Research Article |
Corresponding author: Ralph S. Peters ( r.peters@zfmk.de ) Academic editor: Petr Janšta
© 2018 Jennifer Werner, Ralph S. Peters.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Werner J, Peters RS (2018) Taxonomic revision of the genus Oodera Westwood, 1874 (Hymenoptera, Chalcidoidea, Pteromalidae, Cleonyminae), with description of ten new species. Journal of Hymenoptera Research 63: 73-123. https://doi.org/10.3897/jhr.63.12754
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The world species of Oodera Westwood, 1874 (Chalcidoidea: Pteromalidae: Cleonyminae: Ooderini) are revised. We examined 115 specimens of this rarely collected genus and based on morphological characters assign 110 specimens to 20 recognised species, of which the following ten are described as new: Oodera circularicollis sp. n. (Morocco), O. felix sp. n. (Central African Republic), O. fidelis sp. n. (Vietnam), O. florea sp. n. (Thailand), O. heikewernerae sp. n. (Botswana and South Africa), O. leibnizi sp. n. (Papua New Guinea, Malaysia and Phillippines), O. mkomaziensis sp. n. (Tanzania), O. namibiensis sp. n. (Namibia), O. niehuisorum sp. n. (Egypt and Israel), and O. srilankiensis sp. n. (Sri Lanka). Oodera monstrum Nikol’skaya, 1952, syn. n., is synonymised under O. formosa (Giraud, 1863). Five specimens could not be assigned to species and are treated as Oodera sp. Redescriptions are provided for all previously described valid species. Oodera albopilosa Crosby, 1909 is excluded from Oodera and transferred to Eupelmus Dalman, 1820 (Eupelmidae) as E. albopilosa (Crosby, 1909) n. comb. Oodera rufimana Westwood, 1874 and O. obscura Westwood, 1874 are treated as nomina dubia because we were unable to locate type specimens and the original descriptions are not sufficiently informative to clarify the taxonomic status of these names. Several specimens from North America are identified as introduced specimens of the European species O. formosa. We provide images and diagnostic characters for all 20 included species and an identification key to species.
Taxonomy, parasitoid wasps, identification key, new species
Oodera Westwood, 1874 is a morphologically extraordinary and conspicuous genus in Chalcidoidea. The group is currently classified as the monotypic tribe Ooderini in the Cleonyminae, a subfamily of the polyphyletic Pteromalidae (
It included 14 species prior to this revision.
The phylogenetic position and proper classification of Oodera remain unclear. When
All Oodera species are thought to be parasitoids of woodboring beetles (mainly Buprestidae and Scolytinae) but only very few host-parasitoid records are actually verified or even vouchered (
The genus has never been taxonomically revised. Of the 14 valid species described prior to this revision, all were described in isolation or with just another species. The only taxonomic actions so far, apart from new combinations to transfer species to Oodera, were by
Oodera are rarely collected and are, accordingly, also rare in scientific collections. The taxonomic revision presented here is based on 115 specimens. Most specimens are old (>10 years), dry-pinned material that does not allow routine DNA extraction and molecular sequencing. Our revision is based solely on morphological characters. We made high-resolution images of all relevant available type and non-type material and used a variety of qualitative features as well as morphometric ratios that describe body proportions as diagnostic characters to differentiate between species. Based on this, we describe new species, clarify the status of previously described species and redescribe them, and provide an identification key to all species considered valid after the taxonomic revision.
This study is based on material borrowed from the following museums (with abbreviations used in the text):
ZIRAS Zoological Institute of Russian Acedemy of Science, St. Petersburg, Russia.
The terminology in this study follows
Both sexes were used for description of all characters, except for shape and size of the metasoma and the size of the ovipositor. Detailed character definitions are given in Table
List of characters used in this study, with character definition and abbreviation as also used in Suppl. material
Abbreviation | Character | Definition |
---|---|---|
bdy.l | Body length | Length of body, excluding the ovipositor in females, dorsal view |
hea.h | Head height | Distance between lower edge of clypeus and lower edge of anterior ocellus, frontal view |
hea.l | Head length | Maximum length of head, lateral view |
hea.w | Head width | Maximum width of head, frontal view |
eye.h | Eye height | Height of eye, lateral view |
eye.l | Eye length | Length of eye, lateral view |
msp.l | Malar space | Distance between the points where malar sulcus touches mouth margin and lower edge of eye, lateral view |
lof.h | Lower face height | Distance between lower margin of torulus and margin of clypeus |
eya.d | Eye to antennal toruli | Distance between the lower margin of eye and lateral margin of antennal toruli, frontal view |
tor.d | Distance antennal toruli | Distance between the inner margins of the toruli, frontal view |
eye.d | Eye distance | Shortest distance between eyes, dorsal view |
pol.l | POL | Shortest distance between posterior ocelli, dorsal view |
ool.l | OOL | Shortest distance between posterior ocellus and eye margin, dorsal view |
cor.l | Corona length | Maximum length of corona, frontal view |
cor.w | Corona width | Maximum width of corona, frontal view |
scp.l | Scape length | Length of scape exclusive of radicle, outer aspect |
pdl.l | Pedicel length | Length of pedicel, outer aspect |
flg.l | Flagellum length | Length of all flagellomers |
fun.l | Funicule length | Length of flagellum excluding clava |
clv.l | Clava length | Length of clava, outer aspect |
no.l | Pronotum length | Length of pronotum along median line, dorsal view |
no.w | Pronotum width | Maximum width of pronotum, dorsal view |
msn.l | Mesonotum length | Length of mesonotum along median line from anterior edge of mesoscutum to posterior edge of mesoscutellum, dorsal view |
msc.w | Mesoscutum width | Maximum width of mesoscutum just anterior to tegulae, equals mesonotum width, dorsal view |
