–2.0; T2LW 1.25–1.45; T2WW 2.25–2.4. Thyridium 2.0–2.5 times as long as wide. T2 mostly minutely punctate reticulate on a finely granulate background, anterior part finely granulate. Dorsal valve of ovipositor with 6 teeth. Ventral valve with 9–11 teeth. OST 3.75–4.45. — Color (Figs 1K, 16): Head mostly black: labrum, clypeus, mandible except distal part, mouth parts, usually spot on malar space, facial and frontal orbits, dorsal part of genal orbit and f6–13 dorsally, whitish. Mesosoma mostly light orange to orange: mesoscutum black; propleuron, pronotum mostly, spot on medial anterior part of propodeum, dark brown; dorsal lateral part of pronotum, subtegular ridge, lateral parts of scutellum and usually posterior medial spot on propodeum, whitish. Fore and mid leg orange, sometimes fore coxa and trochanter whitish. Hind leg mostly dark brown; coxa, and sometimes ventral part of femur orange; t1 posteriorly, t2–t4, and distal part of t5, whitish. Wing hyaline. Metasoma dark brown; T1–T2 narrowly and T6–T7 extensively posteriorly whitish. — MALE. Unknown.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from Cameroon, Nigeria and Republic of the Congo (Fig. 24E).

Material examined.

6 ♀♀. Holotype. CAMEROON • ♀; Nkoemvom; 13.VII–24.VIII.1980; Ms. D. Jackson (NHMUK). Dry pinned. Distal flagellomeres from f29 of right antenna missing. — Paratypes. CAMEROON • 2 ♀♀; same data as holotype (NHMUK) • 1 ♀; same data except 25.IX–19.XI.1979 (NHMUK). NIGERIA • 1 ♀; Oni River; 70 miles. E of Lagos; 15.V.1910; W.A. Lamborn/1911–422 (NHMUK). REUBLIC OF THE CONGO • 1 ♀; Brazzaville, Djoumouna; 22.X.1968; M. Grjebin Coll. (MNHN).

Stenarella nigromaculata sp. nov.

Figures 1G, 17, 24F, 28

Etymology.

From the Latin adjective niger, -gra, -grum, meaning “black”, and the noun macula, -ae, meaning “spot, stain”, in reference to the dark spots at the fore wing, which are unique of this species within the genus.

Diagnosis.

Stenarella nigromaculata sp. nov. can be distinguished from all other Stenarella species by the combination of the following characters: mesoscutum mostly smooth and shiny (Fig. 17E); mesosoma entirely black (Fig. 17A, D, E); wing hyaline with distal half dark brown (Fig. 17A, C).

Figure 17. 

Morphology of Stenarella nigromaculata sp. nov., female (holotype). A Habitus, lateral view; B head, frontal view; C fore wing; D head and mesosoma, lateral view; E head, mesoscutum and scutellum, dorsal view; F metasoma and ovipositor, lateral view.

Description.

FEMALE. Fore wing 11.1 mm long. — Head (Fig. 17A, B, D, E): Mandible, clypeus and face densely covered with very long setae. Mandible punctate, MLW 1.95. Malar space short, MSM 0.45. Clypeus with sparse but distinct punctures, CWL 2.9. Clypeus and face centrally mostly flat. Face coarsely rugose punctate. Frons sparsely punctate, with complete longitudinal carina. Vertex rugose punctate between posterior ocelli. Antenna with 36 flagellomeres. — Mesosoma (Figs 1G, 17A, C–E): Pronotum mostly with distinct longitudinal striae, dorsal posterior part smooth and shiny. Epomia strong, reaching pronotal depression. Mesoscutum almost entirely smooth and shiny. Notaulus reaching 0.65 of mesoscutum with distinct transverse striae. Scutellum rugose reticulate. Mesopleuron and metapleuron moderately setose. Mesopleuron dorsally with sinuous striae, ventrally mostly rugulose. Epicnemial carina weak (better seen in ventrolateral view), reaching 0.6 of distance to subtegular ridge. Sternaulus shallow evenly upcurved, reaching anterior 0.6 of mesopleuron. Metapleuron rugulose striate. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum coarsely rugulose reticulate. Propodeal spiracle elliptic, SLW 2.5. Posterior area of propodeum covered by moderately coarse, unevenly spaced transverse striae. Posterior transverse carina of propodeum absent. Areolet small, closed distally by an unpigmented vein 3rs-m, APH 0.50. Vein 2m-cu antefurcal to vein 3rs-m. Hind wing vein Cua longer than crossvein cu-a, HW1C 3.0. — Metasoma (Fig. 17A, F): Posterior end of S1 placed approximately opposite to spiracle of T1. T1LW 3.1; T1WW 1.85; T2LW 1.3; T2WW 2.25. Thyridium about 1.25 times as long as wide. T2 with shallow and moderately sparse punctures over finely granulate background. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 7 teeth. OST 3.55. — Color (Figs 1G, 17): Head mostly black; labrum, base of mandible, facial and frontal orbits, genal orbit dorsally, f6–f10 and proximally f11, white. Mesosoma black; a spot on posterior medial part of propodeum yellowish. Fore leg mostly pale orange; coxa, trochanter femur ventrally and distal tarsi, brown to dark brown. Mid leg mostly dark brown, femur dorsally orange. Hind leg mostly dark brown to black; base of femur narrowly orange; t2–t3 whitish. Wing hyaline with distal half dark brown. Metasoma black; T1–T3 narrowly banded posteriorly and T6–T8 posterolaterally, whitish. — MALE. Unknown.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from Central African Republic (Fig. 24F).

