Research Article |
Corresponding author: Tapani Hopkins ( taesho@utu.fi ) Academic editor: Bernardo Santos
© 2019 Tapani Hopkins, Heikki Roininen, Simon van Noort, Gavin R. Broad, Kari Kaunisto, Ilari E. Sääksjärvi.
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.
Citation:
Hopkins T, Roininen H, van Noort S, Broad GR, Kaunisto K, Sääksjärvi IE (2019) Extensive sampling and thorough taxonomic assessment of Afrotropical Rhyssinae (Hymenoptera, Ichneumonidae) reveals two new species and demonstrates the limitations of previous sampling efforts. ZooKeys 878: 33-71. https://doi.org/10.3897/zookeys.878.37845
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Tropical forest invertebrates, such as the parasitoid wasp family Ichneumonidae, are poorly known. This work reports some of the first results of an extensive survey implemented in Kibale National Park, Uganda. A total of 456 individuals was caught of the subfamily Rhyssinae Morley, 1913, which in the Afrotropical region was previously known from only 30 specimens. Here, the six species found at the site are described and the Afrotropical Rhyssinae are reviewed. Two new species, Epirhyssa johanna Hopkins, sp. nov. and E. quagga sp. nov., are described and a key, diagnostic characters, and descriptions for all 13 known Afrotropical species are provided, including the first description of the male of Epirhyssa overlaeti Seyrig, 1937. Epirhyssa gavinbroadi Rousse & van Noort, 2014, syn. nov. is proposed to be a synonym of E. uelensis Benoit, 1951. Extensive sampling with Malaise traps gave an unprecedented sample size, and the method is recommended for other poorly known tropical areas.
Africa, Ichneumonidae, Kibale National Park, Uganda Malaise trapping 2014–2015
Like many taxa, the parasitoid wasps of the family Ichneumonidae are poorly known in the tropics. So much so, that the family was once assumed to be exceptionally species-poor in the equatorial region and to peak in species richness in mid latitudes (
The ichneumonid subfamily Rhyssinae is especially poorly known in the Afrotropical region. It was reviewed in 2014 (
One possible reason for the lack of rhyssine specimens is that adequately inventorying tropical ichneumonids appears to need long-term, extensive sampling (for reasons summarised in
In this and a sister paper, we report the first results of an extensive one-year sampling of Afrotropical ichneumonids in Kibale National Park, Uganda. In the sister paper we report the ecological results for the subfamily Rhyssinae, including descriptions of the habitat use and phenology of the species (
We sampled ichneumonids with 34 Malaise traps for a full year (2014–2015) in Kibale National Park, Uganda. The traps were placed in a wide variety of habitats ranging from primary forest to clear-cut former plantations and farmland, and the total sampling effort was roughly 382 trap months (11662.13 trap days, of which 271.16 trap days were unrepresentative of a normal catch). We describe the sampling in greater detail in
We processed the samples at the Zoological Museum of the University of Turku, Finland. We separated the ichneumonoid wasps (families Ichneumonidae and Braconidae) from the Malaise samples, then pinned the subfamily Rhyssinae and sorted specimens into species. We did not pin the hand netted rhyssines; instead, we stored these specimens individually in 96% ethanol. The samples are deposited at the Zoological Museum.
Layer photographs were taken using a Canon 7 D mark 2 digital camera, attached to an Olympus SZX 16 stereomicroscope. Photographs were captured using the programmes Deep Focus 3.1 and Quick Photo Camera 2.3. Photographs were finally combined with the program Zerene and edited in Photoshop CC. Additional images were acquired at
Because earlier diagnostic characters (
Morphological terminology largely follows
We generated an identification key automatically, based on the nine diagnostic characters (
Diagnostic characters for the thirteen known Afrotropical rhyssine species. The figure shows the frons median carinae (A converge, B diverge, C absent), frons lateral carinae (A present, B absent), hypostomal flange (D wider than second maxillary palp, E narrower than or comparable to second maxillary palp), subalar prominence (F flanged, G no flange), mesopleuron margin (H flanged, G no flange) and female apical horn (I ellipse, J half-ellipse). Not shown are the epicnemial carina (laterally absent, only just reaches the mesopleuron, reaches high onto the mesopleuron), areolet (present, absent) and tergite 3 structure (mostly striate, mostly punctate, mostly smooth). The images are of http://mus.utu.fi/ZMUT.5766, http://mus.utu.fi/ZMUT.2520 (A, F), http://mus.utu.fi/ZMUT.5853 (B, D, H, J), http://mus.utu.fi/ZMUT.5788 (C, E, I) and http://mus.utu.fi/ZMUT.5663 (G). Image F has been flipped horizontally.
