http://dx.doi.org/10.5852/eit.2014.91

www. europeani ournaloftaxonomy. eu

2014 • Rousse R & Van Noort S.

This work is licensed under a Creative Commons Attribution 3.0 License.

Research article

urn:lsid:zoobank.org:pub:F6B6E184-448A-44EA-BFlB-D91122097847

A review of the Afrotropical Rhyssinae (Hymenoptera:

Ichneumonidae) with the descriptions of five new species

Pascal ROUSSE 1 - 2 - 4 -* & Simon VAN NOORT 1 - 3 - 5

1 Natural History Department, Iziko South African Museum, PO Box 61,

Cape Town 8000, South Africa

2 Stellenbosch University, Department of Botany and Zoology, Evolutionary Genomics Group,

Private Bag XI, Stellenbosch 7602, South Africa

3 Department of Biological Sciences, University of Cape Town, Private Bag,

Rondebosch 7701, South Africa

* Corresponding author: [email protected]

4 urn:lsid:zoobank.org:author:B06C2640-700A-429B-AA2F-lBE09251C845

5 urn: lsid: zoobank, org: author: 7 C CD 166F -F1FA-43D A-B5 82-4E 84E AF 5 9 AD 1

Abstract. The Afrotropical Rhyssinae are reviewed. A total of 12 species are reported from the region,

including five new species: Epirhyssa brianfisheri sp. nov., E. gavinbroadi sp. nov., E. shaka sp. nov., E.

villemantae sp. nov. and E. tombeaodiba sp. nov. The generic status of E. brianfisheri sp. nov. is discussed

since this species could also be considered to be an extra-limital Triancyra species, emphasizing the

putative paraphyletic status of Epirhyssa. Epirhyssaghesquierei Seyrig, 1937, E. overlaeti Seyrig, 1937

and E. uelensis Benoit, 1951 are newly reported from Cameroon. We provide illustrated diagnoses and

identification notes. Finally, we discuss the apparent scarcity of African rhyssines compared to other

regions.

Keywords. Parasitoid wasps, systematics, taxonomy, Africa, descriptions, identification keys

Rousse P. & Van Noort S. 2014. A review of the Afrotropical Rhyssinae (Hymenoptera: Ichneumonidae) with the

descriptions of five new species. European Journal of Taxonomy 91: \-A2. http://dx.doi.org/10.5852/eit.2014.91

Introduction

In many respects, Rhyssinae are atypical amongst Ichneumonidae. First, unlike most ichneumonids,

they are rather well known to the public because of their spectacular habitus. The subfamily includes

some of the largest species of ichneumonid wasps, exhibiting often striking color patterns. They also

have very long ovipositors used to access their wood-boring hosts. For example, species of Megarhyssa

Ashmead, 1900 often have an ovipositor that has a length of more than 20 cm, 3x longer than the

entire body, i.e., relatively and absolutely some of the longest known in insects (Townes 1975). They

are readily identifiable: the upper mandibular tooth is truncate and chisel-shaped, the mesoscutum is

flattened and crossed by deep transverse rugosities, and the last tergite of the female bears an apical

horn-like projection. Traditionally included as a tribe within Pimplinae, they were then considered as a

European Journal of Taxonomy 91: 1 —42 (2014)

separate monophyletic subfamily, mostly defined by the above autapomorphies, within the pimpliforme

subfamily group (Gauld 1991; Wahl & Gauld 1998).

Second, their diversification history appears to be an exception to the general rule that most taxa

in the tree of life arose in tropical areas and then diversified into the temperate areas (Jansson et al.

2013). The genus Rhyssa , considered to be the most basal lineage of Rhyssinae, is almost exclusively

Holarctic, whereas the more derived genera are primarily or exclusively tropical; the origin of other

pimpliform subfamilies (e.g., Diacritinae, Poemeniinae, Acaenitinae) was hypothesized to be in the

Northern Hemisphere for similar reasons (Wahl & Gauld 1998). However, their actual diversity in the

tropics, especially in the Neotropical and Afrotropical regions, appears to have been previously greatly

underestimated (Quicke 2012; Veijalainen et al. 2013).

The generic composition of Rhyssinae in tropical and temperate regions may be correctly assessed by

long-term sampling efforts. Rhyssinae are indeed exceptional for a third reason: due to their conspicuous

habitus, their moderate diversity, and their potential economic impact for biocontrol of wood-boring

pests, they received significantly more attention than most of the other ichneumonid subfamilies.

Available reviews cover the following regions: Oriental and Australasian (Baltazar 1961; Kamath &

Gupta 1972; Sheng & Sun 2010), Nearctic (Townes et al. 1960), Neotropical (Porter 1978; Gauld 1991;

Gauld et al. 1997), and Palaearctic (Horstmann 2002; Kasparyan & Khalaim 2007). They total about

250 species in eight genera (Yu et al. 2012). Nearly half of these species belong to the circumtropical

genus Epirhyssa Cresson, 1865. All host records are coleopteran or symphytan larvae (Yu et al. 2012).

They have also been reported to develop as facultative hyperparasitoids of other Symphyta parasitoids

(Hanson 1939), and are able to oviposit and develop on many surrogate hosts (Spradbery 1968). Indeed,

as idiobiont ectoparasitoids, the entire host selection process is only conditioned by what they can locate,

led mostly by chemical and mechanical stimuli (Spradbery 1970): any potential host they eventually

find is thus reduced to a mere “bag of meat” without any immune response to deal with, making the host

range potentially broader.

In accordance with the general rule for Ichneumonidae, their diversity in the Afrotropical region has been

poorly investigated. To date, only seven species have been reported from the region, of which six belong

to Epirhyssa (Yu et al. 2012). This is a relatively poor richness in comparison with the 48 currently

described Neotropical species (all Epirhyssa ), to which many others have still to be added (Saaksjarvi

et al. 2004), and the 140 Oriental and/or Australasian species (52 Epirhyssa). Rhyssinae are expected to

be most species-rich in lowland tropical rainforests (Gauld 1991), and hence these seven Afrotropical

species reflect the poor knowledge we have of the subfamily on this continent. This paper therefore aims

firstly to review the currently known species, by gathering, translating and updating their descriptions

(Seyrig 1937; Benoit 1951, 1952), secondly to add five new species from undetermined material housed

in African and European museums, and then to provide proper identification and description tools for

the expected numerous Rhyssinae species still to be collected in the Afrotropical Region. The scarcity

of available specimens is discussed.

Material and Methods

Photographs

Specimens were point mounted on black, acid-free card for examination (using a Teica M205C stereo

microscope with a TED light source), photography and long-term preservation. Images were taken using

the EntoVision® multiple-focus imaging system. This system combines a Leica® Ml6 microscope with

a JVC® KY-75U 3-CCD digital video camera attached that feeds image data to a notebook computer.

The program Cartograph®5.6.0 was then used to merge an image series (representing typically 10-

15 focal planes) into a single in-focus image. Lighting was achieved using techniques summarized in

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Buffington et al. (2005), Kerr et al. (2008) and Buffington & Gates (2009). All images presented in this

paper are available at http://www.waspweb.org

List of Depositories

BMNH

MNHN

MRAC

NMSA

SAMC

Natural History Museum, London, UK (Gavin Broad)

Museum National d’Histoire Naturelle, Paris, France (Claire Villemant)

Museum Royal de l’Afrique Centrale, Tervueren, Belgium (Eliane de Coninck)

KwaZulu-Natal Museum, Pietermaritzburg, South Africa (Burgert Muller)

Iziko South African Museum, Cape Town, South Africa (Simon van Noort)

Nomenclature and abbreviations

Morphological terminology follows Wahl & Sharkey (1993), but wing venation nomenclature follows

Gauld (1991). Most morphological terms are also defined on the HymAToL website ( http://www.

hymatol.org ) and HAO website ( http://portal.hvmao.Org/proiects/32/publ ic/ontologv/ l. If not indicated

otherwise, all geographical records are extracted from the Taxapad database (Yu et al. 2012). The

following morphometric abbreviations are used:

POL

OOL

body length, from toruli to metasomal apex ( mm )

antenna length, from base of scape to flagellar apex (mm)

fore wing length, from tegula to wing apex (mm)

clypeus transversality index (maximum width of clypeus / median height)

malar line index (malar line / basal mandibular width)

post-ocellar index (shortest distance between posterior ocelli / ocellus maximal diameter)

oculo-ocellar index (shortest distance between eye and posterior ocellus / ocellus maximum

diameter)

length index of flagellomere n (length / width of flagellomere n)

tergite 1 elongation index, (median length of tergite 1 / apical width (applies to female only

if not otherwise specified))

ovipositor sheath-tibia index (length of ovipositor sheath / length of hind tibia)

The first three measurements (absolute measures) were measured on all specimens in the type series,

with data for the primary type reported separately in brackets if necessary. The relative indices were

measured on the primary type specimen only.

Results

Five new species were found in the undetermined material from BMNH, NMSA and SAMC. The key to

and the descriptions of all species are provided below. Additional online dichotomous and matrix keys

are available at www.waspweb.org .

European Journal of Taxonomy 91 : 1 —42 ( 2014 )

Key to species

Areolet closed

2.0 mm

1. Fore wing with areolet closed, triangular and strongly petiolate (A); body orange and dark brown,

wings dark brown (B); tropical Africa. Megarhyssa babaulti Seyrig, 1937

Areolet open

- Fore wing with areolet open (a); body and wings colorations variable but usually wings hyaline

and body yellow interspersed with black markings (b).2 (. Epirhyssa )

2. Face smooth to sparsely punctate (A-B), without transverse rugosities

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

- Face transversely rugulose punctate or transversely striate (a-b)

3. Wings hyaline, fore wing slightly infuscate apically (A); Madagascar. E. migratoria Seyrig, 1932

- Wings with dark patterns, yellowish to black (a, b); tropical Africa

European Journal of Taxonomy 91: 1 —42 (2014)

4. Wings blackish with subapical lighter patch (A); epicnemial carina reaching level of mesopleural

pit (B). E. villemantae sp. nov.