msc.l | Mesoscutum length | Length of mesoscutum along median line from posterior edge of pronotum to posterior edge of mesoscutum, dorsal view |
sct.l | Mesoscutellum length | Length of mesoscutellum along median line from posterior edge of mesoscutum to posterior edge of scutellum, dorsal view |
sct.w | Mesoscutellum width | Maximum width of mesoscutellum, dorsal view |
sam.l | Mesoscutellum anterior margin | Length of part of mesoscutellum from anterior margin of mesoscutellum to imaginary transverse line connecting posterior margins of axillae, measured along median line, dorsal view |
ppd.l | Propodeum length | Length of propodeum measured along median line from anterior edge of propodeum to posterior edge of nucha, dorsal view |
ppd.w | Propodeum width | Maximum width of propodeum, dorsal view |
tb1.l | Protibia length | Length of protibia along median line, outer aspect |
fm1.l | Profemur length | Length of profemur, from distal end of trochanter to tip of profemur, measured along median line, outer aspect |
fm1.w | Profemur width | Maximum width of profemur, outer aspect |
fwi.l | Fore wing length | Maximum length of fore wing, measured from end of humeral plate to tip of wing |
fwi.w | Fore wing width | Maximum width of fore wing |
cc.l | Costal cell length | Length of the costal cell, measured from the basal constriction that delimits the apex of the humeral plate of the wing to the point at which the submarginal vein touches the leading edge of the wing |
mav.l | Marginal vein length | Length of marginal vein, distance between the point at which the submarginal vein touches the leading edge of the wing and the point at which stigmal vein and postmarginal vein unite |
pmv.l | Postmarginal vein length | Length of postmarginal vein, distance between the point at which stigmal vein and postmarginal vein unite, apically to where the vein appears to end |
stv.l | Stigmal vein length | Length of stigmal vein, distance between the point at which stigmal vein and postmarginal vein unite apically, and the distal end of the stigma |
mts.l | Metasoma length | Length of metasoma along median line, from end of nucha to tip of metasoma, excluding ovipositor, dorsal view |
mts.w | Metasoma width | Maximum width of metasoma, dorsal view |
ovp.l | Ovipositor length | Length of the visible part of ovipositor, dorsal view |
We calculated a number of ratio characters to describe shape and dimensions of body parts, and assigned ratios to categories to allow easier diagnosis and use in the identification key. These ratios and categories – used throughout the text – are given in Table
Characters and categories used in the text, with respecitive ranges of measured ratios.
Character | Categories and Ranges | ||
---|---|---|---|
Body shape (mesonotum length/mesonotum width = mesoscutum width) | robust | medium | slender |
<1.4 | 1.4–1.44 | >1.44 | |
Head shape in lateral view (head height/head length) | round | oval | elongated |
<1.45 | 1.45–1.6 | >1.6 | |
Eye size (eye height/head height) | small | large | |
<0.6 | ≥0.6 | ||
Corona shape (corona length/corona width) | thick | medium | slim |
<4.1 | 4.1–7.0 | >7.0 | |
Mesoscutellum shape (mesoscutellum length/mesoscutellum width) | normal | slender | |
<0.75 | ≥0.75 | ||
Mesoscutellum anterior margin (part anterior to virtual transverse line between posterior margins of axillae/mesoscutellum length) | hardly convex | convex | |
≤0.33 | >0.33 | ||
Propodeum size (propodeum length/mesoscutum length) | small | medium | large |
<0.08 | 0.08–0.15 | >0.15 | |
Profemur shape (profemur length/profemur width) | robust | medium | elongated |
<1.99 | 2.0–2.2 | >2.2 | |
Marginal vein length (marginal vein length/postmarginal vein length) | short | medium | long |
<0.9 | 0.9–1.10 | >1.10 | |
Metasoma length (metasoma length/body length) | short | medium | long |
<0.46 | 0.46–0.5 | >0.5 | |
Ovipositor length (ovipositor length/metasoma length) | short | rather long | |
<0.16 | ≥0.16 | ||
Body length (in mm) | small | medium | large |
<7.0 | 7.0–9.0 | >9.0 |
For morphological examination and measurements we used an Olympus SZX12-ILLK stereomicroscope with an eyepiece micrometer (1 mm divided in 100 units). Magnification ranged from 11.2× (16× eyepiece, 0.7× objective) to 144× (16× eyepiece, 9× objective). For photographs, we used a BK Lab Imaging system (Dun, Inc. 2016).
Photos were taken with a Canon EOS 7D, serially in 30 to 40 different focal distances. The camera was equipped with a Tamron Tele-Converter SP AF 1.4X, a Canon AC Adapter Kit ACK E6-(CBCB) and with different objectives (CF1B, CF2, CF3, CF4 or micro/macro-objective) that were selected depending on specimen or character size. For the lighting, the camera’s own flash light and three movable light sources were used. A dull Plexiglas tube was placed over the specimen to avoid reflection from direct light.
Adobe Lightroom 5 was used for initial evaluation and for storing the photos. Then, the photos were stacked with Zerene Stacker and Helicon Focus and modified with GIMP 2.8.14 and Adobe Photoshop CS6. For scaling, photos of a 1 mm scale were made with all used objectives.
The image plates were made with PowerPoint 2010.
We examined a total of 115 specimens (24 males, 85 females, four of undetermined sex due to missing metasoma (= 113) plus two females examined from images). All 113 examined and measured specimens were given a unique code number referring to this study, composed of a species name abbreviation (e.g., OFo for O. formosa) and a consecutive number that was added to the specimen on a separate label. All measurements and calculated ratios for each specimen are given in Suppl. material
Both sexes. Comparatively large-bodied (3.6–17 mm). Parascrobal area of head raised and with crest-like structure (= corona) (Fig.
Four different types of corona structure in Oodera, in dorso-frontal view. Antennal scrobes are left of corona, eye is right of corona; a interrupted (exemplar species imaged: O. namibiensis sp. n.) b continuous (exemplar species imaged: O. formosa) c square (exemplar species imaged: O. longicollis) d three- to four-part (exemplar species imaged: O. leibnizi sp. n.). For additional definitions of corona types see main text.