We caught 448 rhyssines by Malaise sampling and eight by hand netting. They belonged to six species of which two are new. We provide a key to all 13 known Afrotropical species below. We also provide diagnostic characters and descriptions or updates to descriptions for all species. The key, diagnostic characters, and descriptions are also available in table form in the supplementary material (
1 | Fore wing with a closed areolet (Fig. |
Megarhyssa babaulti Seyrig, 1937 |
– | Fore wing without an areolet (Fig. |
2 |
2 | Subalar prominence with a lateral flange (Fig. |
Epirhyssa uelensis Benoit, 1951 |
– | Subalar prominence without a lateral flange (Fig. |
3 |
3 | Dorsal margin of mesopleuron with a raised flange (Fig. |
4 |
– | Dorsal margin of mesopleuron without a raised flange (Fig. |
5 |
4 | Apical horn of metasoma shaped like an ellipse in posterior view (Fig. |
Epirhyssa overlaeti Seyrig, 1937 |
– | Apical horn of metasoma shaped like a half-ellipse in posterior view (Fig. |
Epirhyssa ghesquierei Seyrig, 1937 |
5 | Epicnemial carina laterally absent, does not reach mesopleuron | 6 |
– | Epicnemial carina present on mesopleuron | 7 |
6 | Frons with median carinae that diverge before continuing towards the lateral ocelli (cf. Fig. |
Epirhyssa migratoria Seyrig, 1932 |
– | Frons smooth, without median carinae (Fig. |
Epirhyssa johanna sp. nov. |
7 | Frons with median carinae that diverge before continuing towards the lateral ocelli (cf. Fig. |
Epirhyssa villemantae Rousse & van Noort, 2014 |
– | Frons with median carinae that converge before the ocelli or without median carinae (Fig. |
8 |
8 | Tergite 3 densely striate (Fig. |
Epirhyssa quagga sp. nov. |
– | Tergite 3 mostly smooth or mostly punctate (Fig. |
9 |
9 | Tergite 3 mostly smooth | 10 |
– | Tergite 3 mostly punctate, over 50% of surface (Fig. |
11 |
10 | Epicnemial carina only just reaches the mesopleuron. [Hypostomal carina raised into a low flange, yellow-orange species with yellow and infuscate wings; Fig. |
Epirhyssa maynei Benoit, 1952 |
– | Epicnemial carina long, reaches the approximate height of the mesopleural pit. [Hypostomal carina raised into an elevated flange, yellow species with black mesosternum; Fig. |
Epirhyssa leroyi Benoit, 1951 |
11 | Hypostomal carina raised into an elevated flange (Fig. |
Epirhyssa brianfisheri Rousse & van Noort, 2014 |
– | Hypostomal carina raised into a low flange (Fig. |
12 |
12 | Apical horn of metasoma shaped like a half-ellipse in posterior view (Fig. |
Epirhyssa tombeaodiba Rousse & van Noort, 2014 |
– | Apical horn of metasoma shaped like an ellipse in posterior view (Fig. |
Epirhyssa shaka Rousse & van Noort, 2014 |
Diagnostic character traits used in the identification key. A Areolet (Megarhyssa babaulti holotype) B open areolet (http://mus.utu.fi/ZMUT.5788) C subalar prominence flange (http://mus.utu.fi/ZMUT.2520) D subalar prominence without flange and dorsal margin of mesopleuron with flange (http://mus.utu.fi/ZMUT.5853) E dorsal margin of mesopleuron without flange (
Subfamily Rhyssinae Morley, 1913
Diagnosis. The subfamily Rhyssinae can be recognised by a combination of the transverse rugae covering much of the mesoscutum; the short, broad mandibles and small, rectangular clypeus; the long ovipositor; and the female 8th metasomal tergite being produced posteriorly as a truncate horn-like projection. Other genera that present a potential confusion risk, such as Pseudorhyssa Merril (Pimplinae), Certonotus Kriechbaumer, and Apechoneura Kriechbaumer (Labeninae) are not present in the Afrotropical region.
Hierax Tosquinet, 1903: 255.
Rhyssonota Kriechbaumer, 1890: 489.
Sychnostigma Baltazar, 1961: 75.
The genus Epirhyssa is easily recognised in the Afrotropical region as the species lack the fore wing areolet (vein 3rs-m is missing), whereas the areolet is closed by vein 3rs-m in Megarhyssa Ashmead, the only other rhyssine genus found in the Afrotropical region.
Epirhyssa can be distinguished from other rhyssine genera by the lack of an areolet (cf. Rhyssella Rohwer, Lytarmes Cameron), the lack of an anterior glymma on tergite 1 (cf. Rhyssa Gravenhorst), the upper tooth being slightly wider than the lower and not subdivided (cf. Triancyra Baltazar, Myllenyxis Baltazar) and the pterostigma being angled where it meets the metacarpus (compared to gradually merging in Cyrtorhyssa Baltazar) (
Afrotropical region: Central African Republic, Cameroon, Democratic Republic of Congo, Madagascar, Nigeria, South Africa, Uganda.
Australasian region: Papua New Guinea.
Nearctic region: Mexico, U.S.A.
Neotropical region: Argentina, Brazil, Bolivia, Costa Rica, Cuba, Ecuador, Guatemala, Guyana, Nicaragua, Paraguay, Peru, Trinidad.
Oriental region: China, India, Indonesia, Japan, Malaysia, Myanmar, Philippines, Singapore, Vietnam.
Palaearctic region: Russia.
Type material: CENTRAL AFRICAN REPUBLIC:
• 1 ♀, holotype; Préfecture Sangha-Mbaéré, Réserve Spéciale de Forêt Dense de Dzanga-Sangha (12.7 km, 326 degrees NW of Bayanga); 3°00.27'N, 16°11.55'E; alt. 420 m; 13 May 2001; Simon van Noort leg.; Sweep; CAR01–S158; Lowland rainforest;
Known material: One specimen (1 ♀, see
This species can be distinguished from other Afrotropical Rhyssinae by the combination of an elevated hypostomal flange, the absence of a raised flange on the dorsal margin of the mesopleuron, an elliptical apical horn of the metasoma, and a finely punctate (over 50% of surface) tergite 3. In practice its colour pattern makes it instantly recognisable.
Head : frons with median carinae converging before continuing towards median ocellus, without lateral carinae; hypostomal carina raised into an elevated flange, its height greater than the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence without a lateral flange; mesopleuron without a flange along the dorsal margin; epicnemial carina reaches the approximate height of the mesopleural pit.
Metasoma : tip of apical horn elliptical in posterior view; tergite 3 punctate.