- Wings yellowish with apex blackish and an additional subapical transverse blackish band on fore wing

(a); epicnemial carina very short, hardly extending onto mesopleuron (b). E. maynei Benoit, 1952

tubercle absent

Median subapical

. -

---

, ;

Metasoma with large apico-lateral

yellow spots on tergites 1-5

5. Metasoma with large apico-lateral yellow spots on tergites 1-5 (A); clypeus without median subapical

tubercle (B); body and ovipositor large to very large (B > 25mm; OT >6). E. overlaeti Seyrig, 1937

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

- Metasoma usually uniformly colored (a); clypeus with a distinct subapical tubercle (b); body and

ovipositor smaller (B < 25mm; OT < 6) .6

Anterior sclerotized part

of sternite 1 with a pair of

submedian tubercles

Tubercles also present

on sternites 2-4

6. Anterior sclerotized part of sternite 1 with a pair of submedian tubercles, tubercles also present on

sternites 2-A (A); general coloration dark testaceous to black interspersed with pale yellow (B).

. E. brianfisheri sp. nov.

- Sternite 1 without a pair of submedian tubercles, tubercles present on sternites 2-A only (a); general

coloration pale to striking yellow with isolated black markings (b).7

European Journal of Taxonomy 91: 1 —42 (2014)

- Apex of subtegular ridge convex, without lateral flange (a) .9

0.25

Clypeus punctate

1 »

Tergite 1 slender >1.4x longer

than apically wide

8. Clypeus punctate (A); tergite 1 of female slender, more than 1,4x longer than apically wide (B).

. E. gavinbroadi sp. nov.

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Tergite 1 stout <1 Ax longer than wide

0.25 mm

Clypeus almost smooth ^

- Clypeus almost smooth, hardly sculptured (a); tergite 1 of female stout, less than 1.4x longer than

wide (b) . E. uelensis Benoit, 1951

9. Tergite 2 densely punctate over its entire surface and tergite 1 more than 1.5x longer than apically

wide (A); general coloration pale yellow (B); Southern Africa. E. shaka sp. nov.

Tergite 2 smooth to superficially

punctate laterally

0.25 mm

- Tergite 2 smooth to superficially punctate laterally (a), if ambiguous (some E. tombeaodiba) then

tergite 1 stouter, less than 1.5x longer than apically wide; general coloration bright yellow (b);

tropical Africa.10

European Journal of Taxonomy 91: 1 —42 (2014)

10. Fore wing with 2m-cu basal to rs-m (A); mesosternum black (B) . E. leroyi Benoit, 1951

Fore wing with 2m-cu apical to rs-m (a); mesosternum yellow (b) .11

lypeus shallowly

longitudinally strigose

0.25 mm

Tergite 1 broad

-■.

inpp

Carinae

converge

in front of

median

ocellus

1.0 mm

11. Clypeus shallowly longitudinally strigose (A); frons with two medial carinae converging in front of

median ocellus (B); tergite 1 of female stout, less than 1.4x longer than apically wide (C).

. E. tombeaodiba sp. nov.

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Clypeus smooth

—

0.25

Medial carinae diverge

toward lateral ocelli

Tergite 1 slender

1.0 mm

Clypeus smooth (a); frons with two submedian carinae, diverging on upper frons toward

lateral ocelli (b); tergite 1 of female slender, more than 1.4x longer than apically wide (c)

. E. ghesquierei Benoit, 1951

Taxonomic descriptions

Class Hexapoda Blainville, 1816

Order Hymenoptera Linnaeus, 1758

Suborder Apocrita Latreille, 1810

Superfamily Ichneumonoidea Latreille, 1802

Lamily Ichneumonidae Latreille, 1802

Subfamily Rhyssinae Morley, 1913

Genus Epirhyssa Cresson, 1865

Hierax Tosquinet, 1903: 255.

Rhyssonota Kriechbaumer, 1890: 489.

Sychnostigma Baltazar, 1961: 75.

Diagnosis (updated from Townes 1969 and Gauld 1991)

Small to large insects (fore wing length 6-25 mm), usually orange or yellow interspersed with black

maculae; clypeus very small, not subdivided, often with a subapical ridge bearing median and/or lateral

weak to strong tubercles; ventral margin of clypeus thin, slightly concave, often with lateral corners

produced; mandible not twisted, stout, strongly constricted basally, then weakly tapered, apically

bidentate with lower tooth pointed (may appear chisel-like when worn) and upper tooth blunt or chisel¬

like; occipital carina complete to nearly absent, when ventrally present joining hypostomal carina above

mandibular base; hypostomal carina strongly raised above mandible base; subocular sulcus absent,

but malar area granulate; pronotum long, mid-dorsally with a deep invagination and with thin margin

projecting beyond invagination; antero-lateral margin of pronotum broadly rounded in front, its lower

corner acute and slightly in-turned; epomia weak to moderately strong, contiguous with and diverging

from anterior pronotal margin; mesoscutum with strong transverse rugosities; notauli anteriorly strong

and meeting in centre of mesoscutum, defining an anteriorly abruptly rounded median lobe; mesopleuron

moderately long, epicnemial carina most often present laterally and reaching to or above lower corner

of pronotum, its dorsal end evanescent; mesopleural suture weakly angled centrally; submetapleural

carina weak to strong, more or less complete; propodeum of moderate length, without carina dorsally

but pleural carina distinct; tarsal claws simple, large; fore wing with areolet open, 2m-cu slightly basal

to distinctly apical to rs-m; hind wing with distal abscissa of Cu present, joining Cu&cu-a near or at

junction with M; mid trochantellus with a ventral longitudinal ridge; metasoma moderately slender,

tergites smooth to finely sculptured; tergite 1 fused with its sternite and without glymma; apical margins

of tergites 3-5 hardly to strongly concave; female with sternites 2-A (but see E. brianfisheri sp. nov.)

with an anterior pair of tubercles, and with an apical truncate horn-like process on last tergite, ovipositor

European Journal of Taxonomy 91: 1 —42 (2014)

stout, laterally compressed, 4-5x longer than hind tibia; male with metasoma slenderer and subgenital

plate elongate, posteriorly rather concave.

Species richness and distribution (Yu et al. 2012)

Epirhyssa is mostly circumtropical with some Far-Eastern species occurring up to Northern Japan; it

includes 113 valid species with only six reported from the Afrotropical Region, to which we add here

five new species.

Epirhyssa brianfisheri sp. nov.

um:lsid:zoobank.org:act:lC872313-52C3-4D81-B8AD-94AA2C724EAC

Fig. 1

Diagnosis

Overall color mottled, pale yellow and dark brown to black; wings hyaline, apically infuscate; face

rugulose-punctate, strongly protruding medially; clypeus sparsely punctate, with median subapical

tubercle strong and lateral subapical tubercle weak, ventral margin strongly produced laterally; Irons

rugulose with a blunt Y-shaped mid-longitudinal carina; antenna with 33 flagellomeres; mesopleuron

with shallow transverse striations; epicnemial carina hardly reaching level of mesopleural pit; subtegular

ridge convex; fore wing with 2m-cu opposite rs-m; tergite 1 slender; sternite 1 with an additional pair

of basal tubercles; tergites 1-2 smooth, following tergites densely and finely punctate; apical margins of

tergites 3-5 moderately concave. CT 3.3; ML 0.7; POL 2.0; OOL 1.5; FIj 4.2; Fl 15 1.8; F1 J0 1.2; T1 1.7;

OT 3.9.

Differential diagnosis

Species characterized by the exceptional mottled coloration and the presence of antero-basal tubercles

on sternite 1 {cf. taxonomic remarks). All other species of Epirhyssa lack the antero-basal tubercles.

Etymology

Dedicated to Brian Fisher, friend and expert organizer of expeditions into central Africa, including the

CAR expedition that discovered this species.

Type material

Holotype

CENTRAL AFRICAN REPUBLIC: $, Prefecture Sangha-Mbaere, Reserve Speciale de Foret Dense de

Dzanga-Sangha, 12.7 km, 326° NW of Bayanga, 3 o 00.27’N 16°11.55’E, 420 m, 13 May 2001, S. van

Noort, Sweep CAR01-S158, Lowland Rainforest, SAM-HYM-P048018 (SAMC).

Description

Female (holotype)

B 18.6: A 11.0: F 12.8.

Color. Head pale yellow with black to dark brown parts: mandible, malar space, median tubercle of

clypeus, a thin mid-longitudinal line on upper face, frons, vertex, and most of occiput; mesosoma pale

yellow with most of mesoscutum black, and testaceous to brown parts: anterior margin of pronotum,

propleuron, most of mesopleuron but subtegular ridge and two lateral patches, mesosternum, ventral

margin of metapleuron and anterior third of propodeum; metasoma dark brown with pale yellow

markings: tergites 1-2 medially, tergite 3-4 posteriorly; sternites pale yellow, more or less extensively

testaceous anteriorly; antenna dark brown with scape somewhat pale maculated; ovipositor sheath

brown; fore and mid legs testaceous with coxae and tibiae mostly pale yellow, hind leg testaceous with

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

coxa partly pale yellow and femur with an outer longitudinal pale yellow stripe; wings hyaline, fore

wing anteriorly infuscate from pterostigma to apex.

I Central African Republic,

Prefecture Sangha-Mbaere.

Reserve Speciale de Foret

Dense de Dzanga-Sangha,

112 7km .'26° N~W Bayanga

pO'N 16°11 55'E, 420i

13.V.2001, S vanNoort,

Sweep. CAR01-S158,

U.owland Rainforest

del. S. Van Nonrt 2013

SAM-HYM

P048018

IMAGED

WaspWeb

SAMC 2013

2.0 mm

I-1

1.0 mm

I-1

-1

Fig. 1. Epirhyssa brianfisheri sp. nov., holotype, §. A. Habitus, lateral view (inset: data labels). B.

Mesosoma, lateral view. C.. Metasoma, dorsal view. D. Head, anterior view. E. Mandibles, anterior

view.

