Research Article |
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Corresponding author: Bernardo F. Santos ( bernardofsantos@gmail.com ) Academic editor: Brendon Boudinot
© 2025 Bernardo F. Santos, Santiago Bordera.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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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.
Darwin wasp, Cryptini, Osprynchotina, mud-nesting, biogeography
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.
All of these species share the following characters that have been used in the keys and descriptions of Townes’ revisionary works (
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
Stenarella was placed by
In spite of their broad distribution and distinctive morphology, no new species of Stenarella have been described since 1943, even though
In this work we included 328 specimens assigned to Stenarella from the following institutions:
Morphological terminology follows
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 (
To assess the monophyly of Stenarella and the relationships among its species, we generated molecular data from genomic ultraconserved elements (UCEs;
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 (
Stenarella
Szépligeti, 1916: 307. Description. Type species: Ichneumon gladiator Scopoli, 1763 (= Ichneumon domator
Orientostenaraeus
Uchida, 1930: 321. Type species: Orientostenaraeus chinensis Uchida, 1930 (= Mesostenus insidiator
Parasilsila Cheesman, 1936: 368. Type species: Parasilsila trilineata Cheesman, 1936 (= Mesostenus victoriae Cameron), by original designation.
Ovipositor very long, OST >2.7 (Fig.
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).
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.
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.
Widely distributed in the Old World, Australia, and introduced into North America. The known distribution records are expanded from 47 countries (
Females
| 1 | Mesoscutum densely punctate or granulate (e.g., Figs |
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 |
7 |
| 2 | Mesoscutum mostly granulate, lateral lobes without punctures (Fig. |
2. S. brykella Roman |
| 2’ | Mesoscutum densely punctate or punctate reticulate (e.g., Figs |
3 |
| 3 | Mesosoma black, with well defined whitish marks (Figs |
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 |
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. |
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. |
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. |
5. S. domator (Poda) |
| 5’ | Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped (Figs |
6 |
| 6 | Vertex finely granulate with very sparse setiferous punctures (Fig. |
4. S. delicata sp. nov. |
| 6’ | Vertex densely punctate on a smooth or finely granulate background (Fig. |
16. S. tripartita (Brullé) |
| 7 | Mesosoma entirely black (Fig. |
14. S. nigromaculata sp. nov. |
| 7’ | Mesosoma black and orange or entirely orange (e.g., Figs |
8 |
| 8 | Posterior area of propodeum transversely striate, striae regularly separated and reaching pleural sulcus (Fig. |
9 |
| 8’ | Posterior area of propodeum mostly rugose, rugulose or irregularly strigose | 12 |
| 9 | Mesosoma light orange without black marks (Fig. |
1. S. aurea sp. nov. |
| 9’ | Mesosoma mostly black and orange (e.g., Figs |
10 |
| 10 | Pronotum coarsely and longitudinally striated (Fig. |
15. S. saaksjarvi sp. nov. |
| 10’ | Pronotum mostly smooth or shallowly striated at pronotal depression (Figs |
11 |
| 11 | Mesopleuron, metapleuron and propodeum dark orange to brownish orange (Fig. |
8. S. hopkinsi sp. nov. |
| 11’ | Mesopleuron, metapleuron and propodeum light orange to orange (Fig. |
13. S. nigriscuta sp. nov. |