Type material: DEMOCRATIC REPUBLIC OF CONGO:
• 1 ♀, holotype; Eala [0°4.22'N, 18°18.15'E]; Nov. 1935; “J. Ghesquière”; “R. Dét. G 3330”;
• 1 ♂, paratype; Bambesa; Dec. 1946; “P.L. Benoit”;
Non-type material: CAMEROON:
• 1 ♂; Korup; Dec. 1980–Jan. 1981; Mrs D. Jackson leg.;
• 1 ♂; Korup; 1981; Mrs D. Jackson leg.;
UGANDA:
• 1 ♀; Kibale National Park, Kanyawara, Site R93, Malaise trap R93T1; 0.5653N, 30.3568E (WGS84); alt. 1510 m (GPS, WGS84); 23 Sep. 2014–7 Oct. 2014; Tapani Hopkins leg.;
• 1 ♀; same data as preceding; Site K30S, Malaise trap K30ST4; 0.5414N, 30.3755E (WGS84); alt. 1420 m (GPS, WGS84); 25 Aug. 2015–11 Sep. 2015;
• 1 ♀; same data as preceding; Site K15, Malaise trap K15T2; 0.5843N, 30.3644E (WGS84); alt. 1470 m (GPS, WGS84); 4 May 2015–20 May 2015;
• 1 ♀; same data as preceding; Site CC, Malaise trap CCT1; 0.5497N, 30.3673E (WGS84); alt. 1450 m (GPS, WGS84); 15 Dec. 2014–29 Dec. 2014;
• 1 ♀; same data as preceding; 24 Feb. 2015–10 Mar. 2015;
• 1 ♀; same data as preceding; 13 Jan. 2015–27 Jan. 2015;
• 1 ♂; Kibale National Park, Kanyawara, Site K31, Malaise trap K31T4; 0.5362N, 30.3486E (WGS84); alt. 1460 m (GPS, WGS84); 29 Dec. 2014–16 Jan. 2015; Tapani Hopkins leg.;
• 1 ♂; same data as preceding; Site CC, Malaise trap CCT1; 0.5497N, 30.3673E (WGS84); alt. 1450 m (GPS, WGS84); 30 Jun. 2015–14 Jul. 2015;
Non-type material (only diagnostic characters checked): UGANDA:
• 109 ♀; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
• 47 ♂; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
Known material: 168 specimens (164 Ugandan, 4 other):
112 ♀, 44 ♂; Ugandan specimens caught by Malaise trap, data above and also in supplementary material (
3 ♀, 5 ♂; Ugandan hand-netted specimens, data above and also in supplementary material (
1 ♀, 3 ♂; see
This species can be distinguished from other Afrotropical Rhyssinae by the combination of a half-elliptical apical horn of the metasoma and a mostly smooth tergite 3.
Head : frons with diverging median carinae, without clear lateral carinae; hypostomal carina raised into an elevated flange, its height greater than the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence without a lateral flange; mesopleuron with an elevated flange along the dorsal margin; epicnemial carina reaches the approximate height of the mesopleural pit.
Metasoma : tip of apical horn half-elliptical in posterior view; tergite 3 mostly smooth.
Apart from the diagnosis, we provide the following additional or updated character traits to the description in
Female.
Body length 11.4 mm–17.2 mm. Frons rugulose or smooth, often with more or less distinct rugae that fan out from the median carinae towards the ocelli. Antenna with 32–34 flagellar segments. Tergites mostly smooth, but with variable structure on some tergites (4–7 pubescent and anterior margins of 3–5 slightly punctate or striate in Ugandan specimens, 3–6 shallowly punctate with anterior striations in other specimens), tergite 1 2.2–2.5 times as long as apically wide. The Ugandan specimens are more orange than yellow in colour, generally have no dark spots on the lateral lobes of the mesoscutum, and the colour of their interocellar area varies from orange (most frequent) to black.
Male. Body length 11.5 mm–14.1 mm. Antenna with 31–33 flagellar segments. Anterior margin of tergite 3 sometimes neither punctate nor striate. Tergite 1 2.5–3.0 times as long as apically wide. Males are smaller than females on average.
Democratic Republic of Congo, Cameroon. New record: Uganda.
In Uganda, this species was most abundantly caught in primary forest near decaying wood, during the dry season (
Epirhyssa ghesquierei female (http://mus.utu.fi/ZMUT.5853), a species found in Uganda. 7 Habitus 8 face and clypeus 9 frons 10 hypostomal flange 11 mesopleuron dorsal margin 12 apical horn of metasoma 13 tergites 1–7. Scale bars: 0.5 mm (8–13), 1 mm (7).
Type material: UGANDA:
• 1 ♀, holotype; Kibale National Park, Kanyawara, Site R03, Malaise trap R03T2; 0.5403N, 30.3608E (WGS84); alt. 1490 m (GPS, WGS84); 7 May 2015–21 May 2015; Tapani Hopkins leg.;
Known material: One specimen (1 ♀, Ugandan specimen, data above).
This species can be distinguished from other Afrotropical Rhyssinae by the combination of a smooth frons without median carinae and a laterally absent epicnemial carina. No other species has the same colour pattern.
Head : frons without median carinae, without lateral carinae; hypostomal carina raised into a low flange, its height slightly less than or equivalent to the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence without a lateral flange; mesopleuron without a flange along the dorsal margin; epicnemial carina laterally absent.
Metasoma : tip of apical horn elliptical in posterior view; tergite 3 mostly smooth.
Body length 8.4 mm.
Head : Frons without median carinae, without lateral carinae, smooth or very faintly striate. Occipital carina interrupted dorsally and interrupted or extremely faint near hypostomal carina. Hypostomal carina raised into a low flange, its height slightly less than or equivalent to maximum width of second maxillary palp segment. Face smooth or very faintly punctate. Clypeus sparsely punctate, with a median apical tubercle. Antenna with 28 flagellar segments.
Mesosoma : Subalar prominence without a lateral flange. Mesopleuron without a flange along the dorsal margin. Epicnemial carina laterally absent. Fore wing with 2m-cu distal to rs-m.