Female. Always with exserted part of ovipositor visible in dorsal view (Figs
For a detailed description of the genus see
Mesosoma. Pronotum in dorsal view appearing somewhat pentagonal or almost circular. Mesoscutellum sculpture variable dorsally, always with longitudinally strigose parts, with smoother, coriaceous apical rim distinguished by furrow or carina.
Wings. Postmarginal vein slightly shorter to slightly longer than marginal vein.
Metasoma. Length of ovipositor sheaths varying from less than 10% of metasoma length to 110% of metasoma length.
Note that some of the characters given by
Note: If combinations of characters are given, connected with and and separated by comma (not semicolon), then all characters have to be present; in the alternative path at least one of the characters has to be different.
1 | Ovipositor longer than 0.25× metasoma length; with bright blue or green colour on mesosoma (Fig. |
2 |
– | Ovipositor distinctly shorter than 0.25× metasoma length; mesosoma not brightly coloured | 3 |
2(1) | Ovipositor longer than metasoma; corona structure three- to four-part; body length >15 mm (Fig. |
O. gracilis Westwood (Indonesia) |
– | Ovipositor shorter than metasoma; corona structure square; body length <15 mm (Fig. |
O. longicollis (Cameron) (China, Indonesia, Malaysia, Myanmar, Philippines, Vietnam) |
3(1) | Head width about 7× eye distance (Fig. |
O. madegassa Bouček (Madagascar) |
– | Head width at most 4.38× eye distance; usually small- to medium-sized (≤9.0 mm) | 4 |
4(3) | Eyes small (eye height <0.6× head height), and head oval (head height 1.45–1.6× head length), and small-sized (<7.0 mm) | 5 |
– | Different combination of characters | 6 |
5(4) | Pronotum virtually round; profemur robust (profemur length <1.99× width) (Fig. |
O. circularicollis sp. n. (Morocco) |
– | Pronotum pentagonal, with broadest part before midlength; profemur medium (profemur length 2.15× width) (Fig. |
O. pumilae Yang (China) |
6(4) | Head round (height <1.45× length), and marginal vein long (marginal vein length >1.10× postmarginal vein length), and small-sized (<7.0 mm) | 7 |
– | Different combination of characters | 10 |
7(6) | Profemur elongated (profemur length >2.2× width); fore wing at least weakly infumate in part | 8 |
– | Profemur medium (profemur length 2.0–2.2× width); fore wing hyaline (Fig. |
O. florea sp. n. (Thailand) |
8(7) | Anterior margin of mesoscutellum convex (part anterior to imaginary transverse line connecting posterior margins of axillae more than 1/3 mesoscutellum length, 0.37–0.39); mesoscutellum completely lineate; pronotum pentagonal with posterior part hardly narrowing towards mesoscutum (Figs |
O. fidelis sp. n. (Vietnam) |
– | Anterior margin of mesoscutellum hardly convex (part anterior to imaginary transverse line connecting posterior margins of axillae at most 1/3 mesoscutellum length, 0.17–0.33); mesoscutellum lineate only in anterior two thirds; pronotum oval or pentagonal, if pentagonal then with posterior part distinctly narrowing towards mesoscutum | 9 |
9(8) | Pronotum oval (Fig. |
O. tenuicollis (Walker) (Indonesia) |
– | Pronotum pentagonal with posterior part distinctly narrowing towards mesoscutum (Fig. |
O. leibnizi sp. n. (Malaysia, Papua New Guinea, Philippines) |
10(6) | Body slender (mesonotum length 1.45× mesonotum width), and eyes large (eye height 0.64× head height), and marginal vein long (marginal vein length 1.17× postmarginal vein length) (Figs |
O. felix sp. n. (Central African Republic) |
– | Different combination of characters | 11 |
11(10) | Medium- to large-sized (≥7.0 mm), and either uniformly dark-coloured, or if not uniformly dark-coloured then fore wing hyaline | 12 |
– | Small-sized (<7.0 mm) or if medium-sized (7.0–9.0 mm) then in part with distinct dark green or coppery colour and fore wing partly infumate | 14 |
12(11) | Body robust or medium (mesonotum length ≤1.44× width) (Figs |
O. namibiensis sp. n. (Namibia) |
– | Body slender (mesonotum length >1.44× width) (Figs |
13 |
13(12) | Head and mesosoma uniformly black with tinges of dark green and purple (Figs |
O. mkomaziensis sp. n. (Tanzania) |
– | Head and mesosoma dark blue and green-blue (Figs |
O. ahoma (Mani & Kaul) (India) |
14(11) | Head round (head height <1.45× length) or if oval (head height 1.45–1.6× length) then either mesoscutellum only partially lineate and fore wing partly infumate (some O. formosa) or mesoscutellum only partially lineate and with distinct blue on head and mesosoma (O. srilankiensis males) | 15 |
– | Head oval or elongated (head height ≥1.45× length) and without other character combination | 16 |
15(14) | Propodeum large (propodeum length >0.15× mesoscutum length) (Fig. |
O. srilankiensis sp. n. (Sri Lanka) |
– | Propodeum usually medium (propodeum length 0.08–0.15× mesoscutum length) (Fig. |
O. formosa (Giraud) (Southern and Central Europe, Russia, eastern USA (introduced)) |
16(14) | Marginal vein short (marginal vein length <0.9× postmarginal vein length), and propodeum medium to large (propodeum length ≥0.08× mesoscutum length), and fore wing hyaline, and mesoscutellum at least partly areolate or rugolose | 17 |
– | Different combination of characters | 18 |
17(16) | Mesoscutellum densely lineate in anterior half to two-thirds and areolate in posterior one-third to half (Fig. |
O. niehuisorum sp. n. (Egypt, Israel) |
– | Mesoscutellum lineate in anterior two-thirds and rugulose in posterior one-third (Fig. |
O. hoggarensis Hedqvist (Algeria) |
18(16) | Marginal vein long (marginal vein length 1.19× postmarginal vein length); corona short (corona length 0.45× eye height) (Fig. |
O. regiae Yang (China) |
– | Marginal vein short to medium (marginal vein length <1.07× postmarginal vein length); corona longer (corona length always >0.52× eye height) | 19 |
19(18) | Fore wing hyaline (Fig. |
O. heikewernerae sp. n. (Botswana, South Africa) |
– | Fore wing partly infumate (Fig. |
O. magnifica (Risbec) (Senegal) |
Lycisca ahoma Mani & Kaul, 1973: 53–55.