Material examined.

1 ♀. Holotype. CENTRAL AFRICAN REPUBLIC • ♀; Prefecture Sangha–Mbaéré; Dzanga–Ndoki National Park; 38.6km 173º S. Lidjombo, 2º21.60’N, 16º09.20’E; 350m; 26–27.V.2001; S. van Noort Leg; Malaise trap, lowland rainforest, CAR01–M231 (SAMC). Dry pinned. Distal flagellomeres from f29 of right antenna missing; mid and hind right legs removed for DNA sequencing.

Stenarella saaksjarvi sp. nov.

Figures 1I, 18, 25A, 28

Etymology.

Named after hymenopterist Ilari Sääksjärvi (ZMUT), as homage to his distinguished entomological career with significant contributions to the systematics and ecology of Ichneumonidae. Noun in genitive case.

Diagnosis.

Stenarella saaksjarvi sp. nov. can be distinguished from all other Stenarella species by the combination of the following characters: pronotum longitudinally and coarsely striated (Fig. 18B–D); mesoscutum mostly smooth and shiny (Fig. 18E); posterior area of propodeum transversely striate, striae regularly separated and reaching pleural sulcus (Fig. 1I); face (except orbits) and ventral part of genal orbit black (Fig. 18B–D, G); mesosoma mostly black or black and orange (Fig. 18A–E); fore coxae dark brown or black with white dorsal and distal longitudinal whitish stripe (Fig. 18B–D).

Figure 18. 

Morphology of Stenarella saaksjarvi sp. nov., female. A Habitus, lateral view (holotype); BD head and mesosoma, colour variation, lateral view; E head, mesoscutum and scutellum, dorsal view (holotype); F fore wing (holotype); G head, frontal view (holotype); H metasoma and ovipositor, lateral view (holotype).

Description.

FEMALE. Fore wing 5.8–11.3 mm long. — Head (Fig. 18A–E, G): Mandible moderately covered with very long setae. Clypeus and face sparsely setose. Mandible punctate at base, MLW 2.10–2.35. Malar space short, MSM 0.30–035. Clypeus sparsely punctate, CWL 2.75–2.80. Clypeus and face centrally mostly flat. Face centrally slightly rugulose, elsewhere smooth. Frons sparsely punctate, with interrupted longitudinal carina. Vertex mostly smooth and shiny with sparse punctures. Antenna with 30–36 flagellomeres. — Mesosoma (Figs 1I, 18A–F): Pronotum covered by uniform longitudinal striae, the striae fading dorsally leaving small dorsal smooth area. Epomia weak, reaching pronotal depression. Mesoscutum almost entirely smooth and shiny, except for distinct set of longitudinal striae between posterior end of notauli. Notaulus narrow and deep with minute dense transverse striae, reaching 0.75 of mesoscutum. Scutellum weakly rugulose punctate dorsally. Mesopleuron and metapleuron densely setose. Mesopleuron dorsally rugulose–striate, ventrally finely rugulose. Epicnemial carina distinct, reaching 0.9 of distance to subtegular ridge. Sternaulus shallow and wide, sinuous, reaching posterior rim of mesopleuron. Metapleuron finely strigose. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum finely transversely strigose. Propodeal spiracle elliptic, SLW 2.5–2.75. Posterior area of propodeum covered by more or less uniformly spaced transverse striae. Posterior transverse carina of propodeum absent. Areolet small, closed distally by an unpigmented vein 3rs-m, APH 0.40–0.45. Vein 2m-cu antefurcal to vein 3rs-m. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 2.20–2.35.— Metasoma (Fig. 18A, H): Posterior end of S1 placed distinctly posterior to spiracle of T1. T1LW 2.9–3.35; T1WW 2.25–2.3; T2LW 1.10–1.45; T2WW 2.2–3.1. Thyridium about 3 times as long as wide. T2 minutely punctate reticulate on a finely granulate background, except for small granulate portion on anterior part. Dorsal valve of ovipositor with 4 teeth. Ventral valve with 10 teeth. OST 4.75–5.15. — Color (Figs 1I, 18): Head mostly black: labrum, mandible except apex, clypeus dorsally, facial, frontal and genal orbits dorsally, whitish. Antenna mostly dark brown, f6–f11(12) dorsally white, Mesosoma mostly orange: propleuron, pronotum mostly, ventral part of epicnemium and mesopleuron, mesosternum, metasternum, metapleuron ventrally, marks on anterior part of median lobe and center of lateral lobes of mesoscutum and dorsal part of propodeum dark brown to black; dorsal rim of pronotum, subtegular ridge and lateral parts of scutellum and posterior dorsal mark on propodeum, whitish. Legs mostly dark brown; mid femur dorsally and most part of hind femur orange; dorsal posterior longitudinal strip on fore tibia and ventral part of fore and mid trochanters, whitish. Wing hyaline. Metasoma mostly black: posterior narrow bands on T1–T2, posterior rim of T3 slightly and dorsal part of T6–T8 extensively, whitish. — MALE. Unknown.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from Uganda (Fig. 25A).

Material examined.