Metasoma : Tip of apical horn elliptical (flattened ellipse) in posterior view. Tergites mostly smooth, anterior of tergites 2–6 medially striate, tergite 1 1.2 times as long as apically wide.
Colour : General colour orange. Other colour: white face, lower 1/4 of genae, lateral frons, black mandibles, median frons, occiput, upper genae, and median spot on apical half of tergite 6 and entire tergite 7, dark brown hind tarsi. Antennae black. Ovipositor sheaths dark testaceous. Wings hyaline, faintly infuscate near apex.
Male. Unknown.
Dedicated to Johanna Hopkins, the first author’s wife. This species is known from only one, quite exceptional, female specimen.
Uganda.
Only one specimen was caught during 382 trap months of sampling, in a habitat (successional forest logged 2002–2004) that generally yielded few rhyssines.
Epirhyssa johanna female (holotype http://mus.utu.fi/ZMUT.4920), a new species from Uganda. 14 habitus 15 face and clypeus 16 frons 17 hypostomal flange 18 mesopleuron dorsal margin 19 apical horn of metasoma 20 tergites 1–7. Scale bars: 0.5 mm (15–20), 1 mm (14).
Type material: DEMOCRATIC REPUBLIC OF CONGO:
• 1 ♀, holotype; Bambesa [03°28'N 25°43'E]; Dec. 1933; “J.V. Leroy”;
• 1 ♀, paratype; Ubangui-Bumba; Dec. 1939; H. de Saeger;
Known material: Two specimens (2 ♀, see
This species can be distinguished from other Afrotropical Rhyssinae by the combination of converging median carinae on the frons, the absence of lateral carinae on the frons, an epicnemial carina that reaches high onto the mesopleuron, and a mostly smooth tergite 3. No other species is known to have a black mesosternum.
Head : frons with median carinae converging before continuing towards median ocellus, without lateral carinae; hypostomal carina raised into an elevated flange, its height slightly greater than the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence without a lateral flange; mesopleuron without a flange along the dorsal margin; epicnemial carina reaches the approximate height of the mesopleural pit.
Metasoma : tip of apical horn elliptical in posterior view; tergite 3 mostly smooth.
Democratic Republic of Congo.
Non-type material: CENTRAL AFRICAN REPUBLIC:
• 1 ♂; Préfecture Sangha-Mbaéré, Parc National de Dzanga-Ndoki (38.6 km, 173 degrees S of Lidjombo); 2°21.60'N, 16°09.20'E; alt. 350 m; 27 May 2001; Simon van Noort leg.; Handnet; CAR01-H25; Lowland rainforest;
Known material: Three specimens (0 Ugandan, 3 other):
1 ♂, holotype; see
1 ♂, paratype; see
1 ♂; see
This species can be distinguished from other Afrotropical Rhyssinae by the epicnemial carina only just reaching the mesopleuron. The fore wing colour pattern makes it instantly recognisable.
Head : frons with median carinae converging before continuing towards median ocellus, without lateral carinae; hypostomal carina raised into a low flange, its height slightly less than or equivalent to the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence without a lateral flange; mesopleuron without a flange along the dorsal margin; epicnemial carina short, barely extending onto mesopleuron.
Metasoma : tip of apical horn of unknown shape (no females are known); tergite 3 mostly smooth.
Type material: MADAGASCAR:
• 1 ♀, paralectotype; Rogez, Forêt Cote Est; Mar. 1932; “A Seyrig”;
Known material: Eight specimens (0 Ugandan, 8 other):
1 ♀, lectotype; see
3 ♂; see
1 ♀, paralectotype; previously unpublished specimen, data above in material examined.
1 ♀, 2 ♂; previously unpublished specimens, same data as paralectotype except collection dates Apr. 1931, Sep. 1932, Dec. 1932;
This species can be distinguished from other Afrotropical Rhyssinae by the combination of diverging median carinae on the frons, the lateral absence of the epicnemial carina, and an open areolet. Its yellow and black colour pattern is distinctive.
Head : frons with median carinae diverging towards ocelli, without lateral carinae; hypostomal carina raised into a low flange, its height slightly less than or equivalent to the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence without a lateral flange; mesopleuron without a flange along the dorsal margin; epicnemial carina laterally absent.
Metasoma : tip of apical horn elliptical in posterior view; tergite 3 mostly smooth.
Non-type material: UGANDA:
• 1 ♀; Kibale National Park, Kanyawara, Site CC, Malaise trap CCT1; 0.5497N, 30.3673E (WGS84); alt. 1450 m (GPS, WGS84); 10 Mar. 2015–24 Mar. 2015; Tapani Hopkins leg.;
• 1 ♀; same data as preceding; Site K31, Malaise trap K31T4; 0.5362N, 30.3486E (WGS84); alt. 1460 m (GPS, WGS84); 16 Jul. 2015–30 Jul. 2015;
• 1 ♀; same data as preceding; Site CC, Malaise trap CCT1; 0.5497N, 30.3673E (WGS84); alt. 1450 m (GPS, WGS84); 18 Nov. 2014–2 Dec. 2014;
• 1 ♀; same data as preceding; Site R93, Malaise trap R93T2; 0.5654N, 30.3593E (WGS84); alt. 1510 m (GPS, WGS84); 20 May 2015–1 Jun. 2015;
• 1 ♀; same data as preceding; Site HILL, Malaise trap HILLT1; 0.5486N, 30.3614E (WGS84); alt. 1520 m (GPS, WGS84); 30 Jul. 2015–13 Aug. 2015;
• 1 ♂; Kibale National Park, Kanyawara, Site K31, Malaise trap K31T2; 0.5427N, 30.3482E (WGS84); alt. 1460 m (GPS, WGS84); 9 Apr. 2015–23 Apr. 2015; Tapani Hopkins leg.;
• 1 ♂; same data as preceding; Site CC, Malaise trap CCT1; 0.5497N, 30.3673E (WGS84); alt. 1450 m (GPS, WGS84); 24 Feb. 2015–10 Mar. 2015;
• 1 ♂; same data as preceding; Site K31, Malaise trap K31T3; 0.5360N, 30.3469E (WGS84); alt. 1450 m (GPS, WGS84); 26 Aug. 2015–12 Sep. 2015;
Non-type material (only diagnostic characters checked): UGANDA:
• 56 ♀; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
• 14 ♂; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
• 1 unknown sex; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
Known material: 81 specimens (79 Ugandan, 2 other):
61 ♀, 17 ♂, 1 U; Ugandan specimens, data above and also in supplementary material (
1 ♀, holotype; see
1 ♀; see
Epirhyssa overlaeti male (http://mus.utu.fi/ZMUT.1280) and female (34: http://mus.utu.fi/ZMUT.2291, 37: http://mus.utu.fi/ZMUT.5766), a species found in Uganda. 32 Habitus 33 face and clypeus 34 frons 35 hypostomal flange 36 mesopleuron dorsal margin 37 apical horn of metasoma 38 tergites 1–7. Scale bars: 0.5 mm (33–38), 1 mm (32).