Oodera
ahoma
;
FEMALE (N = 1). Medium-sized (8.40 mm). Head and mesosoma dark blue to green-blue. Fore wing hyaline. Body slender (mesonotum 1.51× as long as wide). Head round (1.41× as high as long). Eyes small (0.57× as high as head) (Fig.
FEMALE. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
Asia. India: female holotype, Lumding, Assam, leg. R.O., 25.05.1942, det. Mani and Kaul as Lycisca ahoma (
Hosts may be Agrilus sp. (Buprestidae) (
India.
FEMALE (N = 3). Small-sized (5.88–6.48 mm). Head and mesosoma blackish with dark green, purple and yellow parts. Fore wing partly infumate. Body robust to medium (mesonotum 1.22–1.43× as long as wide). Head oval (1.45–1.51× as high as long). Eyes small (0.54–0.56× as high as head) (Fig.
FEMALE. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
AFRICA. Morocco: female holotype, Granja del Muluya, Kebdana-Marruecos, leg. P. Alcaide VII. 1953, ex larva ‘unreadable word’ Punica, ex coll. V. Delucchi (
Unknown. The label data point towards an association with Punica (pomegranate).
Morocco.
Named for its unusual, round pronotum.
FEMALE (N = 1). Medium-sized (7.20 mm). Head and mesosoma mostly blackish and dark greenish. Fore wing partly infumate. Body slender (mesonotum 1.45× as long as wide). Head oval (1.59× as high as long). Eyes large (0.64× as high as head) (Fig.
FEMALE. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
AFRICA. Central African Republic: female holotype, Prefecture Sangha-Mbaéré, Parc Ntional de Dzanga-Ndoki, 38.6 km 173 °S Lidjombo, leg. S. van Noort, 26–27.05.2001 (
Unknown.
Central African Republic.
Named after the Latin adjective felix, meaning lucky.
Unfortunately, the holotype was damaged during examination for this study (metasoma detached from body). The images show the still undamaged specimen.
BOTH SEXES (N = 3). Small-sized (6.48–6.80 mm). Head and mesosoma dark green to blue. Fore wing partly infumate. Body robust to slender (mesonotum 1.29–1.47× as long as wide). Head round (1.33–1.41× as high as long). Eyes large (0.61–0.67× as high as head) (Fig.
FEMALE. Metasoma medium (0.45–0.46× as long as body). Ovipositor short (0.13–0.13× as long as metasoma) (Fig.
BOTH SEXES. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
ASIA. Vietnam: female holotype, Ha Tinh Huong son, 900 m, 18°22'N/105°13'E, leg. L. Herman, MT, 05.05.1998 (
Unknown.
Vietnam.
Named after the latin adjective fidelis, meaning faithful, because of the distinct blueish colour of head and mesosoma and blue being the symbolic colour of trust.
FEMALE (N = 1). Small-sized (4.80 mm). Head and mesosoma dark green and green-blue. Fore wing hyaline. Body robust (mesonotum 1.32× as long as wide). Head round (1.41× as high as long). Eyes large (0.66× as high as head) (Fig.
FEMALE. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
ASIA. Thailand: female holotype, Loei, Phu Kradueng NP, labelled “Deciduous”, leg. S. Glong-lasae, 02–10.10.2006 (
Unknown.
Thailand.
Named after the latin adjective florea, meaning flowery. The species is, in our subjective view, particularly beautiful and flimsy, like a flower.
Heydenia formosa Giraud, 1863: 21–22.
Stellophora
formosa
;
Oodera
formosa
;
Oodera
bestia
Nikol’skaya, 1952: 487–488. Synonymy by
Oodera
monstrum
Nikol’skaya, 1952: 487–488, syn. n. (Figs
BOTH SEXES (N = 35). Usually small-sized (3.60–7.12 mm, with only 1 of 27 medium-sized). Head and mesosoma dark green and coppery. Fore wing partly infumate. Body robust to slender (mesonotum 1.19–1.47× as long as wide). Head usually round (1.30–1.48× as high as long, with only 3 of 32 with head oval). Eyes usually large (0.55–0.68× as high as head, with only 4 of 32 with eyes small) (Fig.
FEMALE. Metasoma short to long (0.43–0.55× as long as body). Ovipositor usually short (0.09–0.17× as long as metasoma, with only 2 of 26 with metasoma rather long) (Fig.
BOTH SEXES. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
Europe. Bulgaria: male, Slencev Brjag, leg. Kocourek, 26.07.1968, det. Z. Bouček 1976 (
Hosts: Buprestidae (Agrilus sp., A. graminis, A. suvorovi, A. viridis, Capnodis sp.), Cleridae (Tillus unifaciatus), Ptinidae (Ptinus germanus), Scolytinae (Hylesinus sp.); Plant associates: Fabaceae (Robinia sp.). The host and associates records are taken from
Southern and Central Europe, northernmost location in Germany, Rhineland-Palatinate, representing first record from Germany; Russia. Introduced to the eastern United States (Kentucky, Virginia, New Jersey) and recently found in eastern Canada (Ontario).
