Corresponding author: A. A.Valerio (
Academic editor: Gavin Broad
The genera
The genera
The taxonomic status of both genera has largely been stable since their description, in part because they have received little attention.
Representation of theexternal shape of T1 horn (T1H), shape of the ovipositor shaft (OS) and invaginated intersegmental membrane (IM) for
The current study builds on our recent taxonomic treatment of
The contributions of the individual authors are as follows; A.A. Valerio: character definition, character development, character coding, imaging, species concept development; key development, capture of specimen data, phylogenetic analysis and manuscript preparation; A.D. Austin: species concept development, manuscript preparation, key development, and character coding; L. Masner: generic characters, character definition, character development, and specimen acquisition; N.F. Johnson: generic characters, distributional information and manuscript preparation.
The specimen localities, except for the holotypes, are not included in full in the article’s text to avoid repetition of the data and to save publication space. The full collecting data for specimens are available at
Appendix I list terms associated with identifiers in the
A total of 41 morphological characters were scored for the 55 ingroup and four outgroup taxa (see Appendices II and III). Only character 9 (shape of the medial area of the vertex) was uninformative under the parsimony criterion. Characters 37 (ovipositor shape) and 41 (number of antennal segments) where coded directly from specimens for some species, and from the literature for others (e.g.
Maximum parsimony analysis resulted in 10,000 trees of 348 steps (CI=0.18, RI=0.58). The strict consensus tree is largely unresolved except for a monophyletic group containing all 13
50% majority rule consensus tree (with frequencies above each node) for species of
Phylogenetic consensus tree under Implied Weighting scheme for species of
In both trees the same group of Australian species is sister to
Additionally, the topology of the IW trees for K=5 and K=7 were similar to that for K=3, with
Although the relationships inferred from the data are not particularly robust, it is difficult to entertain a scenario in which all
Distributional map of Old World
Like other members of the
Although nothing is known about the ovipositional behavior of
The previously described species
1 | Fore wing with tubular stigmal vein present, without dark infuscate patch or bands at this position ( |
2 |
– | Fore wing with stigmal vein always absent, with dark infuscate patch or band at this position ( |
|
2 | Notauli completely absent ( |
3 |
– | Notauli always present ( |
6 |
3 | Mesoscutum and mesoscutellum conspicuously flattened in lateral view ( |
|
– | Mesoscutum and mesoscutellum convex in lateral view ( |
4 |
4 | Mesoscutum and mesoscutellum rugulose-granulate ( |
|
– | Mesoscutum and mesoscutellum granulate, never rugulose-granulate ( |
5 |
5 | Posterior sublateral areas of mesoscutum with cicatrose sculpture, remainder of mesoscutum with dense, weak granulate sculpture ( |
|
– | Mesoscutum covered with fine, weak granulate sculpture ( |
|
6 | Netrion appearing absent, lateral margins of pronotum with extensive, conspicuous, longitudinal costae ( |
7 |
– | Netrion well-defined, lateral margins of pronotum at most with fine costae around margin of netrion ( |
12 |
7 | Gena mostly smooth, shining, sometimes coriaceous in upper 1/3 ( |
|
– | Gena weakly rugulose ( |
8 |
8 | Notauli present as smooth furrows ( |
|
– | Notauli crenulate throughout length ( |
9 |
9 | Frons and mesoscutum rugulose, rugulae broad ( |
|
– | Frons smooth, mesoscutum granulose, never rugulose; notauli variable in width, crenulae present or not; body color variable | 10 |
10 | Body black or dark brown; mesoscutellum smooth medially, setation conspicuously long and somewhat sparse ( |
|
– | Body yellow or brown; mesoscutellum without smooth areas, setation short and dense; mesoscutum completely sculptured; crenulae of notauli weak or broad ( |
11 |
11 | Body brown ( |
|
– | Body completely yellow ( |
|
12 | Gena smooth ventrally ( |
|
– | Gena completely sculptured ventrally; lateral propodeal areas variable in sculpture | 13 |
13 | Compound eyes unusually small, dorsoventral length less than 1/3 head height ( |
|
– | Compound eyes of normal size, dorsoventral length nearly 1/2 head height ( |
14 |
14 | Occipital and orbital carinae contiguous or separated at most by distance subequal to width of occipital carina; head in lateral view elongate; occipital carina well-defined laterally ( |
15 |
– | Occipital carina and orbital carinae separated by distance greater than width of occipital carina ( |
18 |
15 | Mesoscutellum medially depressed posteriorly ( |
|
– | Mesoscutellum convex; mesoscutum weakly granulate ( |
16 |
16 | S2 anterior carina absent; lagrimal absent; metasoma yellow or pale brown (except T1 horn) contrasting with dark brown color of remainder of body (Madagascar) | |
– | S2 anterior carina present; lagrimal present ( |