This species can be distinguished from other Afrotropical Rhyssinae by the combination of converging median carinae on the frons, lateral carinae on the frons, and the absence of a lateral flange on the subalar prominence. In practice its unique colour pattern and large size make it instantly recognisable.
Head : frons with strong median carinae converging before continuing towards median ocellus, with lateral carinae curving towards lateral ocelli; hypostomal carina raised into an elevated flange, its height slightly greater than the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence without a lateral flange; mesopleuron with a raised flange along the dorsal margin; epicnemial carina reaches the approximate height of the mesopleural pit.
Metasoma : tip of apical horn almost circular in posterior view; tergite 3 mostly smooth.
Body length 16.4 mm–25.1 mm. Males seem slightly smaller than females on average.
Head : Frons with strong median carinae converging before continuing towards median ocellus, with lateral carinae curving towards lateral ocelli, often with faint traces of lateral rugae. Occipital carina interrupted dorsally. Hypostomal carina raised into an elevated flange, its height slightly greater than maximum width of second maxillary palp segment. Face punctate. Clypeus punctate, without a clear median apical tubercle. Antenna with 38–40 flagellar segments.
Mesosoma : Subalar prominence without a lateral flange. Mesopleuron with a raised flange along dorsal margin. Epicnemial carina reaches approximate height of mesopleural pit. Fore wing with 2m-cu distal to rs-m.
Metasoma : Tergites 1–5 mostly smooth, 6–7 smooth or sparsely pubescent, tergite 1 1.8–2.2 times as long as apically wide.
Colour : General colour mottled dark testaceous, testaceous and pale yellow, pale yellow patches of tergites 1 and 2 are fused (cf. female). Antennae black. Wings hyaline, infuscate near apex.
Additional or updated characters. Apart from the diagnosis and description of the male, we provide the following additional or updated character traits to the description in
Female. Body length 14.7 mm–37.8 mm. Frons often with faint traces of lateral rugae. Face punctate, transversely rugulose-punctate or transversely striate. Antenna with 38–43 flagellar segments (38–41 in Ugandan specimens). Tergites 1–5 mostly smooth, 6–7 often pubescent, anterior margins of tergites 3–6 (sometimes only 4–5) sightly striate or punctate, tergite 1 1.4–1.8 times as long as apically wide. The colour patches vary in extent, with the pale yellow patches of tergite 1 fused in small individuals.
Democratic Republic of Congo, Cameroon. New record: Uganda.
In Uganda, this species was mostly caught in primary forest near decaying wood (
Epirhyssa overlaeti was earlier known from only two females. We describe the male for the first time.
Type material: UGANDA:
• 1 ♀, holotype; Kibale National Park, Kanyawara, Site CC, Malaise trap CCT1; 0.5497N, 30.3673E (WGS84); alt. 1450 m (GPS, WGS84); 11 Aug. 2015–25 Aug. 2015; Tapani Hopkins leg.;
• 1 ♀, paratype; same data as preceding; Site R03, Malaise trap R03T2; 0.5403N, 30.3608E (WGS84); alt. 1490 m (GPS, WGS84); 21 May 2015–4 Jun. 2015;
• 1 ♀, paratype; same data as preceding; Site K31, Malaise trap K31T3; 0.5360N, 30.3469E (WGS84); alt. 1450 m (GPS, WGS84); 23 May 2015–4 Jun. 2015;
• 1 ♀, paratype; same data as preceding; Site K30S, Malaise trap K30ST4; 0.5414N, 30.3755E (WGS84); alt. 1420 m (GPS, WGS84); 2 Dec. 2014–15 Dec. 2014;
• 1 ♀, paratype; same data as preceding; Site R01, Malaise trap R01T2; 0.5501N, 30.3561E (WGS84); alt. 1600 m (GPS, WGS84); 27 Mar. 2015–10 Apr. 2015;
• 1 ♂, paratype; Kibale National Park, Kanyawara, Site CC, Malaise trap CCT1; 0.5497N, 30.3673E (WGS84); alt. 1450 m (GPS, WGS84); 2 Jun. 2015–17 Jun. 2015; Tapani Hopkins leg.;
• 1 ♂, paratype; same data as preceding; Site K30S, Malaise trap K30ST2; 0.5392N, 30.3771E (WGS84); alt. 1480 m (GPS, WGS84); 5 May 2015–19 May 2015;
• 1 ♂, paratype; same data as preceding; Site K15, Malaise trap K15T4; 0.5845N, 30.3674E (WGS84); alt. 1510 m (GPS, WGS84); 4 May 2015–20 May 2015;
Non-type material (only diagnostic characters checked): UGANDA
• 18 ♀; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
• 20 ♂; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
Epirhyssa quagga female (holotype http://mus.utu.fi/ZMUT.5788), a new species from Uganda. 39 Habitus 40 face and clypeus 41 frons 42 hypostomal flange 43 mesopleuron dorsal margin 44 apical horn of metasoma 45 tergites 1–7. Scale bars: 0.5 mm (40–45), 1 mm (39).