European Journal of Taxonomy 91: 1 —42 (2014)

Head. Densely setose with long hairs, hairs shorter on upper back head; face rugulose punctate, strongly

protruding medially; clypeus sparsely punctate, subapical ridge with median tubercle strong and lateral

tubercle weak, ventral margin strongly produced laterally; mandible stout; frons transversely rugose with

a blunt Y-shaped mid-longitudinal carina below median ocellus, inner margins of toruli not expanded

backwards; vertex and temple sparsely punctate but inter-ocellar area rugose; ocellar triangle wide;

occipital carina mid-dorsally interrupted; antenna with 33 flagellomeres.

Mesosoma. Entirely covered with dense and long hairs; mesosoma moderately densely punctate, but

mesonotum deeply transversely striate, anterior half of pronotum smooth and mesopleuron somewhat

transversely strigose medially; epomia moderate; epicnemial carina hardly reaching level of mesopleural

pit; apex of subtegular ridge convex, not flanged laterally; submetapleural carina strong, slightly and

evenly narrowed toward apex.

Wings. Fore wing with 2m-cu opposite rs-m, cu-a almost opposite Rs&M, and Rs strongly bowed

forwards; hind wing with distal abscissa of Cu joining Cu&cu-a close to junction with M, with a basal

group of two hamuli on distal abscissa of Rl.

Metasoma. Tergite 1 moderately slender; stemite 1 with a basal pair of submedian tubercles, additional

to usual ones on sternites 2—4; tergites 1-2 mostly smooth, following tergites densely and finely punctate

with fine transverse striations medially; apical margins of tergites 3-5 moderately concave.

Male

Unknown.

Distribution

Central African Republic.

Taxonomic remarks

The presence of tubercles on sternite 1 is quite unexpected in Epirhyssa , while it is one of the features

defining the Oriental genus Triancyra Baltazar, 1961. However, we refrain from placing this new species

in Triancyra for the following reasons. Besides the geographical separation from the known distribution

area of Triancyra , the majority of morphological characters exhibited by this new species do not fit

the current generic definition for Triancyra. For example, E. brianfisheri sp. nov. exhibits subequal

mandibular teeth (vs. upper tooth narrower in Triancyra), a pterostigma that is long and slender (vs. up

to 5x longer than high), hind wing with a basal group of two hamuli (vs. usually three), tergite 1 long and

slender (vs. stout, 1.3x longer than wide) (Townes, 1969). Hence, we are of the opinion that the species

fits more comfortably within Epirhyssa than in Triancyra, but the possession of additional tubercles

in Epirhyssa species underlines the need for a global phylogenetic revision of the whole subfamily.

Epirhyssa is indeed poorly defined and probably not monophyletic (G. Broad, pers. comm.).

Epirhyssa gavinbroadi sp. nov.

urn:lsid:zoobank.org:act:94C30C69-E07C-4C48-BB79-3B3D903665EE

Fig. 2

Diagnosis

Bright yellow overall with isolated dorsal black markings; wings hyaline, slightly infuscate apically; face

rugulose-punctate, moderately protruding dorsally; clypeus punctate, with median subapical tubercle

moderate, ventral margin strongly produced laterally; frons smooth with a sharp mid-longitudinal carina

reaching median ocellus and lateral curved carina toward lateral ocellus; antenna with more than 35

flagellomeres; mesopleuron with epicnemial carina reaching level of mesopleural pit; subtegular ridge

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Fig. 2. Epirhyssa gavinbroadi sp. nov., holotype, $. A. Habitus, lateral view (inset: data labels). B.

Mesosoma, lateral view. C. Metasoma, dorsal view. D. Head, anterior aview. E. Mandibles, anterior

view.

European Journal of Taxonomy 91: 1 —42 (2014)

concave, laterally flanged; fore wing with 2m-cu slightly apical to rs-m; tergite 1 slender; tergites 1-5

smooth, following tergites densely and finely punctate; apical margins of tergites 3-7 strongly concave.

CT 2.7; ML 0.5; POL 1.3; OOL1.6; 4.2; Fl 15 1.8; T1 1.7; OT 5.2.

Differential diagnosis

A mostly bright yellow species with hyaline wings, otherwise characterized by the combination of the

punctate clypeus, the concave and flanged subtegular ridge, and the sharp mid-longitudinal carina on the

frons, which separates this species from other yellow species with hyaline wings (these species have a

strigose or hardly sculptured clypeus, and all of them except E. uelensis have no flange on the subtegular

ridge).

Etymology

Dedicated to Gavin Broad for his constant support and his most valuable advice.

Type material

Holotype

CAMEROON: $, Nkoemvon, 02°48’N 11°08’E, 16 Mar.M May 1980. Ms D. Jackson (BMNH).

Description

Female (holotype)

B 15.7; ANA; F 12.1.

Color. Bright yellow overall with black markings: mandible, mid-apical tubercle on clypeus, vertex,

temple, occiput mid-dorsally, mesoscutum but postero-lateral margins and a central macula, and a small

mid-posterior macula on tergites 3-6; flagellum and ovipositor sheath dark brown; wings hyaline,

slightly infuscate apically, venation brown.

Head. Face rugulose punctate, moderately protruding dorsally; clypeus punctate, with a moderate

subapical median tubercle, ventral margin strongly produced laterally; mandible stout; frons smooth;

inner margins of toruli expanded backwards into two converging lamellar carinae, carinae then fused

into a sharp mid-longitudinal carina reaching median ocellus; frons also with an additional lateral curved

carina from posterior margin of torulus to next to lateral ocellus; inter-ocellar area rugose, remainder of

vertex and temple finely punctate; occipital carina mid-dorsally evanescent; antenna with more than 35

flagellomeres (apices broken).

Mesosoma. Mesosoma moderately densely punctate, but mesonotum deeply transversely striate and

anterior half of pronotum smooth; epomia very weak; epicnemial carina reaching level of mesopleural

pit; apex of subtegular ridge concave, strongly flanged laterally; submetapleural carina slightly and

evenly narrowed toward apex.

Wings. Fore wing with 2m-cu slightly apical to rs-m, cu-a slightly apical to Rs&M, and Rs strongly

bowed forwards; hind wing with distal abscissa of Cu joining Cu&cu-a close to junction with M.

Metasoma. Tergite 1 moderately slender; tergites 1-5 mostly smooth, apex of tergite 5 and following

tergites finely and densely punctate; apical margins of tergites 3-7 strongly concave.

Male

Unknown.

Distribution

Cameroon.

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Epirliyssa gliesquierei Seyrig, 1937

Fig. 3

Diagnosis

Bright yellow overall with isolated dorsal black markings; wings hyaline, slightly infiiscate apically;

face deeply rugulose-punctate, moderately protruding dorsally; clypeus nearly smooth, with median

subapical tubercle strong, ventral margin produced laterally; frons laterally punctate and medially

rugulose, with two weak submedian carinae diverging below median ocellus toward lateral ocelli;

antenna with 34 flagellomeres; mesopleuron with epicnemial carina reaching above level of mesopleural

pit; subtegular ridge convex; fore wing with 2m-cu slightly apical to rs-m; tergite 1 slender; tergites 1-2

smooth, following tergites shallowly punctate with basal transverse striations; apical margins of tergites

straight. CT 2.7; ML 0.7; POL 1.1; OOL 1.3; Fl t 4.1; Fl 15 2.2; Fl 33 1.1; T1 2.5; OT 4.2.

Differential diagnosis

A mostly bright yellow species with hyaline wings, otherwise characterized by the combination of the

pair of submedian carinae on the frons diverging towards the lateral ocelli and the slender tergite 1. The

other yellow species with hyaline wings have a stouter tergite 1 in the female, less than twice as long as

apically wide.

Material examined

Holotype

DEMOCRATIC REPUBLIC OF CONGO: ?, Eala [00°40’N 18 0 17’E], Nov. 1935, J. Ghesquiere, R.

Det. G 3330 (MRAC).

Paratype

DEMOCRATIC REPUBLIC OF CONGO: 1 S, Bambesa, Dec. 1946, PL. Benoit (MRAC).

Other material

CAMEROON: 1 S, Korup, Dec. 1980-Jan. 1981, Mrs D. Jackson (BMNH); 1 S, Korup, 1981, Mrs D.

Jackson (BMNH).

Description

Female (holotype)

B 14.6; A 12.2; F 12.1.

Color. Bright yellow overall, with black markings on mandible, vertex, upper occiput, and mesoscutum

margins; antenna and ovipositor sheath dark brown; wings hyaline, apex hardly infiiscate, venation pale

brown.

Head. Face subquadrate, deeply rugulose punctate and moderately protruding dorsally; clypeus almost

smooth, with a strong subapical median tubercle, ventral margin produced laterally; mandible stout;

frons laterally moderately punctate, medially smoothly rugulose; inner margins of toruli expanded

backwards into two weak sinuate submedian carinae, carinae then diverging and reaching lateral ocelli;

vertex and temple slightly and finely punctate; occipital carina complete but mid-dorsally faint; antenna

with 34 flagellomeres.

European Journal of Taxonomy 91: 1 —42 (2014)

Mesosoma. Mesosoma shallowly and moderately densely punctate, but mesonotum deeply transversely

striate and anterior half of pronotum smooth; epomia very weak; epicnemial carina long, its evanescent

dorsal end reaching above level of mesopleural pit; apex of subtegular ridge convex, not flanged laterally;

submetapleural carina thick, evenly narrowed toward apex.

lS4v itt-ei

'Type

IMAGED

WaspWeD

SAMC 2013

Eala IP^ TYP , E

,.V/ - 1935 IK | UAM

J. GhesquiAre

2.0 mm

I-1

•v-i-.

0.49 mm

I-1

>//

Fig. 3. Epirhyssa ghesquierei Seyrig, 1937, holotype, $. A. Habitus, lateral view (inset: data labels). B.

Mesosoma, lateral view. C. Metasoma, dorsal view. D. Head, anterior view. E. Mandibles, anterior view.

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Wings. Fore wing with 2m-cu slightly apical to rs-m, cu-a slightly apical to Rs&M, and Rs wealdy

bowed forwards; hind wing with distal abscissa of Cu joining Cu&cu-a close to junction with M.