17 |
17 | Ventral portion of antennal scrobe with fan-like carinae arising from malar space ( |
|
– | Ventral portion of antennal scrobe without fan-like carinae ( |
|
18 | Central keel of frons absent ( |
19 |
– | Central keel of frons present ( |
20 |
19 | S2 anterior carina interrupted medially (similar to |
|
– | S2 anterior carina uninterrupted ( |
|
20 | Notauli crenulate ( |
21 |
– | Notauli smooth ( |
22 |
21 | Ocelli large ( |
|
– | Ocelli small ( |
|
22 | Vertex slightly depressed mesally ( |
|
– | Vertex flat or slightly rounded, without mesal depression; antennal scrobe varying from completely smooth to completely rugulose; lateral ocelli variable in size | 23 |
23 | Antennal scrobe with transverse costae, covering at least 1/2 of frons ( |
24 |
– | Antennal scrobe without such extensive transverse elements, mostly smooth or only with coriaceous or granulose sculpture ( |
29 |
24 | Fan-like costae present on frons, with a well-developed furrow at lower lateral areas of scrobe ( |
|
– | Fan-like sculpture absent on frons, if weakly indicated, then without a furrow at lower lateral areas of scrobe ( |
25 |
25 | Vertex in anterior view flat ( |
|
– | Vertex in anterior view convex; lateral surface of T1 horn transversely carinate, carinae becoming more curved dorsally, posterior surface with longitudinal carinae dorsally and remainder of T1 horn smooth or with very few narrow transverse costae on upper 1/3; T1 horn thin, elongate | 26 |
26 | Lateral ocelli small ( |
|
– | Lateral ocelli large, distance between occipital carina and lateral ocellus less than 0.8× diameter of lateral ocellus | 27 |
27 | S2 anterior carina absent ( |
|
– | S2 anterior carina present (similar to |
28 |
28 | Body yellow ( |
|
– | Body dark brown ( |
|
29 | S2 anterior carina interrupted medially ( |
30 |
– | S2 anterior carina present, continuous ( |
32 |
30 | S2 anterior carina absent ( |
|
– | S2 anterior carina present, interrupted medially ( |
31 |
31 | Body dark brown; T1 laterally with abundant, fine, longitudinal costae, costae slightly curved, effaced (Congo, Guinea, Kenya, Madagascar, South Africa, Uganda, Zimbabwe) | |
– | Body completely yellow; T1 laterally with sparse, coarser, longitudinal costae, costae very straight, well-defined (Cameroon) | |
32 | Mesosoma orange to light brown, contrasting with dark brown head and metasoma ( |
|
– | Body uniformly colored from yellow to dark brown ( |
33 |
33 | Mesoscutum between notaulus and tegula almost completely smooth ( |
|
– | Mesoscutum and mesoscutellum covered by well-defined sculpture, without smooth areas | 34 |
34 | Lagrimal present, conspicuous ( |
35 |
– | Lagrimal absent or reduced ( |
36 |
35 | Head yellow; T1 light orange with horn contrastingly dark brown ( |
|
– | Head dark brown; T1 entirely yellow ( |
|
36 | Body completely black ( |
|
– | Body color a combination of yellow and dark brown, or completely yellow ( |
This species is named in reference to its broad distribution across southern and east Africa.
Holotype female:
The holotype has the posterior right leg detached from the body and glued to the triangle; the right side wings are detached and in a gelatin capsule; the left antenna is missing. Most of the paratypes are in perfect condition.
In some female specimens the color pattern differs from that described above, the metasoma is a slightly lighter tone of dark brown compared to the head or the mesosoma (i.e. OSUC 238763). Additionally, the mesal area of T2 normally is covered by fine, dense, longitudinal costae (as in the holotype) but some specimens exhibit a smooth area with weak coriaceous sculpture in the background (in the absence of longitudinal costae), or a narrow smooth area with the lateral areas of the tergite with weak, longitudinal costae mixed with weak coriaceous sculpture in the background.
This is the only species of
This species is named to honor the Renaissance Italian naturalist Ulisse Aldrovandi. The epithet is used as noun in the genitive case.
Holotype female:
The holotype is in good condition except that the right hind wing and right antennal clava are missing.
The species
This species is named after the great British fossil hunter Mary Anning. The epithet is a noun in the genitive case.
Holotype female:
The female holotype is in perfect condition.
This is one of two species within a group of taxa in which the netrion is obscured and its marginal crenulae are not well-defined, the mesoscutum is completely sculptured, the mesoscutellum is never with smooth areas, and has short, dense setation.
This species is named after the collection locality of the species, Australia. The epithet is used as an adjective.
On tree trunk of
Holotype female:
The holotype is in perfect condition; all paratypes are in good condition except for specimen OSUC 239143 which is missing the metasoma.
This species is very similar to
This species is named after the amazing German biologist Karl Ernst von Baer. The epithet is a noun in the genitive case.
Holotype female:
The female holotype is in perfect condition.