6 ♀♀. Holotype. UGANDA • ♀; Kabarole district, Kibale National Park, Fort Portal; 4–18.VI.2015; T. Hopkins leg. Malaise; ZMUT.7076 / http://mus.utu.fi/ZMUT.11402 (ZMUT). Dry pinned. Distal flagellomeres from f31 of right antenna, and hind left leg, missing. — Paratypes. UGANDA • 1 ♀; same data as holotype except 16–30.I.2015; K31T3 – 150130 / http://mus.utu.fi/ZMUT.11367 (ZMUT) • 1 ♀; same data except 23.V–04.VI.2015; K31T4 / http://mus.utu.fi/ZMUT.11392 ZMUT.7534 (ZMUT) • 1 ♀; same data except 30.I.2015–13.II.2015; K31T4, http://mus.utu.fi/ZMUT.11397, ZMUT.7535 (ZMUT) • 1 ♀; same data except 28.I.2015; sweeping, http://mus.utu.fi/ZMUT.11395 (ZMUT) • 1 ♀; same data except Kanywara, Makerere University Biological Field station; 1484 m; primary mid–alt rainforest near stream; 0°34.405′N 30°21.646′E; 12–26.VIII.2008, S. van Noort leg.; Malaise; UG08–KF10–M20 / SAM–HYM–P051187 (SAMC).

Stenarella tripartita (Brullé, 1846)

Figures 1E, 19, 22D, 25B

Mesostenus tripartitus Brullé, 1846: 222. Lectotype ♀ (MNHN, designated by Townes and Townes (1973); examined).

Stenarella tripartita: Townes and Townes (1973).

Diagnosis.

Stenarella tripartita can be distinguished from all other Stenarella species by the combination of the following characters: vertex densely punctate (Fig. 19D); mesoscutum densely and coarsely punctate on a granulate background (Fig. 19D); propodeum relatively long, dorsal profile in lateral view evenly convex and sloped (Fig. 19B, 22D); areolet closed, vein 2m-cu antefurcal to vein 3rs-m (Fig. 19E); genal orbit completely yellow (Fig. 19A, B, 22D) mesosoma mostly orange, subtegular ridge yellow (Fig. 19A, B, 22D).

Figure 19. 

Morphology of Stenarella tripartita, female. A Habitus, lateral view; B head and mesosoma, lateral view; C head, frontal view; D head and mesosoma, dorsal view; E fore wing; F metasoma and ovipositor, lateral view.

Description.

FEMALE. Fore wing 6.5–10.5 mm long. — Head (Fig. 19A–D): Mandible, clypeus and face moderately setose. Mandible weakly striate, MLW 1.85–2.2. Malar space short, MSM 0.4–0.45. Clypeus sparsely punctate CWL 2.7–2.9. Clypeus and face centrally mostly flat. Face centrally distinctly wrinkled, elsewhere granulate–punctate. Frons densely punctate, with complete longitudinal carina Vertex densely punctate on a smooth or finely granulate background. Antenna with 32–33 flagellomeres. — Mesosoma (Figs 1E, 19A, B, D, E): Pronotum covered by uniform longitudinal striae, dorsal lateral part densely punctate. Epomia strong, reaching pronotal depression. Mesoscutum coarsely and densely punctate on a granulate background, with strong rugae between posterior part of notauli. Notaulus reaching 0.75 of mesoscutum, with strong transverse striae. Scutellum densely punctate. Mesopleuron and metapleuron moderately setose. Mesopleuron dorsally rugulose–striate, ventrally finely rugulose. Epicnemial carina weak (better seen in ventrolateral view), reaching 0.6 of distance to subtegular ridge. Sternaulus shallow and sinuous, reaching anterior 0.7–0.8 of mesopleuron. Metapleuron rugulose punctate. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum rugulose punctate. Propodeal spiracle elliptic, SLW 2.5–3.0. Area between anterior and posterior transverse carina rugulose. Posterior area of propodeum strongly rugulose dorsally, reticulate rugose laterally. Posterior transverse carina of propodeum distinct and complete, distance to anterior transverse carina 2.0 times the distance from anterior carina to anterior margin of propodeum. Areolet closed distally by an unpigmented vein 3rs-m, APH 0.45–0.55. Vein 2m-cu antefurcal to vein 3rs-m. Hing wing vein Cua distinctly longer than crossvein cu-a, HW1C 2.75–2.8. — Metasoma (Fig. 19A, F): Posterior end of S1 placed anteriorly or approximately opposite to spiracle of T1. T1LW 2.3–2.35; T1WW 2.7–2.85; T2LW 0.95–1.1; T2WW 2.1–2.15. Thyridium about 2 times as long as wide. T2 minutely punctate reticulate on a granulate background, anterior part granulate. Dorsal valve of ovipositor with 5 teeth. Ventral valve with 9 teeth. OST 2.3–3.4. — Color (Figs 1E, 19): Head mostly black, often orange posteriorly to ocelli; clypeus, labrum, mandibles except apex, facial frontal and genal orbits and scape ventrally, whitish; f7–f10 dorsally and f6 and f11 partially, white. Mesosoma mostly orange; dorsal margin of pronotum, subtegular ridge, whitish; propleuron, mesosternum and metasternum dark brown. Legs mostly dark orange to dark brown; fore coxa posteriorly and all trochanters ventrally, whitish; hind coxa, with dorsal orange stripe; hind t2–4 dorsally white. Wing hyaline. Metasoma mostly black: T1 brown; T1–T2 and T5–T8 posteriorly whitish. — MALE (Fig. 22D). Fore wing 5.3–8.3 mm long. — Head: MSM 0.45–0.5. CWL 3.2. Antenna with 30–32 flagellomeres; tyloids on f12(13)–f15(16). — Mesosoma: SLW 3.5–3.65. APH 0.35–0.45. Vein 2m-cu antefurcal or interstitial to vein 3rs-m. HW1C 1.85–2.0. — Metasoma: Posterior end of S1 placed anteriorly or approximately opposite to spiracle of T1. T1LW 4.45–5.3; T1WW 1.45–1.75; T2LW 2.4–3.3; T2WW 1.8–1.9. — Color: As in female. Flagellum without white band. Fore and mid coxa extensively and all trochanters ventrally white; all tarsi dark brown. Other characters as in female.