Comparison of the female holotype (examined from images) and the male paratype of O. monstrum (Figs
The specimens from North America were assumed to be O. formosa by
In general, we found distinguishing O. formosa from many other species of Oodera to be rather difficult. This is mainly due to the fact that O. formosa is the only species of Oodera represented by a significant number of specimens from several series (except for O. longicollis (Cameron) which is, however, very easily distinguished from all other Oodera species). The specimens show intraspecific variation that can be quite staggering for some characters. Only examination of this larger series allowed us to recognise the variation as intraspecific. For many other species, only single or a few specimens were available or (small) uniform series that originate from the same host or region. The variation of O. formosa is reflected in several diagnostic characters for which we add the term “usually” if the vast majority of examined specimens exhibits this character but a minority does not. We made these additions (also in the diagnoses of few other species) to make diagnoses and key more easily applicable for the reader. With a combination of characters, O. formosa is well separated from all other species.
Oodera gracilis Westwood, 1874: 145.
FEMALE (N = 1). Large-sized (17.00 mm). Head and mesosoma bright dark blue and green-blue with blackish parts. Fore wing partly infumate. Body slender (mesonotum 1.45× as long as wide). Head elongated (1.67× as high as long). Eyes large (0.60× as high as head) (Fig.
Habitus (dorsal) II a O. gracilis Westwood (imaged specimen: OGr01; ovipositor not visible in full length) b O. heikewernerae sp. n. (OHe01, holotype) c O. hoggarensis Hedqvist (OHo02, paratype) d O. leibnizi sp. n. (OLe01, holotype) e O. longicollis (Cameron) (OLo01) f O. madegassa Bouček (OMad01, holotype). Scale bars: 1 mm.
FEMALE. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
ASIA. Indonesia (North Sulawesi): female, Indonesia, Sulawesi Utara, Dumoga-Bone N.P., leg. G. Else, 09.06.1985, det. Z. Bouček (compared with holotype in Oxford Museum) (
Unknown.
Indonesia (Sulawesi), West Papua.
We did not examine the type (described from West Papua). However, the uniqueness of the species and the fact that we examined a specimen compared to the type by Bouček allow us to include this species in the taxonomic revision.
BOTH SEXES (N = 12). Head and mesosoma mostly blackish with blue and coppery parts. Fore wing hyaline. Body robust to medium (mesonotum 1.22–1.40× as long as wide). Head usually oval (1.45–1.65× as high as long, with only 3 of 12 with head elongated). Eyes small to large (0.58–0.67× as high as head) (Fig.
FEMALE. Small-sized (5.31–6.64 mm). Profemur robust to medium (1.71–2.04× as long as wide). Metasoma short to medium (0.39–0.47× as long as body). Ovipositor short to rather long (0.12–0.18× as long as metasoma) (Fig.
MALE. Body length not available. Profemur elongated (2.29× as long as wide).
BOTH SEXES. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
AFRICA. Botswana: female holotype, Serowe, Farmer’s Brigade, leg. P. Forchhammer, MT, XII. 1987 (
Unknown.
Botswana and South Africa.
Named in memory of Heike Werner, the mother of the first author, who sadly passed away during completion of this revision.
Oodera hoggarensis Hedqvist, 1967: 186
FEMALE (N = 2). Exact body length not available (both specimens are fragmented), roughly small-sized. Head and mesosoma blackish with light green, purple and yellow parts. Fore wing hyaline. Body robust (mesonotum 1.16–1.2× as long as wide). Head oval (1.52–1.54× as high as long). Eyes small to large (0.59–0.64× as high as head) (Fig.
FEMALE. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
AFRICA. Algeria: female holotype, Timesdelssine, Hoggar, leg. J. Mateu, 23.05.1962, det. K-J. Hedqvist (1966) (
Algeria (Hoggar Mountains).
The species description of O. hoggarensis by
BOTH SEXES (N = 4). Small-sized (5.31–6.06 mm). Head and mesosoma dark green to blue-green. Fore wing partly weakly infumate. Body robust to medium (mesonotum 1.34–1.42× as long as wide). Head round (1.29–1.42× as high as long). Eyes large (0.60–0.66× as high as head) (Fig.
FEMALE. Corona thick (3.64–4.00× as long as wide). Metasoma short (0.45–0.45× as long as body). Ovipositor short (0.08–0.10× as long as metasoma) (Fig.
MALE. Corona medium (4.36× as long as wide).
BOTH SEXES. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
ASIA. Philippines: female holotype, S.O. Luzon, determined as O. ornata by anonymous (
Unknown.
Malaysia, Philippines (both Oriental part of Asia), Papua New Guinea (Australasian part of Asia).
Named in honour of Gottfried Wilhelm Leibniz (1646–1716) on the occasion of his 300th death day. The Zoologisches Forschungsmuseum Alexander Koenig in which this study was done is part of the Leibniz Association, named after Leibniz.
The two specimens from Malaysia and the Philippines were originally identified as O. ornata (valid species name O. longicollis), one of the specimens from Papua New Guinea was identified by Bouček as O. gracilis in 1984, later crossed out by an unknown person. The new species is rather easily distinguished from O. longicollis and O. gracilis, as O. longicollis and O. gracilis exhibit exceptionally long ovipositors and bright metallic colours on head and mesosoma while O. leibnizi sp. n. does not.
Epistenia longicollis Cameron, 1903: 98.
Oodera
longicollis
;
Oodera
ornata
Gahan, 1925: 97. Synonymy with Oodera longicollis and lectotype designation by
BOTH SEXES (N = 14). Head and mesosoma bright green to blue and blackish. Fore wing partly infumate. Body robust to slender (mesonotum 1.28–1.45× as long as wide). Head oval to elongated (1.57–1.72× as high as long). Eyes large (0.60–0.65× as high as head) (Fig.
FEMALE. Medium- to large-sized (8.80–12.60 mm). Propodeum medium to large (0.12–0.22× as long as mesoscutum) (Fig.
MALE. Small- to medium-sized (5.38–9.6 mm). Propodeum very large (0.26 to 0.34× as long as mesoscutum).
BOTH SEXES. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
ASIA. China: female, Matang Rd, 18.05.1920, det. C. R. Vardy (1962) (
Associated with teak (Tectona grandis) (Verbenaceae), presumably as habitat for the parasitoids’ host (unverified record taken from
China (Oriental part), Malaysia, Myanmar, Philippines, Vietnam.
