Corresponding author: Tapani Hopkins (
Academic editor: Bernardo Santos
Tropical forest invertebrates, such as the parasitoid wasp family
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.
Like many taxa, the parasitoid wasps of the family
The ichneumonid subfamily
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
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
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 (
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. |
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– | Fore wing without an areolet (Fig. |
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2 | Subalar prominence with a lateral flange (Fig. |
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– | Subalar prominence without a lateral flange (Fig. |
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3 | Dorsal margin of mesopleuron with a raised flange (Fig. |
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– | Dorsal margin of mesopleuron without a raised flange (Fig. |
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4 | Apical horn of metasoma shaped like an ellipse in posterior view (Fig. |
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– | Apical horn of metasoma shaped like a half-ellipse in posterior view (Fig. |
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5 | Epicnemial carina laterally absent, does not reach mesopleuron |
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– | Epicnemial carina present on mesopleuron |
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6 | Frons with median carinae that diverge before continuing towards the lateral ocelli (cf. Fig. |
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– | Frons smooth, without median carinae (Fig. |
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7 | Frons with median carinae that diverge before continuing towards the lateral ocelli (cf. Fig. |
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– | Frons with median carinae that converge before the ocelli or without median carinae (Fig. |
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8 | Tergite 3 densely striate (Fig. |
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– | Tergite 3 mostly smooth or mostly punctate (Fig. |
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9 | Tergite 3 mostly smooth |
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– | Tergite 3 mostly punctate, over 50% of surface (Fig. |
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10 | Epicnemial carina only just reaches the mesopleuron. [Hypostomal carina raised into a low flange, yellow-orange species with yellow and infuscate wings; Fig. |
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– | Epicnemial carina long, reaches the approximate height of the mesopleural pit. [Hypostomal carina raised into an elevated flange, yellow species with black mesosternum; Fig. |
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11 | Hypostomal carina raised into an elevated flange (Fig. |
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– | Hypostomal carina raised into a low flange (Fig. |
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12 | Apical horn of metasoma shaped like a half-ellipse in posterior view (Fig. |
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– | Apical horn of metasoma shaped like an ellipse in posterior view (Fig. |
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Diagnostic character traits used in the identification key.
The genus
• 1 ♀, holotype; Préfecture Sangha-Mbaéré, Réserve Spéciale de Forêt Dense de Dzanga-Sangha (12.7 km, 326 degrees NW of Bayanga);
This species can be distinguished from other Afrotropical
Central African Republic.
• 1 ♀, holotype; Eala [
• 1 ♂, paratype; Bambesa; Dec. 1946; “P.L. Benoit”;
• 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;
• 1 ♀; same data as preceding; Site K30S, Malaise trap K30ST4;
• 1 ♀; same data as preceding; Site K15, Malaise trap K15T2;
• 1 ♀; same data as preceding; Site CC, Malaise trap CCT1;
• 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;
• 1 ♂; same data as preceding; Site CC, Malaise trap CCT1;
• 109 ♀; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
• 47 ♂; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
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
Apart from the diagnosis, we provide the following additional or updated character traits to the description in
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.
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 (
• 1 ♀, holotype; Kibale National Park, Kanyawara, Site R03, Malaise trap R03T2;
This species can be distinguished from other Afrotropical
Body length 8.4 mm.
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.
• 1 ♀, holotype; Bambesa [
• 1 ♀, paratype; Ubangui-Bumba; Dec. 1939; H. de Saeger;
This species can be distinguished from other Afrotropical
Democratic Republic of Congo.
• 1 ♂; Préfecture Sangha-Mbaéré, Parc National de Dzanga-Ndoki (38.6 km, 173 degrees S of Lidjombo);
1 ♂, holotype; see
1 ♂, paratype; see
1 ♂; see
This species can be distinguished from other Afrotropical
Democratic Republic of Congo, Central African Republic.
• 1 ♀, paralectotype; Rogez, Forêt Cote Est; Mar. 1932; “A Seyrig”;
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
Madagascar.