Metasoma. Tergite 1 slender and smooth, tergite 2 mostly smooth with some superficial punctures

laterally, tergites 3-6 shallowly punctate with basal transverse striations; apical margins of all tergites

straight.

Male (3 specimens)

B 11.8-13.6; A 15.1-16.2; F 14.9-16.0.

Similar to female but tergite 1 slenderer (T1 2.5); the two males from Cameroon are significantly lighter,

without dorsal black markings.

Distribution

Democratic Republic of Congo. New record: Cameroon.

Epirhyssa leroyi Benoit, 1951

Fig. 4

Diagnosis

Bright yellow overall with black markings including entire mesosternum, metasoma fading to brown

apically; wings hyaline, slightly infuscate apically; face rugulose-punctate, strongly protruding medially;

clypeus nearly smooth, with median and lateral subapical tubercles moderate, ventral margin produced

laterally; frons laterally punctate and medially striate, with a blunt Y-shaped mid-longitudinal carina;

mesopleuron with epicnemial carina reaching above level of mesopleural pit; subtegular ridge convex;

fore wing with 2m-cu slightly basal to rs-m; tergite 1 of female stout (male unknown); tergites 1-3

smooth, following tergites shallowly punctate with basal transverse striations; apical margins of tergites

3^1 slightly concave. CT 3.6; MF 0.6; POF 1.6; OOF 1.4; F[ { 3.4; F1 1S NA; T1 1.2-1.4; OF 4.4.

Differential diagnosis

A mostly bright yellow species with hyaline wings, otherwise characterized by the combination of the

black mesosternum, vein 2m-cu slightly basal to rs-m and the stout tergite 1 (at least in the female). All

other yellow Afrotropical Epirhyssa species have the mesosternum concolorous with the remainder of

the mesosoma and the vein 2m-cu slightly apical to rs-m.

Material examined

Holotype

DEMOCRATIC REPUBFIC OF CONGO: ?, Bambesa [03°28’ N 25 0 43’E], Nov. 1933, J.V. Feroy

(MRAC).

Paratype

DEMOCRATIC REPUBFIC OF CONGO: ?, Ubangui-Bumba, Dec. 1939, H. de Saeger (MRAC).

Description

Female (holotype)

B 14.9; ANA; F 11.1.

Color. Bright to dull yellow overall, with metasoma fading to brown toward apex, and with black

markings: mandible, ventral margin of clypeus, ocellar area (macula connected to eye margin), occiput

European Journal of Taxonomy 91: 1 —42 (2014)

IMAGED

WaspV/e&

SAMC 3013

MUSEE DU CONGO

Bambesa

Xu -1933

J. V. Lerov

0.50 mm

I-1

0.5

* ».

0.25 mm

Fig. 4. Epirhyssa leroyi Benoit, 1951, holotype, $. A. Habitus, lateral view (inset: data labels). B.

Mesosoma, lateral view. C. Metasoma, dorsal view. D. Head, anterior view. E. Mandibles, anterior view.

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

mid-dorsally, mesosternum, and a median macula on each mesoscutal lobe; antenna and ovipositor

sheath dark brown; wings hyaline, slightly infuscate apically, venation pale brown.

Head. Face rugulose-punctate, strongly bulging medially; clypeus smoothly sculptured with distinct

lateral and median subapical tubercles, ventral margin strongly produced laterally; mandible short

and stout; frons laterally sparsely punctate, medially transversely striate with a blunt Y-shaped mid¬

longitudinal carina below median ocellus, inner margins of toruli not expanded backwards; temple and

vertex finely punctate, inter-ocellar area somewhat rugose; occipital carina mid-dorsally interrupted;

antenna broken in both specimens.

Mesosoma. Mesosoma shallowly and moderately densely punctate, but mesonotum deeply transversely

striate and anterior half of pronotum smooth; epomia moderate; epicnemial carina long, its evanescent

dorsal end reaching above level of mesopleural pit; apex of subtegular ridge convex, not flanged laterally;

submetapleural carina anteriorly expanded, then distinctly narrowed toward apex.

Wings. Fore wing with 2m-cu slightly basal to rs-m, cu-a slightly apical to Rs&M, and Rs wealdy

bowed forwards; hind wing with distal abscissa of Cu joining Cu&cu-a close to junction with M.

Metasoma. Tergite 1 stout and smooth, tergites 2-3 nearly smooth, following tergites finely and distinctly

punctate with weak basal striations; apical margins of tergites 3-4 slightly concave, others straight.

Male

Unknown.

Distribution

Democratic Republic of Congo.

Epirhyssa maynei Benoit, 1952

Fig. 5

Diagnosis

Bright yellowish-orange overall with dark dorsal markings including apical half of metasoma; wings

yellowish with apices strongly infuscate and an additional infuscate transverse subapical stripe on fore

wing; face sparsely punctate, barely protruding; clypeus nearly smooth, without subapical tubercle,

ventral margin truncate; frons smooth with a blunt Y-shaped mid-longitudinal carina; antenna with 37

flagellomeres; epicnemial carina hardly extending onto mesopleuron; subtegular ridge convex; fore

wing with 2m-cu opposite rs-m; tergite 1 stout; all tergites smooth, with fine isolated punctures; apical

margins of tergites 3-5 strongly concave. CT 2.9; ML 0.5; POF 1.3; OOF 1.3; Fl x 3.7; Fl 15 2.5; Fl 36 1.2;

T1 1.2.

Differential diagnosis

Species characterized by its coloration, with apical half of metasoma black, wings yellowish with black

patterns; otherwise characterized by the combination of the sparsely punctate face, the absence of

subapical protuberances on the clypeus, the ventral margin of the clypeus not produced laterally, and the

epicnemial carina nearly absent laterally. It is the only yellow Afrotropical species with banded wings.

Material examined

Holotype

DEMOCRATIC REPUBFIC OF CONGO: S, Albertville [Kalemie, 05°56’N 29°12 , E], Jul. 1918, R.

Mayne (MRAC).

European Journal of Taxonomy 91: 1 —42 (2014)

2.0

D.25

Ini'

0.5 mm

I-1

WiU-t-jj y 1 1951

(let. P.L.Gr. Benoil

HOLOTTPUS

■ . './■ *- ~ 1

det, Dr.Fahrlnger

*, j IMAGED

i. f j- » WaspWeb

"/* |H SAMC2013

rtneer m

MUSEEDU CONGO

Albertville

XII-1918

R. Mayne

Fig. 5. Epirhyssa maynei Benoit, 1952, holotype, $. A. Habitus, lateral view (inset: data labels). B.

Mesosoma, lateral view. C. Metasoma, dorsal view. D. Head, anterior view. E. Mandibles, anterior view.

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Paratype

DEMOCRATIC REPUBLIC OF CONGO: 1 Bambesa, Dec. 1946, PL. Benoit (MRAC).

Other material

CENTRAL AFRICAN REPUBLIC: 1 Prefecture Sangha-Mbaere, Parc National de Dzanga-Ndoki,

38.6 km, 173°S of Lidjombo, 2°21.60 , N 16°09.2(TE, 350 m, 27 May 2001, S. van Noort, handnet.

Lowland Rainforest, CAR01-H25, SAM-HYM-P049437 (SAMC).

Description

Female

Unknown.

Male (3 specimens)

B 14.2-16.7; A 13.6-14.9; F 11.5-13.3 (holotype: B 16.7; A 14.9; F 13.3).

Color. Bright yellowish orange overall with dark brown to black markings: apical 2/3 of mandible,

frons, vertex, temple, hind leg from femur and metasoma from tergite 4; flagellum dark brown; wings

yellowish, apically strongly infuscate, fore wing with an additional transverse infuscate stripe at

pterostigma level, venation yellow.

Head. Face sub-quadrate, sparsely punctate, barely protruding dorsally but with a short mid-longitudinal

ridge near toruli; clypeus mostly smooth with some punctures along dorsal margin, without distinct

subapical tubercle, ventral margin truncate; mandible stout, both teeth truncate and chisel-like; upper

head quite smooth; frons with a blunt Y-shaped mid-longitudinal carina below median ocellus, inner

margins of toruli not expanded backwards; occipital carina mid-dorsally evanescent; antenna with 34-

37 flagellomeres.

Mesosoma. Mesosoma shallowly and moderately densely punctate, but mesonotum deeply transversely

striate and anterior half of pronotum smooth; epomia very weak; epicnemial carina very short,

hardly extending laterally onto mesopleuron; apex of subtegular ridge convex, not flanged laterally;

submetapleural carina anteriorly moderately expanded, then abruptly narrowed from anterior third.

Wings. Fore wing with 2m-cu opposite rs-m, cu-a distinctly apical to Rs&M, Rs moderately bowed

forwards; hind wing with distal abscissa of Cu joining Cu&cu-a at its anterior third, remote from

junction with M.

Metasoma. Tergite 1 slender; all tergites smooth but with fine isolated punctures; apical margins of

tergites 3-5 strongly concave.

Distribution

Democratic Republic of Congo. New record: Central African Republic.

Diagnosis

Epirhyssa migratoria Seyrig, 1932

Fig. 6

Bright yellow overall with isolated dorsal black markings; wings hyaline, slightly infuscate apically;

face sparsely punctate, almost flat; clypeus nearly smooth, with median and lateral subapical tubercles

moderate, ventral margin produced laterally; frons smooth, with two submedian carinae diverging below

median ocellus toward lateral ocelli; antenna with 36 flagellomeres; epicnemial carina laterally absent;

subtegular ridge convex; fore wing with 2m-cu slightly apical to rs-m; tergite 1 weakly slender in

European Journal of Taxonomy 91: 1 —42 (2014)

female, much more slender in male; tergites 1-3 smooth, following tergites sparsely punctate with basal

transverse striations; apical margins of tergites 3-5 moderately concave. CT 2.5; ML 0.6; POL 2.6; OOL

1.3; Flj 4.4; Fl 15 2.3; Fl 35 1.1; T1 1.4-1.5; OT 5.0.

Fig. 6. Epirhyssa migratoria Seyrig, 1932, holotype, $. A. Habitus, lateral view (inset: data labels). B.