Oodera ornata was synonymised with O. longicollis by
The species description of O. ornata by
Note that we located one additional specimen in the Australian National Insect Collection in Canberra (ANIC). It originated from the same series as specimens already included here (from Philippines, Davao/Mindanao). O. longicollis is a well-defined species and we did not borrow and include this additional specimen.
Oodera madegassa Bouček, 1958: 376.
FEMALE (N = 1). Large-sized (10.30 mm). Head and mesosoma dark blue to blackish. Fore wing partly infumate. Body slender (mesonotum 1.52× as long as wide). Head elongated (1.77× as high as long). Eyes large (0.63× as high as head) (Fig.
FEMALE. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
AFRICA. Madagascar: female holotype, Haute-Vallée de Sambirano, det. Z.
Unknown.
Madagascar.
Stellophora magnifica Risbec, 1951: 239–243.
Oodera
magnifica
;
FEMALE (N = 1). Small-sized (5.75 mm). Head and mesosoma blackish, coppery and reddish. Fore wing partly infumate. Body robust (mesonotum 1.25× as long as wide). Head oval (1.51× as high as long). Eyes large (0.65× as high as head) (Fig.
FEMALE. Colour (Figs
Habitus (dorsal) III a O. magnifica (Risbec) (imaged specimen: OMag01, holotype) b O. mkomaziensis sp. n. (OMk01, holotype) c O. monstrum syn. n. (OFo12, paratype) d O. namibiensis sp. n. (ONa01, holotype) e O. niehuisorum sp. n. (ONi01, holotype) f O. pumilae Yang (OPu01, paratype) g O. regiae Yang (ORe01, paratype) h O. srilankiensis sp. n. (OSr01, holotype) i O. tenuicollis (Walker) (OTe01, holotype). Scale bars: 1 mm.
Head (frontal) III a O. magnifica (Risbec) (imaged specimen: OMag01, holotype) b O. mkomaziensis sp. n. (OMk01, holotype) c O. monstrum syn. n. (OFo12, paratype) d O. namibiensis sp. n. (ONa01, holotype) e O. niehuisorum sp. n. (ONi01, holotype) f O. pumilae Yang (OPu01, paratype) g O. regiae Yang (ORe01, paratype) h O. srilankiensis sp. n. (OSr01, holotype) i O. tenuicollis (Walker) (OTe01, holotype). Scale bars: 1 mm.
Head (lateral) III a O. magnifica (Risbec) (imaged specimen: OMag01, holotype) b O. mkomaziensis sp. n. (OMk01, holotype) c O. monstrum syn. n. (OFo12, paratype) d O. namibiensis sp. n. (ONa01, holotype) e O. niehuisorum sp. n. (ONi01, holotype) f O. pumilae Yang (OPu01, paratype) g O. regiae Yang (ORe01, paratype) h O. srilankiensis sp. n. (OSr01, holotype) i O. tenuicollis (Walker) (OTe01, holotype). Scale bars: 1 mm.
Mesosoma (dorsal) II a O. gracilis Westwood (imaged specimen: OGr01) b O. heikewernerae sp. n. (OHe01, holotype) c O. hoggarensis Hedqvist (OHo02, paratype) d O. leibnizi sp. n. (OLe01, holotype) e O. longicollis (Cameron) (OLo01) f O. madegassa Bouček (OMad01, holotype). Scale bars: 1 mm.
Mesosoma (dorsal) III a O. magnifica (Risbec) (imaged specimen: OMag01, holotype) b O. mkomaziensis sp. n. (OMk01, holotype) c O. monstrum syn. n. (OFo12, paratype) d O. namibiensis sp. n. (ONa01, holotype) e O. niehuisorum sp. n. (ONi01, holotype) f O. pumilae Yang (OPu01, paratype) g O. regiae Yang (ORe01, paratype) h O. srilankiensis sp. n. (OSr01, holotype) i O. tenuicollis (Walker) (OTe01, holotype). Scale bars: 1 mm.
Mesoscutellum and propodeum (dorsal) I a O. ahoma (Mani & Kaul) (imaged specimen: OAh01, holotype) b O. circularicollis sp. n. (OCi01, holotype) c O. felix sp. n. (OFe01, holotype) d O. fidelis sp. n. (OFi01, holotype) e O. florea sp. n. (OFl01, holotype), f O. formosa (Giraud) (OFo14). Scale bars: 1 mm.
Mesoscutellum and propodeum (dorsal) II a O. gracilis Westwood (imaged specimen: OGr01) b O. heikewernerae sp. n. (OHe01, holotype) c O. hoggarensis Hedqvist (OHo02, paratype) d O. leibnizi sp. n. (OLe03, paratype) e O. longicollis (Cameron) (OLo01) f O. madegassa Bouček (OMad01, holotype). Scale bars: 1 mm.
Mesoscutellum and propodeum (dorsal) III a O. magnifica (Risbec) (imaged specimen: OMag01, holotype) b O. mkomaziensis sp. n. (OMk01, holotype) c O. monstrum syn. n. (OFo12, paratype) d O. namibiensis sp. n. (ONa01, holotype) e O. niehuisorum sp. n. (ONi01, holotype) f O. pumilae Yang (OPu01, paratype) g O. regiae Yang (ORe01, paratype) h O. srilankiensis sp. n. (OSr01, holotype) i O. tenuicollis (Walker) (OTe01, holotype). Scale bars: 1 mm.
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
AFRICA. Senegal: female holotype, M’Bambey, leg. A. Wane, 13.11.1945, det. J.Y. Rasplus (1990) (
Unknown.
Senegal.
The description of O. magnifica by
FEMALE (N = 1). Medium-sized (7.20 mm). Head and mesosoma black, with tinges of dark green and purple. Fore wing hyaline. Body slender (mesonotum 1.50× as long as wide). Head oval (1.56× as high as long). Eyes large (0.63× as high as head) (Fig.