| 12 | Mandible striated at basal half (Fig. |
13 |
| 12’ | Mandible smooth or shallowly punctate at basal half (Fig. |
15 |
| 13 | Notaulus conspicuously wide with coarse spaced transverse keels (Fig. |
17. S. vannoorti sp. nov. |
| 13’ | Notaulus narrow or moderately wide with very fine dense transverse keels (Figs |
14 |
| 14 | Posterior end of S1 placed approximately opposite to spiracle of T1 (Fig. |
6. S. favilla sp. nov. |
| 14’ | Posterior end of S1 placed posteriorly to spiracle of T1 (Fig. |
12. S. natalina sp. nov. |
| 15 | Pronotum at most with short fine striae at pronotal depression, elsewhere smooth (Fig. |
11. S. lissonota (Cameron) |
| 15’ | Pronotum mostly with distinct and coarse longitudinal striae, only dorsal posterior part smooth (Figs |
16 |
| 16 | Posterior end of S1 placed distinctively posterior to spiracle of T1 (Fig. |
3. S. bugalana sp. nov. |
| 16’ | Posterior end of S1 placed approximately opposite to or slightly posterior to spiracle of T1 (Figs |
17 |
| 17 | Posterior area of propodeum mainly strigose (Fig. |
10. S. katanga sp. nov. |
| 17’ | Posterior area of propodeum mainly rugulose (Fig. |
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 |
2 |
| 1’ | Mesoscutum densely punctate (As in Figs |
3 |
| 2 | Head with face, clypeus, basal half of mandible, and ocular orbits except at vertex, whitish (Fig. |
11. S. lissonota (Cameron) |
| 2’ | Head dark brown except white facial orbits narrowly (Fig. |
17. S. vannoorti sp. nov. |
| 3 | Mesosoma black, with numerous well defined whitish marks (Fig. | 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. |
5 |
| 4 | Propodeum with one medial posterior large whitish mark (Fig. |
18. S. victoriae (Cameron) |
| 4’ | Propodeum with two lateral posterior whitish marks (Fig. |
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. |
5. S. domator (Poda) |
| 5’ | Propodeum relatively long, dorsal profile in lateral view evenly convex and sloped (Fig. |
16. S. tripartita (Brullé) |
From the Latin adjective aureus, -a, -um, meaning “golden”, in reference to the light orange, yellowish body color of this species.
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.
FEMALE. Fore wing 5.1–6.2 mm long. — Head (Fig.
Unknown.
Afrotropical. Known records from: Angola, Cameroon, Central African Republic, Gabon, Ivory Coast and Sierra Leone (Fig.
11 ♀♀. Holotype. SIERRA LEONE • ♀; Freetown Peninsula, Freetown; III.1969; Denis Owen (
Stenarella brykella
Roman, 1943: 20. Lectotype ♀ (
Stenarella brykella can be easily distinguished from all other species of Stenarella by the mesoscutum mostly granulate without punctures on lateral lobes (Fig.
FEMALE. Fore wing 8.5–10.6 mm long. Head (Fig.
Unknown.
Afrotropical. Known records from Kenya (Fig.
3 ♀♀. Lectotype. [KENYA] • ♀; “Elgon, Bryk / M.E.H. / 10/8(?) / typus; Stenarella brykella n. sp., A. Roman det.; Lectotype Stenarella brykella Roman, Townes, 1972 /
2 ♀♀. KENYA • 1 ♀; Karen, Nairobi, Keya 6000ft; 25–31.VIII.1971; C. van Someren (
Named after Bugala Island, where the paratype was collected. From the Latinized adjective bugalanus, -a, -um.
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.
FEMALE. Fore wing 8.0–8.7 mm long. — Head (Fig.
Unknown.
Afrotropical. Known records from Uganda (Fig.
2 ♀♀. Holotype. UGANDA • ♀; Kampala; 24.VII.1933; H. Hargreaves (
From the Latin adjective delicatus, -a, -um, meaning “delicate”, referencing the fact that this is the smallest of the Stenarella species.
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.
FEMALE. Fore wing 6.3–7.8 mm long. — Head (Fig.
Unknown.
Afrotropical. Known records from Nigeria and Uganda (Fig.
3 ♀♀. Holotype. UGANDA • ♀; Kampala; X.1965; Denis F. Owen (
Ichneumon domator Poda, 1761: 105. Type lost.