Mesosoma, lateral view. C. Metasoma, dorsal view. D. Head, anterior view. E. Mandibles, anterior view.

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Differential diagnosis

A mostly bright yellow species with hyaline wings, otherwise characterized by the combination of

the flat and sparsely punctate face, the pair of submedian carinae on the frons and the absence of the

epicnemial carina on the mesopleuron. The lateral part of the epicnemial carina is distinct in every other

yellow species with hyaline wings.

Material examined

Lectotype

MADAGASCAR: $, Rogez [48°18’S 48°32 , E], Sep. 30, A. Seyrig, MNHN EY8815.

Paratypes

3 SS, same label data.

Description

Female (holotype)

B 18.2; A 15.1; L 14.7.

Color. Bright yellow overall with infuscate to black markings: mandible, inter-ocellar area (macula

connected to eye margin), dorsal margin of occiput, mesoscutal lobes mid-longitudinally, and scuto-

scutellar groove; antenna and ovipositor sheath mostly dark brown; wings nearly hyaline, apically

slightly infuscate, venation testaceous.

Head. Face sparsely punctate, almost flat; clypeus smoothly sculptured with moderate lateral and median

subapical tubercles, ventral margin moderately produced laterally; mandible stout; upper head quite

smooth to moderately and shallowly punctate, inter-ocellar area somewhat rugose; frons with a pair of

submedian carinae, carinae abruptly diverging below median ocellus and following anterior margin of

vertex, inner margins of toruli hardly produced backwards; occipital carina mid-dorsally interrupted;

antenna with 36 flagellomeres.

Mesosoma. Mesosoma shallowly and moderately densely punctate, but mesonotum transversely striate

and anterior half of pronotum smooth; epomia weak; epicnemial carina absent except for a short ventral

remnant; apex of subtegular ridge convex, not flanged laterally; submetapleural carina strong and even

all along.

Wings. Fore wing with 2m-cu slightly apical to rs-m, cu-a slightly apical to Rs&M, and Rs moderately

bowed forwards; hind wing with distal abscissa of Cu joining Cu&cu-a close to junction with M.

Metasoma. Tergite 1 moderately slender; tergites 1-3 smooth, following tergites finely and sparsely

punctate with basal transverse striations; apical margins of tergites 3-5 moderately concave, others

straight.

Male (3 specimens)

B 17.5-18.2; A14.3-14.9; L 14.0-14.5.

Similar to female but metasoma much more slender (T1 2.0-2.2).

Distribution

Madagascar.

European Journal of Taxonomy 91: 1 —42 (2014)

Epirhyssa overlaeti Seyrig, 1937

Fig. 7

Diagnosis

Overall color mottled, black, yellow and testaceous; wings hyaline, slightly infuscate apically; face

transversely striate, nearly flat; clypeus shallowly and sparsely punctate, without subapical tubercle,

ventral margin strongly produced laterally; frons smooth, with a sharp mid-longitudinal carina reaching

median ocellus and lateral curved carina toward lateral ocellus; antenna with 41M3 flagellomeres;

epicnemial carina reaching lower level of speculum; fore wing with 2m-cu opposite rs-m; tergite 1

slender; all tergites nearly to quite smooth, tergites 4-5 finely transversely striate basally; apical margins

of tergites 3-6 strongly concave. CT 2.5; ML 0.5; POL 1.3; OOL 1.6; F1 T 3.9; Fl 15 2.8; Fl 40 1.3; T1 1.7;

OT 7.2.

Differential diagnosis

Highly distinctive species of mottled color and large to very large size; otherwise characterized by the

combination of the flat and sparsely punctate face, the sharp mid-longitudinal carina on the frons, the

absence of a subapical ridge on the clypeus, and the very long ovipositor. No other known Epirhyssa

species shares this color pattern.

Material examined

Holotype

DEMOCRATIC REPUBLIC OF CONGO: $, Lulua [10°37’S 24°54’E], Kapanga, Apr. 1933, F.G.

Overlaet, R. Det. F 3330 (MRAC).

Other material

CAMEROON: 1 $, Nkoemvon, Jul.-Aug. 1980, Ms. D. Jackson (BMNH).

Description

Female (2 specimens)

B 25.2-37.8; A 15.4-23.2; F 16.1-24.2.

Color. Head yellow with mandible black and lower gena, vertex and upper occiput brownish testaceous;

mesosoma black to dark brownish testaceous interspersed with yellow markings: pronotum dorsally

and ventrally, mesopleuron anteriorly and posteriorly, and mesoscutum, scutellum, metanotum and

propodeum centrally; legs testaceous to dark testaceous with fore and mid coxa yellow; metasoma light

to dark testaceous, tergites 1-5 with apico-lateral large yellow spots; antenna and ovipositor sheath dark

brown; wings nearly hyaline, apically infuscate, venation dark brown.

Head. Face subquadrate, transversely striate and hardly bulging medially; clypeus shallowly and sparsely

punctate, without subapical tubercle, ventral margin strongly produced laterally; mandible stout; upper

head quite smooth; inner margins of toruli expanded backwards into two converging lamellar carinae,

carinae then fused into a sharp mid-longitudinal carina reaching median ocellus; frons also with an

additional lateral curved carina from posterior margin of torulus to next to lateral ocellus; occipital

carina mid-dorsally interrupted; antenna with 41-43 flagellomeres.

Mesosoma. Mesosoma shallowly and moderately densely punctate, but mesonotum transversely

striate and anterior half of pronotum smooth; epomia indistinct; epicnemial carina reaching lower

level of speculum, mesopleural pit indistinct; apex of subtegular ridge convex, not flanged laterally;

submetapleural carina moderate and even all along.

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Wings. Fore wing with 2m-cu opposite rs-m, cu-a slightly apical to Rs&M, and Rs short and moderately

bowed forwards; hind wing with distal abscissa of Cu joining Cu&cu-a at junction with M.

Metasoma. Tergite 1 moderately slender; all tergites smooth to indistinctly sculptured but tergites 4-5

wealdy striate basally; apical margins of tergites 3-6 strongly concave, others straight.

Fig. 7. Epirhyssa overlaeti Seyrig, 1937, holotype, §. A. Habitus, lateral view (inset: data labels). B.

Mesosoma, lateral view. C. Metasoma, dorsal view. D. Head, anterior view. E. Mandibles, anterior view.

European Journal of Taxonomy 91: 1 —42 (2014)

Male

Unknown.

Distribution

Democratic Republic of Congo. New record: Cameroon.

Comments

The species is only known from two female specimens. They exhibit a remarkable variation in size, the

BMNH specimen being one third smaller than the impressively large MRAC holotype.

Epirhyssa sit aka sp. nov.

urn: lsid: zoobank, org: act: CF997FDF - A6 A7 -49 A1-9DCA-C5F781C ACEE1

Fig. 8

Diagnosis

Pale yellow overall with isolated dorsal black markings; wings hyaline, slightly infuscate apically; face

rugulose-punctate, wealdy bulging; clypeus shallowly longitudinally strigose with a strong subapical

median tubercle, ventral margin produced laterally; frons smooth, with a weak mid-longitudinal carina;

antenna with 32 flagellomeres; mesopleuron with epicnemial carina reaching level of mesopleural

pit; subtegular ridge convex; fore wing with 2m-cu slightly apical to rs-m; tergite 1 slender; tergite 1

smooth, following tergites densely punctate; apical margins of all tergites straight. CT 3.8; ML 0.7; POL

1.3; OOL 1.2; F1 1 4.5; Fl 15 1.8; Fl 31 1.0; T1 1.7; OT 3.9.

Differential diagnosis

Pale yellow species of relatively small size, otherwise characterized by the combination of the weak

mid-longitudinal carina on the frons, the entirely punctate tergite 2 and the slender tergite 1.

Etymology

Refers to King Shaka Zulu, another kind of population regulator of the Zululand region. He would

certainly have been proud of giving his name to an ic hn eumon

Type material

Holotype

SOUTH AFRICA: $, Natal. 1.5 Ion E of Mtunzini, 28°57’S 31°45’E, 2831 Dd Umlalazi Nat. Res., Nov.

1978, R. M. Miller, indigenous forest. Malaise tr. (NMSA).

Description

Female (holotype)

B 11.9: A 9.6: F 8.7.

Color. Pale dull yellow overall with black parts: mandible, ocellar area (macula isolated from eye

margin), a median spot on each mesoscutal lobe, scuto-scutellar groove, a mid-longitudinal triangle

at base of tergites 4-6: flagellum dark brown; hind tarsus and ovipositor sheath reddish-brown; wings

hyaline, barely infuscate apically, venation yellowish.

Head. Face subquadrate, weakly bulging mid-dorsally, rugulose punctate and laterally smoother; clypeus

shallowly longitudinally strigose, with a strong and acute subapical median tubercle, ventral margin

laterally produced; mandible stout; frons smooth; inner margins of toruli expanded backwards into two

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

'MAGED

WsstfWeb

SAMC 20H

[S. Africa Nfalal. 1.5 km e

Mluiwini 28310cf

:Uni|glaaJ>j a [ f-( f. .■•jQy 7q7<

in dig £• n o us f or esl M a I a i 1 5 e 1 r

0.50 mm

1.0

in n

0.50

0.25 mm

Fig. 8. Epirhyssa shaka sp. nov., holotype, $. A. Habitus, lateral view (inset: data labels). B. Mesosoma,

lateral view. C. Metasoma, dorsal view. D. Head, anterior view. E. Mandibles, anterior view.

European Journal of Taxonomy 91: 1 —42 (2014)

converging carinae, carinae then fused into a weak mid-longitudinal carina reaching median ocellus;

vertex and temple moderately punctate; occipital carina complete; antenna with 32 flagellomeres.

Mesosoma. Mesosoma shallowly and moderately densely punctate, but mesonotum deeply transversely

striate and anterior half of pronotum smooth; epomia moderate; dorsal faint apex of epicnemial carina

reaching level of mesopleural pit; apex of subtegular ridge convex, not flanged laterally; submetapleural

carina slightly and evenly narrowed toward apex.