Known material: 46 specimens (23 ♀, 23 ♂, Ugandan specimens, data above).
This species can be distinguished from other Afrotropical Rhyssinae by the distinctive pattern of striation on the frons and the densely striate tergite 3. No other species has the same colour pattern.
Head : frons without median carinae, without lateral carinae; hypostomal carina raised into a low flange, its height slightly less than or equivalent to the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence without a lateral flange; mesopleuron without a flange along the dorsal margin; epicnemial carina reaches the approximate height of the mesopleural pit.
Metasoma : tip of apical horn elliptical in posterior view; tergite 3 densely striate.
Body length 8.5 mm–14.3 mm (holotype 12.3 mm).
Head : Frons without clear median carinae, without lateral carinae, with faint median rugae that fan out towards ocelli. Occipital carina interrupted dorsally. Hypostomal carina raised into a low flange, its height slightly less than or equivalent to maximum width of second maxillary palp segment. Face densely and deeply punctate. Clypeus with little or no punctation, with a median apical tubercle. Antenna with 30–33 flagellar segments (32 in holotype).
Mesosoma : Subalar prominence without a lateral flange. Mesopleuron without a flange along dorsal margin. Epicnemial carina reaches approximate height of mesopleural pit. Fore wing with 2m-cu varying from clearly distal to opposite rs-m.
Metasoma : Tip of apical horn elliptical in posterior view. Tergites 1–5 with dense, light, predominantly longitudinal striation and punctation, 6–7 smoother and more pubescent, tergite 1 1.9–2.4 times as long as apically wide (1.9 in holotype).
Colour : General colour mottled black and white with metasoma orange testaceous from tergite 3 onwards. Hind tibia dark brown. Antennae black. Ovipositor sheaths black to dark testaceous. Wings hyaline.
Variation : Colour of the dark patches of the legs varies from entirely black to testaceous, the black and white metasomal colour extends onto tergite 3 in some individuals.
Male.
Similar to female. Body length 7.9 mm–10.4 mm. T1 3.2–3.6. Antenna with 29–33 flagellar segments. Males are smaller than females on average.
Refers to the colour pattern which is reminiscent of the plains zebra, especially its extinct subspecies, the quagga.
Uganda.
In Uganda, this species was most abundantly caught during the dry season (
Type material: SOUTH AFRICA:
• 1 ♀, holotype; Natal, 2831 Dd Umlalazi Nat. Res., 1.5 km E of Mtunzini; 28°57'S, 31°45'E; Nov. 1978; R. M. Miller leg.; indigenous forest; Malaise trap;
Known material: One specimen (1 ♀, see
This species can be distinguished from other Afrotropical Rhyssinae by the combination of a low hypostomal flange, an elliptical apical horn of the metasoma, and a punctate (over 50% of surface) tergite 3.
Head : frons with median carinae converging before continuing towards median ocellus, without lateral carinae; hypostomal carina raised into a low flange, its height slightly less than or equivalent to the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence without a lateral flange; mesopleuron without a flange along the dorsal margin; epicnemial carina reaches the approximate height of the mesopleural pit.
Metasoma : tip of apical horn elliptical in posterior view; tergite 3 punctate.
Non-type material: UGANDA:
• 1 ♀; Kibale National Park, Kanyawara, Site K30, Malaise trap K30T2; 0.5566N, 30.3633E (WGS84); alt. 1490 m (GPS, WGS84); 26 Aug. 2015–9 Sep. 2015; Tapani Hopkins leg.;
• 1 ♀; same data as preceding; Site K15, Malaise trap K15T1; 0.5850N, 30.3641E (WGS84); alt. 1490 m (GPS, WGS84); 20 Apr. 2015–4 May 2015;
• 1 ♀; same data as preceding; Site HILL, Malaise trap HILLT2; 0.5478N, 30.3619E (WGS84); alt. 1510 m (GPS, WGS84); 4 Jun. 2015–18 Jun. 2015;
• 1 ♀; same data as preceding; Site K30S, Malaise trap K30ST3; 0.5378N, 30.3777E (WGS84); alt. 1480 m (GPS, WGS84); 19 May 2015–2 Jun. 2015;
• 1 ♀; same data as preceding; Site K30, Malaise trap K30T3; 0.5590N, 30.3617E (WGS84); alt. 1540 m (GPS, WGS84); 14 Jul. 2015–28 Jul. 2015;
• 1 ♀; same data as preceding; Site R93, Malaise trap R93T2; 0.5654N, 30.3593E (WGS84); alt. 1510 m (GPS, WGS84); 9 Mar. 2015–23 Mar. 2015;
Known material: 10 specimens (6 Ugandan, 4 other):
6 ♀; Ugandan specimens, data above and also in supplementary material (
1 ♀, holotype; see
1 ♀, paratype; see
1 ♀, paratype; see
1 ♀, paratype; see
This species can be distinguished from other Afrotropical Rhyssinae by the combination of a half-elliptical apical horn of the metasoma and a punctate (over 50% of surface) tergite 3. Epirhyssa uelensis is also predominantly yellow with black spots, but its subalar prominence has a lateral flange.
Head : frons with weak median carinae converging before continuing towards median ocellus, without lateral carinae; hypostomal carina raised into a low flange, its height slightly less than or equivalent to the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence without a lateral flange; mesopleuron without a flange along the dorsal margin; epicnemial carina reaches the approximate height of the mesopleural pit.