• 1 ♀; Kibale National Park, Kanyawara, Site CC, Malaise trap CCT1;
• 1 ♀; same data as preceding; Site K31, Malaise trap K31T4;
• 1 ♀; same data as preceding; Site CC, Malaise trap CCT1;
• 1 ♀; same data as preceding; Site R93, Malaise trap R93T2;
• 1 ♀; same data as preceding; Site HILL, Malaise trap HILLT1;
• 1 ♂; Kibale National Park, Kanyawara, Site K31, Malaise trap K31T2;
• 1 ♂; same data as preceding; Site CC, Malaise trap CCT1;
• 1 ♂; same data as preceding; Site K31, Malaise trap K31T3;
• 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.;
61 ♀, 17 ♂, 1 U; Ugandan specimens, data above and also in supplementary material (
1 ♀, holotype; see
1 ♀; see
This species can be distinguished from other Afrotropical
Body length 16.4 mm–25.1 mm. Males seem slightly smaller than females on average.
Democratic Republic of Congo, Cameroon. New record: Uganda.
In Uganda, this species was mostly caught in primary forest near decaying wood (
• 1 ♀, holotype; Kibale National Park, Kanyawara, Site CC, Malaise trap CCT1;
• 1 ♀, paratype; same data as preceding; Site R03, Malaise trap R03T2;
• 1 ♀, paratype; same data as preceding; Site K31, Malaise trap K31T3;
• 1 ♀, paratype; same data as preceding; Site K30S, Malaise trap K30ST4;
• 1 ♀, paratype; same data as preceding; Site R01, Malaise trap R01T2;
• 1 ♂, paratype; Kibale National Park, Kanyawara, Site CC, Malaise trap CCT1;
• 1 ♂, paratype; same data as preceding; Site K30S, Malaise trap K30ST2;
• 1 ♂, paratype; same data as preceding; Site K15, Malaise trap K15T4;
• 18 ♀; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
• 20 ♂; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
This species can be distinguished from other Afrotropical
Body length 8.5 mm–14.3 mm (holotype 12.3 mm).
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
Uganda.
In Uganda, this species was most abundantly caught during the dry season (
• 1 ♀, holotype; Natal, 2831 Dd Umlalazi Nat. Res., 1.5 km E of Mtunzini;
This species can be distinguished from other Afrotropical
South Africa (KwaZulu-Natal).
• 1 ♀; Kibale National Park, Kanyawara, Site K30, Malaise trap K30T2;
• 1 ♀; same data as preceding; Site K15, Malaise trap K15T1;
• 1 ♀; same data as preceding; Site HILL, Malaise trap HILLT2;
• 1 ♀; same data as preceding; Site K30S, Malaise trap K30ST3;
• 1 ♀; same data as preceding; Site K30, Malaise trap K30T3;
• 1 ♀; same data as preceding; Site R93, Malaise trap R93T2;
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
Apart from the diagnosis, we provide the following additional or updated character traits to the description in
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.
• 1 ♀, holotype of
• 1 ♀; Kibale National Park, Kanyawara, Site K31, Malaise trap K31T4;
• 1 ♀; same data as preceding; Site K30, Malaise trap K30T3;
• 1 ♀; same data as preceding; Site K31, Malaise trap K31T4;
• 1 ♀; same data as preceding; Site K13, Malaise trap K13T1;
• 1 ♀; same data as preceding; Site K30, Malaise trap K30T2;
• 1 ♀; same data as preceding; Site CC, Malaise trap CCT1;
• 1 ♀; same data as preceding; 24 Feb. 2015–10 Mar. 2015;
• 1 ♀; same data as preceding; Site K30S, Malaise trap K30ST4;
• 1 ♂; Kibale National Park, Kanyawara, Site R93, Malaise trap R93T2;
• 1 ♂; same data as preceding; Site K13, Malaise trap K13T1;
• 150 ♀; Kibale National Park, Kanyawara; Tapani Hopkins leg.;
158 ♀, 2 ♂; Ugandan specimens, data above and also in supplementary material (
1 ♀, holotype of
1 ♀, holotype of
1 ♂, paratype of
1 ♀; see
2 ♀; see
1 ♀; see
This species can be distinguished from other Afrotropical
Apart from the diagnosis, we provide the following additional or updated character traits to the description in
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
• 1 ♀, holotype; “Ilorin Prov.” [
This species can be distinguished from other Afrotropical
Nigeria.
The genus
The genus is cosmopolitan with the largest number of species found in the Oriental and Palaearctic regions.
• 1 ♀, holotype; “reg. Lac Kivu”, “Kadjudju” [Kajuju,
This species can be distinguished from other Afrotropical
Democratic Republic of Congo.
Sampling tropical ichneumonids for a year with numerous Malaise traps gave us an unprecedented sample size. We caught 456 individuals of the subfamily
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
The required research permits were issued by the Uganda National Council of Science and Technology and the Uganda Wildlife Authority.