Wings. Fore wing with 2m-cu slightly apical to rs-m, cu-a slightly apical to Rs+M, Rs moderately

bowed forwards; hind wing with distal abscissa of Cu joining Cu&cu-a close to junction with M.

Metasoma. Tergite 1 moderately slender, sparsely and shallowly punctate; tergite 2 and following densely

punctate; apical margins of all tergites straight. Some other yellow species also sometimes have some

punctures on tergite 2 (E. tombeaodiba ), but the first tergite of the female is then distinctively stouter.

Male

Unknown.

Distribution

South Africa (KwaZulu-Natal).

Epirhyssa tombeaodiba sp. nov.

um:lsid:zoobank.org:act: 134C55D0-CBC2-4788-B199-2197C7344475

Fig. 9

Diagnosis

Bright yellow overall with isolated dorsal black markings; wings hyaline, slightly infuscate apically;

face rugulose-punctate, distinctly protruding dorsally; clypeus shallowly longitudinally strigose with a

strong subapical median tubercle, ventral margin hardly produced laterally; frons smooth, with a weak

mid-longitudinal carina reaching median ocellus; mesopleuron with epicnemial carina reaching level of

mesopleural pit; subtegular ridge convex; fore wing with 2m-cu distinctly apical to rs-m; tergite 1 stout

in female (male unknown); tergite 1 about smooth, following tergites more or less densely punctate;

apical margins of all tergites straight. CT 2.8; ML 0.7; POL 1.3; OOL 1.3; ¥\ { 4.0; Fl 15 2.2; T1 1.2; OT

4.0.

Differential diagnosis

Bright yellow species with wings hyaline, otherwise characterized by the combination of the entirely

punctate tergite 2, the weak mid-longitudinal carina on the frons and the stout tergite 1. It is the only

known yellow Epirhyssa species with both a distinctly sculptured clypeus and a stout tergite 1.

Etymology

Refers to Mrs Jackson’s ambiguous marital status as reported by label data. In the Duala language,

“tombea 6 diba” means “to get married” for a woman.

Type material

Holotype

CAMEROON: $, Nkoemvon [02°48’N 11°08’E], 16 Mar.^1 May 1980, Ms. D. Jackson (BMNH).

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Paratypes

1 5, same label data (BMNH); 1 §, same label data except: 30 Mar-19 Apr. 1980 (BMNH); 1 §, same

label data except: Korap 1981, Mrs D. Jackson (BMNH).

0.50 mm

IMAGED

WsspWeb

SAMC 2013

l6.11i-4.v.19BO. BCsg.

te.O.Jackaon. a

.. . • t.UJT

»■ ..

1.0 mm

0.25 mm

Fig. 9. Epirhyssa tombeaodiba sp. nov., holotype, $. A. Habitus, lateral view (inset: data labels). B.

Mesosoma, lateral view. C. Metasoma, dorsal view. D. Head, anterior view. E. Mandibles, anterior view.

European Journal of Taxonomy 91: 1 —42 (2014)

Description

Female (4 specimens)

B 7.2-8.9; A9.5; F 9.0-10.5. (holotype B: 8.0; ANA; F 9.9).

Color. Bright yellow overall with black parts: mandible, ocellar area (macula connected to eye margin),

a median spot on each mesoscutal lobe, scuto-scutellar groove, a mid-longitudinal macula of variable

length on tergites 4-7; flagellum dark brown; flagellum and ovipositor sheath reddish-brown; wings

hyaline, slightly infuscate apically, venation brown.

Head. Face subquadrate, rugulose-punctate, distinctly protruding dorsally; clypeus longitudinally

strigose, with a strong subapical median lobe, ventral margin hardly produced laterally; mandible stout;

frons, vertex and temple sparsely and shallowly punctate; inner margins of toruli expanded backwards

into two converging weak carinae, carinae then fused into a mid-longitudinal carina reaching median

ocellus; occipital carina mid-dorsally interrupted; antenna with at least 22 flagellomeres (no specimen

with complete flagellum).

Mesosoma. Mesosoma shallowly and moderately densely punctate, but mesonotum deeply transversely

striate and anterior half of pronotum smooth; epicnemial carina reaching level of mesopleural pit; apex

of subtegular ridge convex, not flanged laterally; submetapleural carina slightly enlarged anteriorly, then

evenly narrowed toward apex, strongly notched at anterior third in one paratype.

Wings. Fore wing with 2m-cu distinctly apical to rs-m, cu-a barely apical to Rs&M, and Rs moderately

bowed forwards; hind wing with distal abscissa of Cu joining Cu&cu-a shortly below junction with M.

Metasoma. Tergite 1 moderately stout, nearly smooth; following tergites shallowly and more or less

densely punctate, punctation progressively finer toward apex; apical margins of all tergites straight.

Distribution

Cameroon.

Epirhyssci uelensis Benoit, 1951

Fig. 10

Diagnosis

Bright yellow overall with isolated dorsal black markings; wings hyaline, slightly infuscate apically; face

rugulose-punctate and moderately protruding dorsally; clypeus nearly smooth, with median subapical

tubercle strong, ventral margin produced laterally; frons nearly smooth, with a mid-longitudinal

carina reaching median ocellus and lateral curved carina toward lateral ocellus; antenna with 32-34

flagellomeres; mesopleuron with epicnemial carina reaching level of mesopleural pit; subtegular ridge

concave, laterally flanged; fore wing with 2m-cu slightly apical to rs-m; tergite 1 stout in female,

very slender in male; tergites 1-3 smooth, following tergites shallowly punctate with basal transverse

striations; apical margins of tergites 3-6 moderately concave. CT 2.9; ML 0.6; POL 1.2; OOL 1.3;

3.8; Fl 15 2.2; FL 33 0.9; T1 1.2—1.3; OT 4.1.

Differential diagnosis

A mostly bright yellow species with hyaline wings, otherwise characterized by the combination of the

concave subtegular ridge, the mid-longitudinal carina on the frons and the stout tergite 1 of the female.

The only other species with a concave and flanged subtegular ridge is E. gavinbroadi , in which the

clypeus is distinctly punctate.

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Material examined

Holotype

DEMOCRATIC REPUBLIC OF CONGO: $, Haut-Uele, Paulis [Isiro, 03°28’ N 25°43’E], xii-1947,

PL.G. Benoit (MRAC).

4 HOUtTYFUS

COM.. Mils, ri.NHd E IMAGED

Haut-Uele : Paulis | vfospWeb

• <ii -iy-'i7|fSAMC20l 3

P. L. G.Benoit ■■ 11 '

2.0 mm

I-1

0.50 mm

1.0

’ ■

0.25 mm

0.25

Fig. 10. Epirhyssa uelensis Benoit, 1951, holotype, §. A. Habitus, lateral view (inset: data labels). B.

Mesosoma, lateral view. C. Metasoma, dorsal view. D. Head, anterior view. E. Mandibles, anterior view.

European Journal of Taxonomy 91: 1 —42 (2014)

Paratype

DEMOCRATIC REPUBLIC OF CONGO: 1 Bambesa, Jul. 1933, H.J. Bredo (MRAC).

Other material

CAMEROON: 1 §, Nlcoemvon, 30 Mar —19 Apr. 1980, Ms. D. Jackson (BMNH); 2 5?, same label

data except 13 Jul.M Aug. 1980 (BMNH); 1 §, same label data except Oct.-Nov. 1980 (BMNH).

Description

Female (5 specimens)

B 10.4-12.8; A9.1-12.0;F 8.6-11.5 (holotypeB: 10.4; A9.1; F 8.6).

Color. Bright yellow overall, sometimes with infuscate to black mid-longitudinal stripes on tergites

5-7, and black markings: mandible, ocellar area (macula connected to eye margin), upper occiput and

mesoscutum but lateral margins and a large central macula, sometimes apex of scutellum, basal margin

and apex of propodeum; ovipositor sheath reddish testaceous; wings hyaline, wealdy infuscate apically,

venation brown.

Head. Face rugulose-punctate and moderately protruding dorsally; clypeus smoothly sculptured, with a

strong subapical median tubercle, ventral margin moderately produced laterally and medially; mandible

stout; frons nearly smooth with sparse fine punctures; inner margins of toruli expanded backwards

into two converging weak carinae, carinae then fused into a mid-longitudinal carina reaching median

ocellus; frons also with an additional lateral curved carina from posterior margin of torulus to next

to lateral ocellus; vertex and temples sparsely punctate, inter-ocellar area somewhat rugose; occipital

carina mid-dorsally evanescent; hypostomal carina strongly raised above mandibular base; antenna with

32-34 flagellomeres.

Mesosoma. Mesosoma shallowly and moderately densely punctate, but mesonotum deeply transversely

striate and anterior half of pronotum smooth; epomia moderate; dorsal faint apex of epicnemial carina

nearly reaching level of mesopleural pit; apex of subtegular ridge concave, weakly to distinctly flanged

laterally; submetapleural carina slightly and evenly narrowed toward apex.

Wings. Fore wing with 2m-cu slightly apical to rs-m, cu-a slightly apical to Rs+M, Rs strongly bowed

forwards; hind wing with distal abscissa of Cu joining Cu&cu-a close to junction with M.

Metasoma. Tergite 1 stout and smooth, tergites 2-3 smooth, tergites 3-6 shallowly punctate with weak

transverse striations basally; apical margins of tergites 3-6 moderately concave, others straight.

Male (1 specimen)

B 8.5; A 8.2; F 7.9.

Similar to female but metasoma much slenderer (T1 2.0).

Distribution

Democratic Republic of Congo. New record: Cameroon.