Hosts.

Eumenes maxillosus (De Geer, 1773) (Hymenoptera; Vespidae; Eumeninae); Tricarinodynerus anceps anceps (Gribodo1892) (Hymenoptera; Vespidae; Eumeninae); Rhynchium tropicalis (Hymenoptera; Vespidae; Eumeninae); Sceliphron spirifex L. 1758 (Hymenoptera; Sphecidae).

Distribution.

Afrotropical. Known records from BeninNR, CameroonNR, ChadNR, Ethiopia, Ghana, Guinea, Ivory CoastNR, LiberiaNR, MozambiqueNR, NigeriaNR, Republic of the CongoNR, Senegal, Sierra LeoneNR, TanzaniaNR, Togo and ZimbabweNR (Fig. 25B).

Material examined.

78 ♀♀, 10 ♂♂. BENIN • 2 ♀♀; Djougou, Lieut Brot; 1908 (MNHN)• 6 ♀♀, Porto–Novo; 1912; Waterlot (MNHN). CAMEROON • 1 ♂; Dschang; June–September 1924; Dr. Grommier. • 1 ♀; Prov. Kribi, Campo; 28–29.III.1974; leg. W. Schacht (ZSMC). CENTRAL AFRICAN REPUBLIC • 1 ♀; Kaga–Bandoro [“Congo Français, Fort Crampel”] (MNHN). CHAD • 1 ♀; Kanem region; 1904; Captain Dupertus (MNHN). GHANA • 1 ♀; Ashanti Region, Kumasi–Kwadaso; IV.1970; leg F. Bachmaier. GUINEA • 1 ♀; Conakry; January 1923; Millet–Horsin (MNHN) • 1 ♀; June–July 1924; Mlle Homburger et Morrell (MNHN) • 2 ♀♀; N’Zébéla; 1920; P. Chabanaud (MNHN) • 5 ♀♀; Seredou; January 1958; R. Pujol (MNHN) • 2 ♀♀; Tamara; J. 1913; Serand (MNHN) • 1 ♀; Westerm; 10832 (MFNB). IVORY COAST • 1 ♀; Adioppo Doume; September 1965; Gillon (MNHN) • 1 ♀; 1906; A. du Guiney (MNHN) • 1 ♀; Bouaké; Delattre (MNHN) • 1 ♀; Danané; December 1958; B. Faye and R. Roy (MNHN) • 1 ♀; Lamto Scientific Reserve; 21.7.1964 (MNHN) • 5 ♀♀; Odienne; 1973–1975; V. Villard (MNHN) • 1 ♀; Oumé; 1912 (MNHN) • 10 ♀♀; Toumodi; June–July 1962; Mme. Gillon. (MNHN). LIBERIA • 2 ♂♂, 1 ♀; Nimba County; May 1942; M. Lamotte (MNHN) • 1 ♀; Clay–Ashland, 1895, Mrs. Sharp coll (USNM). MOZAMBIQUE • 1 ♂; Gorongosa National Park; October 1907; G. Vasse (MNHN). NIGERIA • 3 ♀♀; Olokemeji, Ibadan (USNM) • 1 ♀; Moor Plantation, c. 580 ft, 4 mi. W. of Ibadan; 30.IX.1913; W.A. Lamborn (OXUM) • 1 ♀; same date except 31.XII.1913 (OXUM) • 1 ♀; same date except 12.I.1914 (OXUM) • 1 ♂; same date except 19.I.1914 (OXUM) • 1 ♀; same date except 30.I.1914 (OXUM) • 1 ♀; Oni, 70 mi. E Lagos, In house, In verandah, Weat Seas c. mid; 20.IX.1911; W.A.Lamborn (OXUM) • 1 ♀; N. Nigeria, Zungeru, 1911–423; Dr. W. Morrison; Mesostenus lissonotus (NHMUK). REPUBLIC OF THE CONGO •1 ♀; Brazzaville; January 1978; Bilsimodo P. (MNHN) • 2 ♀♀; Brazaville; 1908; E. Roubaud & A.Weiss (MNHN). SENEGAL • 1 ♀; Dakar; June 1960 (MNHN) • 1 ♀; same locality; August 1981; B. Sigwalt (MNHN) • 2 ♀♀, same locality, April 1981; M’bour; V. Viltard (MNHN) • 1 ♀; Nioro du Rip (MNHN) • 1♀; Senegal; 10834 (MFNB) • 2 ♀♀; Sudano–Sahel Region; August–October 1809; F. de Zeltner (MNHN). • 1 ♀; Basse Casamance (=Ziguinchor Region) Pointe Saint–Georges; 3.I.1977 (ZSMC). SIERRA LEONE • 1 ♂; Njala, 27.VII.1930; E. Hargreaves (NHMUK) • 1 ♀ 1 ♂; same data except 1.IX.1930 (NHMUK) • 1 ♂; same data except “at light”; 30.VII.1925; “Pres. By Imp. Insti. Ent.; B.M. 1936–224 (NHMUK) • 1 ♀; same locality except 16.XI–4.XII.1935; Van Zwaluwenburg & McGough (USNM) • 1 ♂; Freetown; X.1966–II.1967; D.F.Owen (USUC). TANZANIA • 1 ♀; Kasanga; March 1958; L. Conradt (MNHN). TOGO • 1 ♀; Bismarckburg; Togoland; R. Büttner S.; 26367 (MFNB). ZIMBABWE • 1 ♀; Bindura, 1913; SAMC–HYM–P000735 • 1 ♂, same data except “Goryphus bisulcatus Morl, Antrotype IV.1919, Named by Claude Morley”; SAMC–HYM–P000734; “Stenarella sp. 1 det. Townes 1970”. UNKNOWN COUNTRY • 1 ♀; “W. Afr”; purchase 1875; W.M. Saunders Coll. (OXUM).