Ichneumon gladiator
Scopoli, 1763: 283. Type lost. Synonymized by
Ichneumon tarsosus
Geoffroy in Fourcroy, 1785: 408. Type lost. Synonymized by
Ichneumon insignitor
Villers, 1789: 164. Type lost. Synonymized by
Ichneumon macrourus
Gmelin, 1790: 2687. Type lost. Synonymized by
Listrognathus transsylvanicus
Kiss, 1924: 60. Holotype ♀ (TMA; not examined). Synonymized by
Stenarella gladiator corsicator
Aubert, 1969: 55. Holotype ♀ (
Mesostenus cruentator
Klug in Waltl, 1835: 80. Lectotype ♀ (MFNB, designated and synonymized by
Ichneumon ensator
Thumberg, 1822: 259. Holotype ♀ (ZIU, not examined). Synonymized by
Ichneumon domator var. ensator Roman, 1912.
Mesostenus juvenilis
Tosquinet, 1896: 214. Holotype ♀ (
Mesostenus ruficollis
Rudow, 1882: 33. Type lost. Synonymized by
Stenarella domator:
Stenarella domator can be distinguished from all other Stenarella species by the combination of the following characters: mesoscutum densely and coarsely punctate (Fig.
FEMALE. Fore wing 5.8–10.4 mm long. — Head (Figs
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.
According to
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.
The electronic catalog Taxapad (
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
129 ♀♀; 10 ♂♂.
Stenarella domator corsicator
(4 ♀♀): FRANCE • 1 ♀; Corsica; C. Villemant (
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 (
Stenarella domator domator
(102 ♀♀, 9♂♂): FRANCE • 2 ♀♀; Allier, Brout–Vernet; August 1928; H. du Buysson (
Stenarella domator ensator
(15 ♀♀, 1♂): ALGERIA • 11 ♀♀, 2 ♂; Annaba; 1963; O. Sichel (
From the Latin favilla, -ae, meaning “embers”, in reference to the irregular pattern of black and ferruginous resembling glowing embers. Noun in apposition.
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.
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).
FEMALE. Fore wing 10.7–11.3 mm long. — Head (Fig.
Unknown.
Afrotropical. Known records from South Africa (Fig.
2 ♀♀. Holotype. SOUTH AFRICA • ♀; KwalaZulu–Natal, Umhlanga; 22.I.1960; G. C. Clark (
From the Latinized adjective fenestralis, -e, meaning “window”, in reference to the amusing fact that one of the specimens was collected at a window.
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.
FEMALE. Fore wing 7.2–12 mm long. — Head (Fig.
Unknown.
Afrotropical. Known records from Angola, Malawi, Zambia and Zimbabwe (Fig.
21 ♀♀. Holotype. ZIMBABWE (“Rhodesia”) • ♀; Harare (“Salisbury”), Chishawasha; III.1979; A. Watsham (
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.
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.
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
Unknown.
Afrotropical. Known records from Uganda and Democratic Republic of the Congo (Fig.
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 (
Mesostenus insidiator
Orientostenaraeus chinensis
Uchida, 1930: 321. Holotype ♀ (Hokkaido University, not examined). Synonymized by
Stenarella insidiator:
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.
FEMALE. Fore wing 7.5–16.3 mm long. — Head (Fig.
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.
Sceliphron madraspatanum Klug, 1801 (Hymenoptera, Sphecidae); Orancistrocerus aterrimus (de Saussure, 1852) (Hymenoptera; Vespidae; Eumeninae).
Oriental. Known records from BruneiNR, Cambodia, China (Liaoninh, Sichuan, Yunnan), IndiaNR, Indonesia, MalaysiaNR, MyanmarNR, Philippines, Taiwan, ThailandNR, Vietnam (Fig.
18 ♀♀, 2 ♂♂. Holotype. INDONESIA • ♀; Sulawesi, Makasar (
Named after the province of Haut–Katanga, where the holotype was collected. Noun in apposition.
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.
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).
FEMALE. Fore wing 7.7–10.8 mm long. — Head (Fig.
Unknown.
Afrotropical. Known records from Angola and Democratic Republic of the Congo (Fig.