Metasoma : tip of apical horn half-elliptical in posterior view; tergite 3 densely punctate.
Apart from the diagnosis, we provide the following additional or updated character traits to the description in
Female.
Body length 7.2 mm–12.7 mm. Frons without rugae or with faint lateral rugae curving towards lateral ocelli. Face punctate or transversely rugulose-punctate. Clypeus longitudinally strigose and sparsely punctate. Antenna with 28–29 flagellar segments. Tergites 1 mostly smooth, 2–7 punctate (2 sometimes only punctate laterally), anterior margins of 5–6 often striate, tergite 1 1.2–1.5 times as long as apically wide. The Ugandan specimens have black anterior median spots on tergites 1–7 (ranging from very small on tergite 1 to reaching posterior margin on tergite 7), not just on tergites 4–7.
Cameroon. New record: Uganda.
Epirhyssa tombeaodiba female (http://mus.utu.fi/ZMUT.5663, 52: http://mus.utu.fi/ZMUT.3234), a species found in Uganda. 50 Habitus 51 face and clypeus 52 frons 53 hypostomal flange 54 mesopleuron dorsal margin 55 apical horn of metasoma 56 tergites 1–7. Scale bars: 0.5 mm (51–56), 1 mm (50).
Epirhyssa gavinbroadi Rousse & van Noort, 2014, syn. nov.
Type material: CAMEROON:
• 1 ♀, holotype of E. gavinbroadi; Nkoemvon [Nko’emvon]; 02°48'N, 11°08'E; 16 Mar. 1980–4 May 1980;
Non-type material: UGANDA
• 1 ♀; Kibale National Park, Kanyawara, Site K31, Malaise trap K31T4; 0.5362N, 30.3486E (WGS84); alt. 1460 m (GPS, WGS84); 16 Jul. 2015–30 Jul. 2015; Tapani Hopkins leg.;
• 1 ♀; same data as preceding; Site K30, Malaise trap K30T3; 0.5590N, 30.3617E (WGS84); alt. 1540 m (GPS, WGS84); 21 Apr. 2015–5 May 2015;
• 1 ♀; same data as preceding; Site K31, Malaise trap K31T4; 0.5362N, 30.3486E (WGS84); alt. 1460 m (GPS, WGS84); 30 Jan. 2015–13 Feb. 2015;
• 1 ♀; same data as preceding; Site K13, Malaise trap K13T1; 0.5932N, 30.3598E (WGS84); alt. 1460 m (GPS, WGS84); 13 Jul. 2015–27 Jul. 2015;
• 1 ♀; same data as preceding; Site K30, Malaise trap K30T2; 0.5566N, 30.3633E (WGS84); alt. 1490 m (GPS, WGS84); 2 Dec. 2014–15 Dec. 2014;
• 1 ♀; same data as preceding; Site CC, Malaise trap CCT1; 0.5497N, 30.3673E (WGS84); alt. 1450 m (GPS, WGS84); 27 Jan. 2015–10 Feb. 2015;
• 1 ♀; same data as preceding; 24 Feb. 2015–10 Mar. 2015;
• 1 ♀; same data as preceding; Site K30S, Malaise trap K30ST4; 0.5414N, 30.3755E (WGS84); alt. 1420 m (GPS, WGS84); 30 Jun. 2015–14 Jul. 2015;
• 1 ♂; Kibale National Park, Kanyawara, Site R93, Malaise trap R93T2; 0.5654N, 30.3593E (WGS84); alt. 1510 m (GPS, WGS84); 20 May 2015–1 Jun. 2015; Tapani Hopkins leg.;
• 1 ♂; same data as preceding; Site K13, Malaise trap K13T1; 0.5932N, 30.3598E (WGS84); alt. 1460 m (GPS, WGS84); 4 May 2015–20 May 2015;
Non-type material (only diagnostic characters checked): UGANDA:
• 150 ♀; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
Known material: 167 specimens (160 Ugandan, 7 other):
158 ♀, 2 ♂; Ugandan specimens, data above and also in supplementary material (
1 ♀, holotype of E. gavinbroadi; see
1 ♀, holotype of E. uelensis; see
1 ♂, paratype of E. uelensis; see
1 ♀; see
2 ♀; see
1 ♀; see
Epirhyssa uelensis female (http://mus.utu.fi/ZMUT.2520), a species found in Uganda. 57 Habitus 58 face and clypeus 59 frons 60 hypostomal flange 61 mesopleuron dorsal margin 62 apical horn of metasoma 63 tergites 1–7. Scale bars: 0.5 mm (58–63), 1 mm (57).
This species can be distinguished from other Afrotropical Rhyssinae by its subalar prominence having a lateral flange. Epirhyssa tombeaodiba is also predominantly yellow with black spots but lacks the flange.
Head : frons with median carinae converging before continuing towards median ocellus, with lateral carinae curving towards lateral ocelli; hypostomal carina raised into an elevated flange, its height greater than the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence with a lateral flange; mesopleuron with a raised flange along the dorsal margin; epicnemial carina reaches the approximate height of the mesopleural pit.
Metasoma : tip of apical horn elliptical in posterior view; tergite 3 mostly smooth.