Stenarella vannoorti sp. nov.

Figures 1L, 20, 22E, 25C, 28

Etymology.

Named after hymenopterist Simon van Noort (SAMC), who collected the holotype of this species and as a homage to his extraordinary 33-year effort to document and understand the Afrotropical entomofauna, including many invaluable contributions to the systematics of Ichneumonidae. Noun in genitive case.

Diagnosis.

Stenarella vannoorti sp. nov. can be distinguished from all other Stenarella species by the combination of the following characters: mandible striated at basal half (Fig. 20C); mesoscutum mostly smooth and shiny (Fig. 20F); notaulus conspicuously wide with coarse spaced transverse keels (Fig. 20F); posterior area of propodeum rugulose strigose (Fig. 1L); propodeal spiracle short, SLW 1.4–1.45 (Fig. 20E). Additionally, female has clypeus and face concave in lateral profile; areolet open (Fig. 20D), and mesosoma mostly orange (Fig. 20E). Male has clypeus and face in lateral profile almost flat; propodeum and metapleuron black and all coxae dark brown to black (Fig. 22E).

Figure 20. 

Morphology of Stenarella vannoorti sp. nov., female (holotype). A Habitus, lateral view (holotype); B head, frontal view; C mandible; D fore wing; E head and mesosoma, lateral view; F head, mesoscutum and scutellum, dorsal view; G metasoma and ovipositor, lateral view.

Description.

FEMALE. Fore wing 8.1 mm long. — Head (Fig. 20A–C, E, F): Mandible, clypeus and face moderately setose. Mandible distinctly striate, MLW 2.2. Malar space moderately long, MSM 0.7. Clypeus sparsely punctate, CWL 2.45. Clypeus and face centrally slightly but distinctly concave. Face centrally weakly rugulose, elsewhere granulate. Frons shiny, granulate and wrinkled posteriorly to antennal sockets, with only traces of longitudinal carina. Vertex granulate with sparse punctures. Antenna with 32 flagellomeres. — Mesosoma (Fig. 1L, 20A, D–F): Pronotum mostly smooth, with distinct punctures along dorsal margin, dorsally and ventrally with longitudinal striae on pronotal depression; anterior rim rugulose punctate. Epomia long and strong, reaching pronotal depression. Mesoscutum mostly smooth and shiny, with dense and shallow punctures on anterior part of median lobe. Notaulus conspicuously wide, reaching 0.65 of mesoscutum with strong transverse striae. Scutellum rugulose punctate. Mesopleuron and metapleuron moderately setose. Mesopleuron mostly rugulose, with short longitudinal striae along epicnemial carina. Epicnemial carina weak but complete through subtegular ridge. Sternaulus deep and sinuous, reaching posterior rim of mesopleuron. Metapleuron rugulose. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum weakly rugulose. Propodeal spiracle elliptic, SLW 1.4–1.45. Posterior area of propodeum rugulose strigose. Posterior transverse carina of propodeum medially indistinct, sublaterally present but weakly discernible from other propodeal striae, distance to anterior transverse carina 1.5 times the distance from anterior carina to anterior margin of propodeum. Areolet very small, open distally, APH 0.35. Hind wing vein Cua distinctly longer crossvein cu-a, HW1C 2.0. — Metasoma (Fig. 20A, G): Posterior end of S1 placed approximately opposite to spiracle of T1; T1LW 2.9; T1WW 1.9; T2LW 1.3; T2WW 2.55. Thyridium about 2 times as long as wide. T2 mostly punctate reticulate on a granulate background, except for small granulate portion on anterior part. Dorsal valve of ovipositor with 4 teeth. Ventral valve with 10 teeth. OST 5.4. — Color (Figs 1L, 20): Head black; clypeus, face, frons and vertex, partially bright orange. Antenna dark brown, f6–f10 dorsally white. Mesosoma mostly orange; propleuron, pronotum anteriorly, sternal groove and posterior part of mesosternum, metasternum and ventral part of metapleuron black. Legs mostly dark brown; fore femur tibia and tarsus clearer. Wing hyaline slightly obscured distally. Metasoma mostly black, T5–T8 posteriorly white. — MALE (Fig. 22E). Fore wing about 8 mm long. — Head: Face and clypeus flat in lateral profile; malar space short, MSM 0.35; CWL 2.5. Frons smooth and shiny, with fine sparse punctures and weak longitudinal carina. Vertex smooth with sparse punctures. Antenna with 32 flagellomeres, tyloids on f14–f16. — Mesosoma: Pronotum mostly smooth, with strong longitudinal striae along pronotal depression, weaker on posterior rim. Scutellum punctate. Anterior area of propodeum rugulose punctate. SLW 1.45. Areolet short, closed distally, APH 0.45. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C about 1.3. — Metasoma: Posterior end of S1 placed posterior to spiracle of T1; T1LW 3.3; T1WW 2.35; T2LW 2.35; T2WW 1.55. Thyridium about 2.25 times as long as wide. — Color: Antenna dark brown, f8–f16 dorsally white, f17 partially. Mesosoma mostly orange; propleuron, pronotum anteriorly, mesosternum, posterior part of mesopleuron, posterior axillae, metapleuron and propodeum, black. Legs mostly dark brown; fore and mid femurs and tibiae orange; hind t2–t4 white. Wing hyaline. Metasoma mostly black, T1–T5 posteriorly white. T6–T8 widely white. Other characters as in female.