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Epirhyssa villemantae sp. nov.

urn:lsid:zoobank. org: act: C701D367-2EF 0-4A73 -8D41 -F 8 F 5EDADAA21

Fig. 11

Diagnosis

Head and pronotum yellowish-orange; remainder of mesosoma black; metasoma orange; wings blackish

with a pale subapical patch; face sparsely punctate and moderately protruding dorsally; clypeus smooth,

with median subapical tubercle moderate, ventral margin truncate; frons laterally smooth, medially

transversely striate, with two submedian carinae diverging toward lateral ocelli; antenna with 36

flagellomeres; mesopleuron with epicnemial carina reaching level of mesopleural pit; subtegular ridge

convex; fore wing with 2 m-cu slightly apical to rs-m; tergite 1 stout in female (male unknown); tergites

1-3 smooth, following tergites moderately punctate with basal transverse striations; apical margins of

all tergites straight. CT 2.0; ME 0.6; POE 1.3; OOL 1.6; Flj 3.6; Fl 15 2.1; FL 33 0.9; T1 1.3; OT 4.4.

Differential diagnosis

A species with unique color pattern, with most of mesosoma black, metasoma orange and wings blackish

with a subapical light patch; otherwise characterized by the sparsely punctate face, the truncate ventral

margin of the clypeus and the stout tergite 1 of the female.

Etymology

Dedicated to Claire Villemant, MNHN curator, hymenopterologist and ichneumonomaniac, adviser,

teacher, supporter, coach and friend.

Type material

Holotype

NIGERIA: §, IlorinProv. [06°48’N 05°18’E], 18 Jun. 192 [sic], De. G.W.S. Macfie, Pres, by Imp. Bur.

Ent. 1921-129 (BMNH).

Description

Female (holotype)

B 17.1; A 13.1; F 13.1.

Color. Head and pronotum yellowish-orange with mandible and ocellar area black; remainder of

mesosoma blackish testaceous; metasoma orange; fore leg and mid leg from tibia yellowish orange,

remainder of legs blackish testaceous; antenna and ovipositor sheath brown; wings strongly infuscate

but with large subapical and small mid-dorsal pale yellowish patches on fore wing.

Head. Face sparsely punctate, almost smooth laterally, and moderately protruding dorsally; clypeus

smooth, with a moderate subapical median tubercle, ventral margin truncate; mandible stout; frons

laterally smooth, medially transversely striate; inner margin of toruli expanded backwards into two

weak sinuate submedian carinae, carinae then diverging and reaching lateral ocelli; frons also with an

additional lateral curved carina from posterior margin of torulus to next to lateral ocellus; vertex and

temples almost smooth, inter-ocellar area somewhat rugose; occipital carina mid-dorsally evanescent;

hypostomal carina strongly raised above mandibular base; antenna with 36 flagellomeres.

Mesosoma. Mesosoma finely, sparsely and shallowly punctate, but mesonotum transversely striate

and pronotum, speculum area and propodeum medially smooth; epomia very weak; dorsal faint apex

of epicnemial carina reaching level of mesopleural pit; apex of subtegular ridge convex, not flanged

laterally; submetapleural carina moderate and even all along.

European Journal of Taxonomy 91: 1 —42 (2014)

Fig. 11. Epirhyssa villemantae sp. nov., holotype, §. A. Habitus, lateral view (inset: data labels). B.

Mesosoma, lateral view. C. Metasoma, dorsal view. D. Head, anterior view. E. Mandibles, anterior view.

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Wings. Fore wing with 2m-cu slightly apical to rs-m, cu-a slightly apical to Rs+M, Rs moderately

bowed forwards; hind wing with distal abscissa of Cu joining Cu&cu-a at its anterior third, remote from

junction with M.

Metasoma. Tergite 1 moderately stout; tergites 1-3 smooth, following tergites moderately punctate with

weak transverse basal striations; apical margins of all tergites straight.

Male

Unknown.

Distribution

Nigeria.

Genus Megarliyssa Ashmead, 1900

Thalessa Holmgren, 1859: 122.

Megalorhyssa Shulz, 1906: 115.

Eurhyssa Derksen, 1941: 721.

Diagnosis (updated from Townes 1969)

Large to very large insects (fore wing length 10-30 mm); clypeus small, transversely rectangular,

ventral margin laterally and sometimes medially bluntly produced; occipital and hypostomal carinae

joining above mandible base; mandibular teeth subequal, lower tooth pointed and upper tooth more or

less chisel shaped; propodeum of moderate length, without carina dorsally but pleural carina distinct;

fore wing with areolet closed except in occasional dwarf males, receiving 2m-cu within its apical half,

pterostigma about 5.5x longer than wide; hind wing with distal abscissa of Cu present, joining Cu&cu-a

near or at junction with M; mid trochantellus with a ventral longitudinal ridge; tarsal claws simple, large;

tergite 1 fused with its sternite and without glymma; tergites 3-6 almost smooth to finely and sparsely

punctate and with isolated aciculate areas; female with sternites 2—4 each with a pair of tubercles near

anterior margin, and with an apical truncate hom-like process on last tergite; male with metasoma

strongly depressed, gonosquama lanceolate, strongly depressed, with a sharp piliferous groove along

inner lower margin and a short subapical piliferous groove on outer face.

Species richness and distribution (Gauld 1984; Yu et al. 2012)

Megarhyssa is mostly a Holarctic and Oriental genus, with one species introduced into Australia and

New Zealand for biocontrol purposes, and a single species reported from the Afrotropical Region.

Megarhyssa babaulti Seyrig, 1937

Fig. 12

Megarhyssa babaulti kapangaensis Seyrig, 1937: 116.

Megarhyssa babaulti kapangaensis var. rubra Benoit, 1951: 385.

Megarhyssa babaulti kapangaensis var. rubrernma Benoit, 1951: 385.

Diagnosis

Head orange with a black part of variable extent; mesosoma orange, posterior third sometimes dark

brown; metasoma mostly black with pale yellow or orange markings of variable extent; flagellum two-

colored, basally black and apically orange; wings blackish with lighter patch of variable extent; face

moderately punctate to smooth, flat; clypeus smooth, with median subapical tubercle weak, ventral

margin strongly produced laterally; frons smooth with a pair of submedian carinae diverging toward

European Journal of Taxonomy 91: 1 —42 (2014)

lateral ocelli; epicnemial carina laterally absent; subtegular ridge convex; fore wing with areolet strongly

petiolate, receiving 2m-cu at its distal apex, opposite 3rs-m; tergite 1 very stout; tergites 1-4 smooth

or mostly so, following tergites finely and densely punctate; apical margins of tergites 3-5 strongly

concave. CT 3.4; ML 0.6; POL 1.2; OOL 1.7; Flj 4.4; Fl 15 2.3; Fl 39 1.0; T1 1.1; OT 4.8.

Fig. 12. Megarhyssa babaulti Seyrig, 1937, holotype, $. A. Habitus, lateral view (inset: data labels). B.

Mesosoma, lateral view. C. Metasoma, dorsal view. D. Head, anterior view. E. Mandibles, anterior view.

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

Material examined

Holotype

DEMOCRATIC REPUBLIC OF CONGO: $, reg. Lac Kivu, Kadjudju [Kajuju, 02°09’S 28°54’E],

1932, G. Babault, MNHN EY8831.

Description

Female (holotype)

B 22.1; A 15.5; F 18.

Color. Head orange with isolated black parts to almost entirely black; antenna basally black, fading to

orange from flagellomere 12; mesosoma orange, posterior third varying from orange to entirely black;

legs orange with mid and fore legs variably orange to dark brown; metasoma black with tergites 1-3

varying from orange to black, tergite 4 and following largely pale yellow laterally, sternites pale yellow;

ovipositor sheath black, apical half fading to reddish-orange; wings blackish with lighter yellowish parts

of variable extension (color variation: Seyrig 1937; Benoit 1951).

Head. Face flat, laterally smooth, medially moderately punctate; clypeus smooth, with a weak median

subapical tubercle, ventral margin strongly produced laterally; upper head almost smooth; frons with a

pair of weak submedian carinae, carinae diverging below median ocellus, inner margins of toruli hardly

produced backwards; occipital carina complete, mid-dorsally weak; antenna with 40 flagellomeres.

Mesosoma. Mesosoma finely and moderately densely punctate, but mesonotum deeply transversely

striate and anterior half of pronotum smooth; epomia indistinct; epicnemial carina absent except for

a faint ventral remnant; apex of subtegular ridge convex, not flanged laterally; submetapleural carina

barely expanded anteriorly and evenly narrowed toward apex.

Wings. Fore wing with areolet strongly petiolate, receiving 2m-cu at its apex, cu-a strongly apical to

Rs&M, Rs strongly bowed forwards; hind wing with distal abscissa of Cu joining Cu&cu-a shortly

above middle of Cu&cu-a.

Metasoma. Tergite 1 stout, nearly as long as apically wide; tergites 1-3 quite smooth, tergite 4 smooth

with some fine punctures basally, following tergites finely and densely punctate; apical margins of

tergites 3-5 strongly concave.

Male

Unknown.

Distribution

Democratic Republic of Congo.

Discussion

We face here an apparent paradoxical observation: despite the historical general interest they have

received and their expected high diversity in the tropics, very few species of Rhyssinae have been

described in Africa, and the African and European museums house only scarce, undetermined material.

Discussing the zoogeographical history of Epirhyssa in the Neotropical region, Porter (1978, 1982)

showed that their abundance, distribution and phenology are strongly related to the humidity level:

dry periods and arid areas are major ecological obstacles for Rhyssinae, hence they are expected to

be highly diversified in lowland tropical humid forests. Nevertheless, the collection efforts carried out

by the Iziko South African Museum in Gabon, Uganda, Central African Republic and South Africa

European Journal of Taxonomy 91: 1 —42 (2014)

Table 1. Summary of the main collecting efforts by Simon van Noort of SAMC in Africa over the last

22 years (MTM: Malaise trap months; YPTM: yellow pan trap months).

Country

Year(s)

MTM

YPTM

Sweeps

Rhyssinae

collected

Gabon

2000

3000

Central African Republic

2001

6000

Uganda

2005 & 2008

Not quantified

South Africa

1992-present

c. 2000

c. 4500

c. 30 000

(Table 1) produced only two specimens, despite the fact that a large proportion of the central African

sampling took place in lowland rainforest. On the other hand, examination of the mere 15 undetermined

specimens gathered for this study revealed that half of them belonged to five new species, four of them

being singletons.