Apart from the diagnosis, we provide the following additional or updated character traits to the description in
Female. Body length 9.8 mm–15.7 mm (15.7 mm E. gavinbroadi holotype, 9.8 mm–15.1 mm other specimens). Frons with more or less distinct rugae parallel to the lateral carinae, often as distinct as the lateral carinae. Clypeus sparsely punctate or smooth. Antenna with 30–35+ flagellar segments (30–33 in Ugandan specimens, at least 35 in E. gavinbroadi type, 30–34 in other specimens). Fore wing with 2m-cu varying from clearly distal to only just distal to rs-m. Tip of apical horn with an extra dorsal projection in one aberrant female (http://mus.utu.fi/ZMUT.5844). Tergites mostly smooth, except with varied amount of punctation on tergites 3–7 (anteriorly punctate 3–6 in many Ugandan specimens, punctate 3–6 in most non-Ugandan specimens, punctate apex of 5 and 6–7 in E. gavinbroadi holotype), anterior margins of 3–5 or 3–6 often striate, tergite 1 1.2–1.7 times as long as apically wide (1.7 in E. gavinbroadi holotype, 1.2–1.5 in other specimens). The Ugandan specimens have black posterior margins of tergites 2–7 (sometimes faint or patchy, especially on anterior tergites) and median stripes on 4–7 (strongest on 5–6). Non-Ugandan specimens sometimes have black median stripes on 5–7, or 3–6 in the E. gavinbroadi holotype.
Male. Body length 8.5 mm–8.7 mm. Antenna with 30 flagellar segments. Tergites 4–7 faintly pubescent, tergite anterior margins smooth, or lightly striate and punctate, tergite 1 1.5–2.0 times as long as apically wide. The two Ugandan males are smaller than females on average.
Democratic Republic of Congo, Cameroon. New record: Uganda.
In Uganda, this species was most abundantly caught in primary forest near decaying wood, during the dry season (
This species is quite variable, but we are unable to find morphological characters that would reliably split it into more than one species. We propose that Epirhyssa gavinbroadi (of which there is only one specimen) is a synonym of E. uelensis. It was mainly distinguished from E. uelensis by the punctate clypeus and slender tergite 1, but the Ugandan specimens generally have a punctate clypeus and a stout tergite 1. These two characters also vary considerably in the Ugandan specimens and seem to represent intraspecific variation. The Ugandan material has a very skewed sex ratio with 158 female specimens collected and only two males.
Type material: NIGERIA:
• 1 ♀, holotype; “Ilorin Prov.” [06°48'N, 05°18'E]; “18 Jun. 192” [18 Jun 1921?]; “De. G.W.S. Macfie” leg.; “Pres. by Imp. Bur. Ent. 1921–129”;
Known material: One specimen (1 ♀, see
This species can be distinguished from other Afrotropical Rhyssinae by the combination of lateral carinae on the frons and the absence of a raised flange on the dorsal margin of the mesopleuron. No other species has the same colour pattern.
Head : frons with median carinae diverging towards ocelli, with lateral carinae curving towards lateral ocelli; hypostomal carina raised into an elevated flange, its height greater than the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence without a lateral flange; mesopleuron without a flange along the dorsal margin; epicnemial carina reaches the approximate height of the mesopleural pit.
Metasoma : tip of apical horn elliptical in posterior view; tergite 3 mostly smooth.
Thalessa Holmgren, 1859: 122.
Megalorhyssa Shulz, 1906: 115.
Eurhyssa Derksen, 1941: 721.
The genus Megarhyssa is easily recognised in the Afrotropical region by the presence of the fore wing areolet (vein 3rs-m is present), whereas vein 3rs-m is missing in Epirhyssa, the only other rhyssine genus found in the Afrotropical region.
Megarhyssa can be distinguished from other rhyssine genera by the presence of an areolet (cf. Triancyra), the lack of an anterior glymma on tergite 1 (cf. Rhyssa), the upper tooth not being subdivided (cf. Myllenyxis), tergite 1 having anterior lateral carinae (cf. Cyrtorhyssa which lacks carinae), the occipital carina joining the hypostomal carina some distance from the mandible base (cf. Lytarmes) and tergites 3–6 not being transversely, non-uniformly aciculate (cf. Rhyssella) (
Afrotropical region: Democratic Republic of Congo.
The genus is cosmopolitan with the largest number of species found in the Oriental and Palaearctic regions.
Type material: DEMOCRATIC REPUBLIC OF CONGO:
• 1 ♀, holotype; “reg. Lac Kivu”, “Kadjudju” [Kajuju, 02°09'S, 28°54'E]; 1932;
Known material: One specimen (1 ♀, see
This species can be distinguished from other Afrotropical Rhyssinae by the presence of a closed areolet. No other species has the same colour pattern.
Head : frons with median carinae diverging towards ocelli, without lateral carinae; hypostomal carina raised into an elevated flange, its height slightly greater than the maximum width of the second maxillary palp segment.
Mesosoma : subalar prominence without a lateral flange; mesopleuron without a flange along the dorsal margin; epicnemial carina laterally absent.
Metasoma : tip of apical horn almost circular in posterior view; tergite 3 mostly smooth.
Sampling tropical ichneumonids for a year with numerous Malaise traps gave us an unprecedented sample size. We caught 456 individuals of the subfamily Rhyssinae, which in the Afrotropical region was earlier known from only 30 published specimen records (
Our results strongly support the idea that the Afrotropical region contains a large number of rhyssine species, most of which have simply not been discovered due to insufficient sampling (
Our results also support the claim by
This study was partly funded by the Finnish Cultural Foundation, Oskar Öflunds Stiftelse, the Helsinki Entomology Society and Waldemar von Frenckells stiftelse (grants to TH). Isaiah Mwesige helped maintain the Malaise traps and ably carried out a whole range of other field work. Our field research was supported by the staff of the Makerere University Biological Field Station. The Muséum national d’Histoire naturelle (Paris collections, Agnièle Touret-Alby) provided images of M. babaulti, and the Royal Museum for Central Africa (Stéphane Hanot) provided images of E. leroyi and data on E. ghesquierei. Countless people helped process the samples, including the staff of the Zoological Museum of the University of Turku, students of the university, and school pupils from throughout the Turku region. These contributions are gratefully acknowledged.
The required research permits were issued by the Uganda National Council of Science and Technology and the Uganda Wildlife Authority.