Hosts.

Unknown.

Distribution.

Afrotropical. Known records from South Africa (Fig. 25C).

Material examined.

1 ♀, 1 ♂. Holotype. SOUTH AFRICA ♀ • Eastern Cape, Asante Sana Game Reserve; 32°163762′S 24°57.309′E; 1186 m; Southern Karoo riviere, riverine woodland; 23.II–7.IV.2010; S. van Noort leg.; Malaise; ASA09–WOO1–M06 / SAM–HYM–P049958 (SAMC). Dry pinned. Distal flagellomeres from f6 of left antenna, mid and hind left legs, missing; flagellomeres from f2 of right antenna and left ovipositor sheath glued on cardboard label. — Paratype. SOUTH AFRICA • 1 ♂; KwalaZulu–Natal, Pietermaritzburg; 27.X.1970; H. & M. Townes (USUC).

Stenarella victoriae (Cameron, 1912)

Figures 1B, 21, 22F, 25D, 28

Mesostenus victoriae Cameron, 1912: 179. Holotype ♀ (NHMUK, examined).

Parasilsila trilineata Cheesman, 1936: 368. Holotype ♀ (NHMUK, examined). Synonymized by Townes et al. (1961).

Diagnosis.

Stenarella victoriae can be distinguished from all other Stenarella species by the combination of the following characters: mesoscutum densely punctate (Fig. 21E); areolet usually open, vein 2rs-m obliterated (Fig. 21C); propodeum medially and posteriorly to anterior transverse carina coarsely transversely striate (Fig. 1B); mesosoma black, with well defined whitish marks (Fig. 21A, D, E, 22F); propodeum with one medial posterior large whitish mark (Figs 1B, 22F).

Figure 21. 

Morphology of Stenarella victoriae, female. A Habitus, lateral view; B head, frontal view; C fore wing; D head and mesosoma, lateral view; E head, mesoscutum and scutellum, dorsal view; F metasoma and ovipositor, lateral view.

Figure 22. 

Habitus (lateral view) Stenarella spp., males. A Stenarella domator; B:Stenarella insidiator; C Stenarella lissonota; D Stenarella tripartita; E Stenarella vannoorti (paratype); F Stenarella victoriae.

Description.

FEMALE. Fore wing 10.0–12.5 mm long. — Head (Fig. 21A, B, D, E): Mandible, clypeus and face moderately setose. Mandible mostly sparsely punctate, MLW 2.1–2.25. Malar space short, MSM 0.40–0.45. Clypeus sparsely punctate, CWL 3.15–3.20. Clypeus slightly convex, face centrally distinctly convex. Face granulate–punctate, dorsomedially rougher, with median minute tubercle between antennal sockets. Frons and vertex mostly densely punctate, frons medially rugulose–punctate, with interrupted longitudinal carina. Antenna with 35–37 flagellomeres. — Mesosoma (Fig. 1B, 21A, C–E): Pronotum mostly smooth, with distinct longitudinal striae along anterior depression and posterior margin, dorsal lateral part strongly and densely punctate. Epomia long and strong, ending distinctly on pronotal collar. Mesoscutum densely punctate. Notaulus reaching 0.80 of mesoscutum, with minute transverse striae. Scutellum mostly punctate with central part smooth. Mesopleuron and metapleuron moderately setose. Mesopleuron dorsally densely punctate, centrally strigose punctate, ventrally rugulose punctate. Epicnemial carina weak (better seen in ventrolateral view), reaching 0.4–0.5 of distance to subtegular ridge. Sternaulus deep and sinuous, reaching posterior rim of mesopleuron. Metapleuron posterodorsally rugose punctate, anteriorly rugulose punctate and posteroventrally strigose punctate. Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped. Anterior area of propodeum densely punctate, with median longitudinal carina distinct. Propodeal spiracle elliptic, SLW 5.0–5.25. Posterior area of propodeum anteriorly rugulose reticulate, posteriorly and medially, coarsely striate transversally, laterally rugulose striate. Posterior transverse carina of propodeum absent. Areolet small, open distally, APH 0.4–0.45. Vein 2rs-m obliterated. Hind wing vein Cua distinctly longer than crossvein cu-a, HW1C 2.75–2.85. — Metasoma (Fig. 21A, F): Posterior end of S1 placed anterior to or approximately opposite to spiracle of T1; T1LW 2.8–3.2; T1WW 2.4–2.5; T2LW 1.1–1.15; T2WW 2.1–2.15. T2 minutely punctate reticulate on a granulate background. Dorsal valve of ovipositor with 8 teeth. Ventral valve with 11 teeth. OST 3.4–3.5. — Color (Figs 1B, 21): Head mostly black: labrum, clypeus, face centrally, mandibles except distal part and eye orbits, whitish and f7–13(14) dorsally white. Mesosoma mostly black; propleuron widely, dorsal lateral margin, spot on medial dorsal part and a ventral anterior spot on pronotum, subtegular ridge, spot between median anterior part of mesopleuron and median posterior part of epicnemium, posterior ventral corner of mesopleuron, mesepisternum, dorsal division of metapleuron, wide dorsal mark on posterior part of metapleuron, central spot on mesoscutum, tegula, scutellum laterally and dorsally, and a large medial posterior spot on propodeum, yellow cream. Legs mostly orange; distal part of hind femur and tibia, base of t1, and t5 dark brown; distal part of t1 and t2–4 whitish. Wing hyaline. Metasoma mostly dark brown to black, T1–T8 posteriorly yellow cream banded. — MALE (Fig. 22F). Fore wing 8.7–10.2 mm long. — Head: MLW 1.9–2.0; MSM about 0.36. CWL 2.6–2.7. Antenna with 32–37 flagellomeres. Tyloids on f14–f18, sometimes on f15–18 or f13–19. — Mesosoma: SLW 5.5–6.0., HW1C 2.2–2.7. — Metasoma: Posterior end of S1 placed posterior to spiracle of T1; T1LW 4.9–5.1; T1WW 1.5–2.0; T2LW 1.4–1.8; T2WW 2.2–2.4. — Color: As female, but antenna with f7–f19 dorsally white, first one and last one partially, some specimens without white band; face and mesosternum entirely whitish. Other characters as in female.