Thus, Rhyssinae appear very rare though greatly diversified in the Afrotropical region. Their rarity may

be explained by ecological hypotheses. Rhyssinae are indeed known to parasitize primarily Siricidae and

Xiphydriidae in the Northern Hemisphere (Yu et al. 2012), w hil e these families are poorly represented in

the tropics. Virtually nothing is known about the ecology of Epirhyssa , but as some Holarctic Rhyssinae

are also known to exploit Cerambycidae, they have been hypothesized to develop mostly on wood¬

boring coleopteran larvae (Gauld 1991). This ecological niche, however, is also exploited by other taxa:

Braconinae, for example, have been diversifying in Africa for a long time, and one plausible hypothesis

is that the competition was disadvantageous for Rhyssinae (G. Broad, pers. comm.).

Somewhat contradictory with his own observations, Porter (1978, 1982) stated that the largest lowland

rainforest of the world in the Amazon Basin was rather depleted of Rhyssinae. Following their own

collection efforts in this region, Saalcsjarvi et al. (2004) and Veijalainen et al. (2013), however, showed

that Porter’s biogeographical assumptions were mainly influenced by sampling bias: intensive local

collection efforts are needed to uncover most of the diversity of Rhyssinae and other hymenopteran

groups. Saalcsjarvi et al. (2004) report a sampling effort of 185 Malaise trap months resulting in 12 new

species. These figures of return relative to sampling effort appear similar to ours from tropical areas of

Africa. We predict that most of the Afrotropical diversity of Rhyssinae has still to be discovered, but

intensive efforts of long-term sampling are required to unveil them.

Acknowledgments

We are grateful to the curators of the above mentioned museums for loaning specimens, and once again

to Gavin Broad for his advice. Simon van Noort was funded by South African NRF (National Research

Foundation) grants: GUN 2068865; GUN 61497; GUN 79004; GUN 79211; GUN 81139. Part of the

South African field work conducted by SvN was funded by the National Science Foundation under

PlatyPBI grant No. DEB-0614764 to N.F. Johnson and A.D. Austin. Pascal Rousse was funded by

SABI (South African Biodiversity Initiative) NRF post-doctoral fellowship GUN 81609, a Claude Teon

Foundation post-doctoral fellowship, and Societe Entomologique de France (Bourse Germaine Cousin).

Cape Nature, the Eastern Cape Department of Environmental Affairs and the Northern Cape Department

of Nature and Environmental Conservation provided collecting permits for South Africa. Fieldwork in

Gabon was funded by the World Bank through WWF-US and WWF-CARPO. Collecting and export

permits were granted by Emile Mamfoumbi Kombila, Directeur de la Faune et de la Chasse, Libreville.

The Ugandan Wildlife Authority and UNCST provided permits to conduct research in Uganda. Field

work in Central African Republic was supported by WWF-US and WWF-CARPO. The Ministers of

Water, Forests and the Environment and the High Commissioners for tertiary Education and Research

of the Central African Republic granted permission to carry out the inventory survey and to export the

ROUSSE R & VAN NOORT S., Review of Afrotropical Rhyssinae

specimens as part of the WWF-US CAR field expedition conducted in 2001. Disclaimer: the attributed

names referring to historical personages are purely functional: neither of the authors approves, nor

supports any aspect of the atypical policy of King Shaka Zulu nor any other monarch of the 19 th Century.

References

Baltazar C.R. 1961. The Philippine Pimplini, Poeminiini, Rhyssini, and Xoridini (Hymenoptera,

Ichneumonidae, Pimplinae). Monographs of the National Institute of Science and Technology 7: 1-130.

Benoit PL.G. 1951. Les Rhyssini ethiopiens (Pimplinae - Ichneumonidae). Revue de Zoologie et de

Botanique Africaines 44: 382-385.

Benoit PL.G. 1952. Notes Ichneumonologiques Africaines II. Revue de Zoologie et de Botanique

Africaines 46: 51-59.

Buffington M.L., Burks R. & McNeil L. 2005. Advanced Techniques for Imaging Parasitic Hymenoptera.

American Entomologist 51: 50-54.

Buffington M.L. & Gates M. 2009. Advanced imaging techniques II: using a compound microscope for

photographing point-mount specimens. American Entomologist 54: 222-224.

Gauld I.D. 1984. An Introduction to the Ichneumonidae of Australia. British Museum of Natural History,

London, UK.

Gauld I.D. 1991. The Ichneumonidae of Costa Rica, 1. Memoirs of the American Entomological Institute

47: 1-589.

Gauld I.D., Wahl D.B., Bradshaw K., Hanson P. & Ward S. 1997. The Ichneumonidae of Costa Rica, 2.

Memoirs of the American Entomological Institute 57: 1—485.

HansonH.S. 1939. Ecological notes on the Sirex wood wasps and their parasites. Bulletin of Entomological

Research 30: 27-76. http://dx.doi.org/10.1017/S00074853000Q4399

Horstmann K. 2002. Revisionen von Schlupfwespen-Arten VI (Hymenoptera: Ichneumonidae).

Mitteilungen der Miinchener Entomologischen Gesellschaft 92: 79-91.

Jansson R., Rodriguez-Castaneda G. & Harding L.E. 2013. What can multiple phylogenies say about

the latitudinal diversity gradient? A new look at the tropical conservatism, out-of-the-tropics and

diversification rate hypotheses. Evolution 7: 1741-1755. http://dx.doi.org/10. Ill 1/evo. 12089

KamathM.K. & Gupta V.K. 1972. Ichneumonologia Orientalis, Part II. The Tribe Rhyssini (Hymenoptera:

Ichneumonidae). Oriental Insects Monographs 2: 1-300.

Kasparyan D.R. & Khalaim A.I. 2007. Pimplinae, Tryphoninae, Eucerotinae, Xoridinae, Agriotypinae,

Lycorininae, Neorhacodinae, Ctenopelmatinae, Phrudinae, Ophioninae, Acaenitinae, Collyriinae,

Mesochorinae. Key to the insects of Russia Far East. Vol. IV. Neuropteroidea, Mecoptera, Hymenoptera

5. Academy of Sciences, Saint-Petersburg.

Kerr PH., Fischer E.M. & Buffington M.L. 2008. Dome lighting for insect imaging under a microscope.

American Entomologist 54: 198-200.

Porter C.C. 1978. A revision of the genus Epirhyssa (Hymenoptera, Ichneumonidae). Studia Entomo-

logica 20: 297-318.

Porter C.C. 1982. Systematics and zoogeography of the neotropic genus Epirhyssa (Hymenoptera:

Ichneumonidae). National Geography Society Research Reports 14: 513-522.

Quicke D.L.J. 2012. We know too little about parasitoid wasp distributions to draw any conclusions

about latitudinal trends in species richness, body size and biology. PLoS One 7: 1-9. http://dx.doi.

org/10.1371/iournal.pone.0032101

European Journal of Taxonomy 91: 1 —42 (2014)

Saaksjarvi I.E., Haataja S., Neuvonen S., Gauld I.D., Jussila R., Salo J. & Burgos A.M. 2004. High

local species richness of parasitic wasps (Hymenoptera: Ichneumonidae; Pimplinae and Rhyssinae)

from the lowland rainforests of Peruvian Amazonia. Ecological Entomology 29: 735-743. http://

dx.doi.org/10,111 l/i.0307-6946.2004.00656.x

Seyrig A. 1937. Sur les Rhyssini de la faune ethiopienne (Hym. Ichneumonidae). Revue de Zoologie et

de Botanique Africaines 30: 112-116.

Sheng M.-L. & Sun S.-P. 2010. Parasitic Ichneumonids on Woodborers in China (Hymenoptera:

Ichneumonidae). Science Press, Beijing, China.

Spradbery J.P 1968. A technique for artificially culturing ichneumonid parasites of woodwasps

(Hymenoptera: Siricidae). Entomologia Experimentalis et Applicata 11: 257-260.

Spradbery J.P. 1970. Host finding by Rhyssa persuasoria (L.), an ichneumonid parasite of siricid

woodwasps. Animal Behaviour 18: 103-114. http://dx.doi.org/10.1016/0003-3472(70)90077-1

Townes H.K. 1969. The genera of Ichneumonidae, part 1. Ephialtinae to Agriotypinae. Memoirs of the

American Entomological Institute 11: 1-300.

Townes H.K. 1975. The parasitic Hymenoptera with the longest ovipositors, with descriptions of two

new Ichneumonidae. Entomological News 86: 123-127.

Townes H.K., Townes M., Walley G.S., Walldey L., Habeck D. & Townes G. 1960. Ichneumon-flies of

America north of Mexico: 2. Subfamilies Ephialtinae, Xoridinae, Acaenitinae. United States National

Museum Bulletin 216: 1-676. http: //www. biodiversity library, org / item/32635#page/5/mode/1 up

Veijalainen A., Saaksjarvi I.E., Erwin T.L., Gomez I.C. & Longino J.T. 2013. Subfamily composition

of Ichneumonidae (Hymenoptera) from western Amazonia: insights into diversity of tropical parasitoid

wasps. Insect Conservation and Diversity 6: 28-37. http://dx.doi.Org/10.llll/j.1752-4598.2012.00185.x

Wahl D.B. & Gauld I.D. 1998. The cladistics and higher classification of the Pimpliformes (Hymenoptera:

Ichneumonidae). Systematic Entomology23: 265-298. http://dx.doi.org/10.1046/i. 1365-3113.1998.00057.x

Wahl D.B. & Sharkey M.J. 1993. Chapter 10. Superfamily Ichneumonoidea. In: Goulet H. & Huber J.T.

(eds) Hymenoptera of the World: An Identification Guide to Families'. 359-509. Agriculture Canada,

Ottawa.

Yu D.S.K., Van Achterberg C. & Horstmann K. 2012. Taxapad2012, Ichneumonoidea 2011. Database

on flash-drive, www.taxapad.com. Ottawa.

Manuscript received: 14 February 2014

Manuscript accepted: 10 June 2014

Published on: 25 July 2014

Topic editor: Koen Martens

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; Botanic Garden Meise,

Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London,

United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History

Museum of Denmark, Copenhagen, De nm ark