FEMALE. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
AFRICA. Tanzania: female, Mkomazi Game Reserve (now Mkomazi National Park), Ibaya Hill, 3°58.40'S 37°47.13'E, leg. S. van Noort, 15.–30.04.1996 (SAM) (OMk01)
Unknown.
Tanzania.
Named after the type locality, the Mkomazi National Park in northeastern Tanzania.
FEMALE (N = 8). Medium- to large-sized (7.36–9.10 mm). Head and mesosoma dark, mostly blackish and reddish. Fore wing partly infumate. Body robust to medium (mesonotum 1.01–1.40× as long as wide). Head round to oval (1.42–1.59× as high as long). Eyes large (0.6–0.66× as high as head) (Fig.
FEMALE. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
AFRICA. Namibia: female holotype, ca. 1500 m (above sea level), ca. 140 km N Okanhandja, 20°50.85'S/16°47.77'E, Holzeintrag (= from wood), leg. M. & O. Niehuis, 01.04.1997 (
Reared from wood, exact tree species unknown.
Namibia.
Named after the type series’ origin from Namibia.
BOTH SEXES (N = 13). Small-sized (4.50–6.48 mm). Head and mesosoma blackish and coppery, with small dark green parts, never with blue. Fore wing hyaline. Head usually oval (1.42–1.59× as high as long, with only 2 of 13 with head round). Eyes large (0.60–0.74× as high as head) (Fig.
FEMALE. Body robust to medium (mesonotum 1.29–1.41× as long as wide). Metasoma short (0.40–0.45× as long as body). Ovipositor short to rather long (0.14–0.18× as long as metasoma) (Fig.
MALE. Body robust to slender (mesonotum 1.27–1.47× as long as wide).
BOTH SEXES. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
AFRICA. Egypt: female holotype, Sinai, Nuweiba, ex Acacia sp. tree, 29°01.17'N/34°40.52'E, leg. O. Niehuis, 11.03.2001, 10 m amsl (= above sea level) (
Reared from Acacia wood, which is likely the habitat of the O. niehuisorum host.
Egypt and Israel.
Named after Manfred and Oliver Niehuis, who collected the specimens of the type series.
Oodera pumilae Yang, 1996: 100, 311.
FEMALE (N = 1). Small-sized (5.81 mm). Head and mesosoma dark green to blue-green. Fore wing partly weakly infumate. Body robust (mesonotum 1.25× as long as wide). Head oval (1.46× as high as long). Eyes small (0.59× as high as head) (Fig.
FEMALE. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
ASIA. China: female paratype, leg. Z.Q. Yang, 02.07.1989, rest of label in Chinese, det. Z.Q. Yang 1995 (
Probably parasitic on Scolytus spp. on elm (Ulmus) trees (
China (Province: Heilongjiang; Palaearctic part of China) (
We were unable to examine the holotype. Given that the paratype we examined is from the same series as the holotype and that it perfectly matches the (short) description, we decided to include the species in our revision, diagnoses and key without examination of the primary type.
Oodera regiae Yang, 1996: 98, 310.
FEMALE (N = 1). Small-sized (6.25 mm). Head and mesosoma dark green to green. Fore wing hyaline. Body robust (mesonotum 1.25× as long as wide). Head oval (1.58× as high as long). Eyes large (0.69× as high as head) (Fig.
FEMALE. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
ASIA. China: female paratype Oodera regiae, 12.1994, det. Z.Q. Yang 1995, rest of labels in Chinese (
Parasitic on the larvae of Xyeloborus sp. (Curculionidae) and Agrilus sp. (Buprestidae) on a walnut (Juglans regiae) tree (Yang, 1996).
China (Province: Shaanxi; Palaearctic part of China) (
The species is characterised by a very short corona (measured as corona length to eye height). In all other species the corona is distinctly longer, with no diagnostically useful differences among them.
We were unable to examine the holotype. Given that the paratype we examined is from the same series as the holotype and that it perfectly matches the (short) description, we decided to include the species in our revision, diagnoses and key without examination of the primary type.
The description of O. regiae Yang, 1996 lists four characters to distinguish this species from O. formosa. First, O. regiae does not have dense hairs on the basal tergite (interpreted as the first gastral tergite), arranged circularly, while O. formosa does. This was not confirmed. The distribution of gastral hairs is identical between the examined O. regiae and specimens of O. formosa. Second, O. regiae is described to have “eyes having sparse pubescence“ (
BOTH SEXES (N = 5). Small-sized (4.00–5.75 mm). Head and mesosoma dark blue to blue-green. Fore wing hyaline. Body robust (mesonotum 1.26–1.33× as long as wide). Eyes large (0.60–0.67× as high as head) (Fig.
FEMALE. Head round (1.30–1.41× as high as long). Metasoma medium (0.47× as long as body). Ovipositor short (0.13–0.14× as long as metasoma) (Fig.
MALE. Head oval (1.53–1.54× as high as long).
BOTH SEXES. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
ASIA. Sri Lanka: female holotype, Mate. Dist. Kibissa, 0,5 mi West of Sigiriya, Jungle, leg. K.V. Krombein, 28.06–04.07.1978 (
Unknown.
Sri Lanka.
Named after the geographic origin of the type series from Sri Lanka.
Some (or maybe all) specimens described here as O. srilankiensis were probably those examined and mentioned by
Eupelmus tenuicollis Walker, 1872: 86.
Oodera
tenuicollis
;
FEMALE (N = 1). Exact body length not available (the only specimen available is missing metasoma), if metasoma is not uniquely long, then small-sized. Head and mesosoma dark blue to green-blue. Fore wing partly infumate. Body slender (mesonotum 1.45× as long as wide). Head round (1.40× as high as long). Eyes large (0.67× as high as head) (Fig.
FEMALE. Colour (Figs
Head (Figs
Mesosoma (Figs
Wings (Fig.
MALE. Unknown.
ASIA. Indonesia: female type (probably holotype), Eupelmus tenuicollis Walker Notes on Chalcidide 5:86.1872, Type female checked RDE 1955 (1955), det. Oodera sp. by R. D. Eady,
Unknown.