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 (
Mesostenus lightfooti
Cameron, 1906: 147. Lectotype ♀ (
Mesostenus levifrons
Cameron, 1906: 148. Holotype ♀ (
Mesostenus lissonotus
Cameron, 1906: 149. Holotype ♀ (
Mesotenus albilabris
Cameron, 1911: 177. Holotype ♀ (
Stenaraeus (Mesostenus) lissonotus var. nigripes
Morley, 1916: 377. Holotype ♀ (
Stenaraeus lightfooti:
Stenaraeus laevifrons
(!):
Stenarella lissonotus: Townes and Townes, 1973.
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.
FEMALE. Fore wing 7.5–9.2 mm long. — Head (Fig.
Unknown.
Afrotropical. Known records from South Africa (Fig.
4 ♀♀, 1 ♂. SOUTH AFRICA • 1 ♀; Eastern Cape, Grahamstown; III.1972; Fred Gess (
Named after the KwaZulu-Natal province, where one of the paratypes was collected. From the Latinized adjective natalinus, -a, -um.
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.
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).
FEMALE. Fore wing 8.0–11.3 mm long. — Head (Fig.
Unknown.
Afrotropical. Known records from South Africa and Tanzania (Fig.
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. (
From the Latin adjective niger, -gra, -grum, meaning “black”, and the noun scutum, -i, in reference to the entirely black mesoscutum in this species.
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.
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).
FEMALE. Fore wing 11.3–12.3 mm long. — Head (Fig.
Unknown.
Afrotropical. Known records from Cameroon, Nigeria and Republic of the Congo (Fig.
6 ♀♀. Holotype. CAMEROON • ♀; Nkoemvom; 13.VII–24.VIII.1980; Ms. D. Jackson (
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.
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.
FEMALE. Fore wing 11.1 mm long. — Head (Fig.
Unknown.
Afrotropical. Known records from Central African Republic (Fig.
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 (
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.
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.
Morphology of Stenarella saaksjarvi sp. nov., female. A Habitus, lateral view (holotype); B–D 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).
FEMALE. Fore wing 5.8–11.3 mm long. — Head (Fig.
Unknown.
Afrotropical. Known records from Uganda (Fig.
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 (
Mesostenus tripartitus
Brullé, 1846: 222. Lectotype ♀ (
Stenarella tripartita:
Stenarella tripartita can be distinguished from all other Stenarella species by the combination of the following characters: vertex densely punctate (Fig.
FEMALE. Fore wing 6.5–10.5 mm long. — Head (Fig.
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).
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.
78 ♀♀, 10 ♂♂. BENIN • 2 ♀♀; Djougou, Lieut Brot; 1908 (
Named after hymenopterist Simon van Noort (
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.
FEMALE. Fore wing 8.1 mm long. — Head (Fig.
Unknown.
Afrotropical. Known records from South Africa (Fig.
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 (
Mesostenus victoriae
Cameron, 1912: 179. Holotype ♀ (
Parasilsila trilineata
Cheesman, 1936: 368. Holotype ♀ (
Stenarella victoriae can be distinguished from all other Stenarella species by the combination of the following characters: mesoscutum densely punctate (Fig.
FEMALE. Fore wing 10.0–12.5 mm long. — Head (Fig.
According to Gauld (1984), this species is “fairly common in Queensland and New South Wales”, and it is well-represented in Australian collections.
Auplopus sp. (Hymenoptera; Pompilidae), Sceliphron sp. (Hymenoptera; Sphecidae).
Australasian. Known records from Australia (Queensland, Tasmania, Victoria) (Fig.
10 ♀♀, 2 ♂♂. AUSTRALIA • 1 ♀; Queensland, Acadia Ridge; 20.Dec.1979, G. Gordhi (
Maximum-likelihood analyses resulted in highly supported trees (average BTP 98.6) and recovered Stenarella a monophyletic group, with full support (Fig.
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
The relationship of Stenarella to other cryptine genera recovered herein was identical to that found by
The species richness of Stenarella in the Afrotropics had already been noted by
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 (
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.
The following curators facilitated visits by BFS to their collections and provided extensive loans of cryptine specimens: Gavin Broad (
Files S1–S3
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.