Figure 23. 

Distribution maps for species of Stenarella. A Stenarella aurea; B Stenarella brykella; C Stenarella bugalana; D Stenarella delicata; E Stenarella favilla; F Stenarella fenestralis.

Figure 24. 

Distribution maps for species of Stenarella. A Stenarella hopkinsi; B Stenarella katanga; C Stenarella lissonota; D Stenarella natalina; E Stenarella nigriscuta; F Stenarella nigromaculata.

Comments

According to Gauld (1984), this species is “fairly common in Queensland and New South Wales”, and it is well-represented in Australian collections.

Hosts.

Auplopus sp. (Hymenoptera; Pompilidae), Sceliphron sp. (Hymenoptera; Sphecidae).

Distribution.

Australasian. Known records from Australia (Queensland, Tasmania, Victoria) (Fig. 25D).

Figure 25. 

Distribution maps for species of Stenarella. A Stenarella saaksjarvi; B Stenarella tripartita; C Stenarella vannoorti; D Stenarella victoriae.

Figure 26. 

Approximate distribution map for Stenarella domator from a compilation of literature sources, online databases and material examined in this study. The upper right inset shows confirmed occurrence points in North America, where the species seems to have been recently introduced.

Figure 27. 

Distribution map for Stenarella insidiator. Records with precise geographic location depicted as red dots; records identified only to country of province depicted as light red coloring of the narrowest geographic area mentioned in the record.

Material examined.

10 ♀♀, 2 ♂♂. AUSTRALIA • 1 ♀; Queensland, Acadia Ridge; 20.Dec.1979, G. Gordhi (USNM) • 1 ♀ 1 ♂; same data except 29.Dec.1979 • 1 ♀; same date except 30.Dec.1979 • 1 ♀; Queensland, Rockhampton; 1870; Thozet, (MNHN) • 1 ♀, Tasmania, Verreaux; March 1947 (MNHN) • 1 ♀ 1 ♂; Queensland, N. of Mt. Molloy, Kingfisher Park, Julatten, 14.XI.1996, G.R. Else • 1 ♀; Queensland, Brisvane, Gap, Ashgrove, August, F.G. Smith (USUC) • 1 ♀; Queensland, Rockhampton; Coll. Bingham (MFNB) • 1♀; Victoria; Musk, 25.II–4.III.2016; A. Campoy Leg.; Rza. 1; Malaise trap (CEUA) • 1♀, 1 ♂; Victoria; same locality but 6–19.I.2017; A. Campoy Leg.; Rza. 24; Malaise trap (CEUA).

3.2. Molecular phylogenetics

Maximum-likelihood analyses resulted in highly supported trees (average BTP 98.6) and recovered Stenarella a monophyletic group, with full support (Fig. 28; File S3). The genus was sister to Nematopodius Gravenhorst, with the next closest genus being Picardiella Lichtenstein.

Figure 28. 

Phylogenetic relationships among species of Stenarella as inferred from a máximum-likelihood analysis of genomic ultraconserved elements. Numbers at nodes correspond to bootstrap support values; values missing from nodes 100% clade support for that metric.

The species of Stenarella were grouped in three major clades, each with full support, and corresponding to the broad scale biogeographic zones in which the genus is distributed. One clade includes the single, widely distributed Palearctic species of the genus, S. domator. Our dataset included specimens from three of the four currently recognized subspecies: the North African S. domator ensnator (Thunberg) was recovered as sister to the other two subspecies. The second clade includes S. insidiator, which is widely distributed in Southeast Asia from India to the Philippines and Indonesia, and the Australian S. victoriae. The third clade included eleven species from the Afrotropical region, with three of the fourteen species recognized herein not successfully sequenced.

Within this Afrotropical clade, a long branch separates S. vannoorti sp. nov. from all other taxa, indicating an early divergence between this single species and all other Afrotropical species. No obvious morphological or distributional pattern was observed in the relationships among species of the clade. For instance, while a subclade comprised of S. fenestralis sp. nov., S. lissonota and S. natalina sp. nov. seems to be restricted to Southern Africa (South Africa, Zimbabwe, Zambia and Angola), the distantly related S. vannoorti is also endemic to South Africa.