Indonesia.
This species is a bit problematic, because only the single type specimen without metasoma is available. However, the rest of the body provides some good diagnostic characters that allow differentiation from all other species of Oodera.
Oodera obscura Westwood, 1874: 146.
Maybe Indonesia. This information is given by
Oodera rufimana Westwood, 1874: 146.
Cambodia.
As for O. obscura (see above), we expected the holotype to be in Oxford as part of the W. W. Saunders collection (both species were described in the same publication). However, even after extensive search at Oxford and London (where other parts of the Saunders collection are located) by the respective curators the type could not be located. The species description is not conclusive and specific. We therefore considered it as a nomen dubium.
Oodera albo-pilosa Crosby, 1909: 86.
This new combination is by authority of G. Delvare (CSIRO, Montpellier), J. Werner and R. S. Peters.
Female holotype, labelled “Eupelmidae?” by
Parasite of an unknown gall-forming fly on branches of Combretum olivaceum (Crosby, 1909).
The species is listed from South Africa and Zambia in
We only examined images of the holotype. However, the female type is clearly a specimen belonging to the subfamily Eupelminae (Eupelmidae), not an Oodera. The species is therefore removed from Oodera and transferred to Eupelmus. The tentative assignment to Eupelmidae was already done by
Five specimens could not be assigned to any of the previously or newly described species (specimens OSp01 (
AFRICA. Namibia: female, Waterberg ca. 1500 m (above sea level), 20°36.58'S/17°10.43'E, Holzeintrag (= from wood), ex larva, leg. M. & O. Niehuis, 01.04.1997 (
This specimen has the same collecting data as one of the paratypes of O. namibiensis sp. n. and was at first included as paratype of this species. Yet, it does not belong to this species because it is, for example, not uniformly dark-coloured on head and mesosoma but has some distinct dark green parts, and the fore wing is not partly infumate but hyaline.
ASIA. Indonesia: male, Sumatra: Tep. Tingi, standing over: Oodera ornata in Hedqvist coll.
It is a single male specimen. It was collected in Indonesia, yet very distant from the only other species from Indonesia O. tenuicollis. It was standing as Oodera ornata (now O. longicollis) in the Hedqvist collection at the
AFRICA. Republic of South Africa: female, Western Cape, Gamkaberg Nature Reserve, 33°39.941'S/21°53.505'E, 315 m, 21.03.–05.05.2009, leg. S. van Noort (
This specimen has very few conspicuous characteristics that might allow placing it close to any other species. It differs from the other South African species O. heikewernerae sp. n. by, for example, a distinctly larger body length (7.7 mm compared to maximum of 6.64 mm in O. heikewernerae), a slightly different corona (no horizontal crests connected by vertical crests vs. some horizontal crests connected by vertical crests in O. heikewernerae), and a different scutellum sculpture (only partially lineate vs. completely lineate in O. heikewernerae). It is tentatively close to O. felix (from Central African Republic), but differs also from this species by, for example, hyaline fore wings (partly infumate in O. felix), small eyes (large in O. felix), and a partly blueish mesosoma (dark green in O. felix).
AFRICA. Tanzania: female. Mkomazi Game Reserve (now Mkomazi National Park), Kisima Plot, 4°06.06'S/38°05.58'E, Acacia/Commiphora bushland, 25.11.–08.12.1995, leg. S. van Noort (
The specimen was collected in the same National park in Tanzania as O. mkomaziensis sp. n. but is different from this species by, for example, distinct light green colour on the head (dark in O. mkomaziensis), partly infumate fore wings (hyaline in O. mkomaziensis), and being distinctly smaller (this specimen: 5.56 mm, O. mkomaziensis: 7.20 mm).
AFRICA. Republic of South Africa: female, Merweville, Laingsburg Distr., leg. H. Zinn (
The specimen is not too different from the other South African species O. heikewernerae sp. n., yet cannot be assigned to this species because of a few rather subtle differences, for example, the colour of the head and mesosoma (light green on head and reddish parts on mesosoma, both not present in O. heikewernerae), and the head shape (round in this specimen, usually oval in O. heikewernerae).
This taxonomic revision of Oodera is based on a comparatively small number of specimens (115) and on morphological data only. Several of the new and previously described species are known from only a single specimen. This is inevitable because of the rarity of specimens in scientific collections. The samples we used were borrowed from 13 different museums, but more museums in Germany and abroad (e.g., Bavarian State Collection for Zoology Munich, Senckenberg Museum Frankfurt, Senckenberg Deutsches Entomologisches Institut Müncheberg, University of Oxford Hope Collection, National Museum of Natural History Stuttgart, Zoological Museum of the University of Zurich, Zoologisches Museum Hamburg, Australian National Insect Collection) were investigated for specimens of Oodera but do not have any (except for one specimen of O. longicollis in Canberra, see part on O. longicollis). Apart from the specimens gathered for this revision that are deposited at the
Species of Oodera seemingly prefer warmer to temperate regions (Fig.
Geographic distribution of the genus Oodera. Countries from which Oodera was recorded are highlighted in dark grey. Stars indicate record localities. Record localities of O. pumilae Yang and O. regiae Yang are not exact but placed in the centre of the respective Chinese province from which the species was recorded.
After comprehensive examination of all available material of this genus, we would like to add some speculation on the function of the modified front legs that are present in both sexes in striking conspiciousness. Raptorial function is highly unlikely. Although Oodera is an extraordinary chalcidoid in many ways, its mouth parts, head shape, etc. are typical for Chalcidoidea, and no other chalcidoid has been reported to be raptorial and carnivorous in the adult stage. Also, the anecdotal descriptions of
We hope that this first revision of the group and the included identification key will foster studies on this beautiful and intriguing group of chalcids that will eventually lead to at least some basic knowledge about the species’ biology and evolution.
Thanks to Gary Gibson (
Table S1. Measurements (in mm) and ratios for all specimens examined