Species of Stenarella tended to be relatively uniform in overall body proportions and major morpho–structural characters, with most differences among species corresponding to details in sculpturing, pilosity, and wing venation. Color patterns vary among species but also within species, as observed particularly among specimens of S. saaksjarvi sp. nov. and among the various regional subspecies of S. domator. The one body proportion that seems to vary substantially among species is the ovipositor length. However, we observed a marked populational difference in ovipositor length within S. insidiator: specimens collected in Borneo seem to have smaller body size overall and a relatively smaller ovipositor (as measured by the ratio between ovipositor length and hind tibia). Figure 29 shows a plot of this measurement against total body size (as estimated by fore wing length) for all but two female specimens, highlighting the difference between the Borneo specimens and the remaining populations of S. insidiator.

Figure 29. 

Relationship between total body size (using fore wing length as a proxy) and relative ovipositor length, highlighting the apparent populational divide in ovipositor length within S. insidiator.

4. Discussion

The relationship of Stenarella to other cryptine genera recovered herein was identical to that found by Santos and Brady (2024) using a combination of UCE and Sanger sequencing data. These relationships contrast with the results of Santos (2017), based on seven genes and morphological data, who recovered Stenarella as sister to a clade comprised of Melcha Cameron + Necolio Cheesman, and only distantly related to Picardiella and Nematopodius.

The species richness of Stenarella in the Afrotropics had already been noted by Townes (1970), who remarked “in the Ethiopian region there are a number of species, elsewhere very few”. We confirmed this observation by finding twelve new Afrotropical species, bringing the total in the region to fifteen, compared to only three species elsewhere in the world. Notably, though, our phylogeny does not suggest that the few species of Stenarella out of the Afrotropics represent a recent range expansion from an Afrotropical stock: the topology of the genus shows an ancient split in two clades corresponding to the Afrotropical and non-Afrotropical component. Hence, the disparity of species richness between the two clades seems to be attributable to a more rapid diversification in the Afrotropical region, though without a dated tree it is impossible to identify for sure what climatic or biogeographic factors may have influenced this faster rate of species accumulation. However, the limits of the widely distributed S. domator and S. insidiator still need to be better investigated. Both taxa show substantial morphological variation throughout its range: S. domator is currently divided into four subspecies, mostly based on coloration differences, while in S. insidiator there is substantial variation in body size and ovipositor length (see above). More in-depth analyses using molecular data with a broader sampling could reveal the presence of cryptic species within both taxa, though it is unlikely that the diversity of Stenarella in the Palearctic and Oriental regions could match that of the Afrotropics.

Notably, in spite of their broad distribution in the Old World, Stenarella have not colonized Madagascar, where the main cryptine attacking nests of mud-nesting wasps is Sphecotonus Seyrig; it had been placed in Osprynchotina but was found to be only distantly related to Stenarella (Santos 2017). At first glance, this would seem to suggest that competitive exclusion may be an important factor shaping cryptine distributions. However, throughout its geographic range, species of Stenarella share the environment with other cryptine taxa that are specialized in parasitizing the pupae of mud–nesting aculeates. Int he Afrotropics, the other “Osprynchotina” are Picardiella and Osprynchotus Spinola; in the Palearctic and Oriental regions we can find species of Nematopodius, Picardiella and Acroricnus Ratzeburg, while in the Australasian region Iaria Cheesman is present.

A similar phenomenon happens in the Neotropics, where in zones past the Andes Mountain range in Chile, the niche of attacking mud–nesting aculeates is filled by Dotocryptus, with no occurrences of the elsewhere abundant and speciose Neotropical genera Photocryptus Viereck and Messatoporus Cushman. In both of these cases, the relative roles of geographic isolation versus competition with other taxa in shaping distributional patterns are still unclear. A better understanding of the natural history of the groups might help to clarify the exact dynamics of competition for resources among species filling apparently similar ecological niches.

5. Acknowledgements

The following curators facilitated visits by BFS to their collections and provided extensive loans of cryptine specimens: Gavin Broad (NHMUK), James Hogan (OXUM), Robert Kula (USNM), Ilari Sääksjärvi (ZMUT), Stefan Schmidt (ZSMC), Simon van Noort (SAMC), David Wahl (USUC) and Kevin Williams (formerly at FSCA). Stefanie Krause kindly helped with logistics involving SB’s visit to the MFNB. Louise Lepert helped with data collection at an early stage of this work. Laura Zara helped with collection of label data. Aisha Mayekiso helped retrieve full label information from database codes from the SAMC collection. Bonnie Blaimer kindly provided access to some of the laboratory reagents for UCE data Generation. Davide dal Pos and an anonymous referee provided valuable comments that improved the original version of the MS. Research funds were provided by the Deutsche Forschungsgemeinschaft (SA 4907/1-1) and a grant from the Global Genome Initiative Peer–Review Awards Program. This study has been carried out in part during two research stays made by SB to MFNB, supported by grants from the University of Alicante (Alicante, Spain) within the framework of the “Programa Propi del Vicerectorat d’Investigació per al Foment de la I+D+I –2022 and 2023 (References ACIE22–03 and ACIE 23–04, respectively)”.

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Supplementary material

Supplementary material 1 

Files S1–S3

Santos BF, Bordera S (2025)

Data type: .zip

Explanation notes: File S1. List of taxa used in the phylogenetic analyses and sequencing metrics, including sequencing reads, assembly results and number UCE loci recovered for each sample. — File S2. Concatenated UCE dataset used in the phylogenetic analyses. — File S3. Maximum-likelihood tree for Stenarella and related taxa.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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