This species may be distinguished from all others with an obscured netrion by the combination of the very wide notauli that have a few, broad and well-defined crenulae; the mesoscutum broadly smooth in its posterior half; and the mesoscutellum mesally with a smooth patch and its conspicuously long but somewhat sparse setae.
This species is named after our entomological colleague Dr Jo Berry from New Zealand. The epithet is a noun in the genitive case.
Recorded as an egg parasitoid of
Holotype female:
The holotype is in perfect condition as are the remainder of the paratypes except for specimen OSUC 238564 which is covered with dust.
This species is very similar to
This species is name after the amazing Bengali painter Nandalal Bose, recipient of the “Padma Vibhushan”. The epithet is a noun in the genitive case.
Holotype female:
The holotype is in perfect condition as are most of the paratypes except for specimen OSUC 238789 which has the left fore wing glued to the point, and OSUC 238790 which has the metascutellum detached from the body.
The species
This species is named after Josephine Cardale, former collection manager of
Holotype female:
The holotype is in perfect condition.
Sculpture of antennal scrobe: largely smooth ventrally, dorsally with sinuate, transverse ridges. Surface of torular triangle: flat. Development of central keel on frons: present, elongate (equal to or greater than 1/3× height of frons), but not reaching anterior ocellus; present, elongate, reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: with sparse, transverse costae mixed with weak, dense granulae. Sculpture of malar space: with sparse, fine, dorsoventral carinae. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: large. Distance between lateral ocellus and occipital carina: 0.5–1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: granulate. Sculpture of occipital carina: weakly crenulate throughout. Distance from occipital carina to orbital carina: at least 2× width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weak, small granulae. Sculpture of gena: granulose.
This species is easily identified based on its unique color pattern; there is no other known species that has the head and metasoma black in combination with an orange mesosoma and yellow legs.
This species is named after the famed British biologist Charles Robert Darwin. The epithet is a noun in the genitive case.
Holotype female:
The holotype is in perfect condition. Paratype OSUC 250602 is smaller in size than other specimens, and also has the mesal area of T2 smooth, the anterior mesal area of T3 mainly smooth with the exception of some weak, sparse longitudinal costae contrasting with the rest of specimens in which the sculpturing of T3 is as follows: anterior third is weakly costate sublaterally, weakly coriaceous mesally but otherwise granulose. Specimen OSUC 339589 has the metasoma lighter in color than other specimens.
This is the only known species that has a longitudinally depressed vertex. In the Neotropics there is one undescribed species (in CNCI) with a very conspicuous longitudinal depression on the vertex, but the general sculpture of the body is very different from
This species is named after the hymenopterist M. C. Day, now retired from the Natural History Museum, London. The epithet is a noun in the genitive case.
Holotype female:
The holotype is in perfect condition.
This is one of two species within a group of taxa that has an obscured netrion, notaular crenulae poorly defined, and the mesoscutum completely sculptured (without smooth areas) and entirely covered by short, abundant setae.
This species is named after the Australian hymenopterist I. D. Galloway. The epithet is a noun in the genitive case
Holotype female:
The holotype is in perfect condition; the paratype has the right hind wing and metasoma detached but glued to a card point.
This species can be easily distinguished from all other
This species is named after the Genting Tea Estate, Malaysia from where the species was collected. The epithet is used as an adjective.
Holotype female:
The holotype is in perfect condition.
This species can be distinguished from all other
This species is named after the African country from which it was collected. The epithet is used as an adjective.
Holotype female:
The holotype is in perfect condition.
Holotype female,
The holotype is slide mounted and partly destroyed (
This species can be separated from all other species with an obscured netrion by the subtriangular area below the eyes (seen in anterior view) that narrows towards the mandibles, unsculptured notauli, and non-ornamented occipital carina.
This species is named after our colleague and arachnologist Dr Mark Harvey from the Western Australian Museum, who also manages to collect numerous parasitic
Holotype female:
The holotype is in good condition except for the right wings which are missing. The paratypes are in good condition except OSUC 237919 which has the metasoma detached and glued to the wings.
In some female specimens the color of the mesoscutum varies from completely yellow to almost completely brown. Additionally, the color of the lateral portions of the T1 horn may vary from completely yellow to almost completely light brown.
This species can be distinguished from the other species from Fiji, all of which lack notauli, by the convex mesosoma, and the short distance between the posterior ocellus and occipital carina (approximately 0.5× the ocellar diameter).
This species is named after our friend and colleague Dr John Heraty, from the University of California, Riverside. The epithet is a noun in the genitive case.
Holotype female:
The holotype and paratypes are in perfect condition.
Male. Unknown.
This species is named after Dr Steve Heydon at the Bohart Museum, who collected this magnificent species. The epithet is a noun in the genitive case.
Holotype female:
The holotype is in perfect condition as are the paratypes except for OSUC 238768 which has the right antenna missing. Some specimens have the body color completely yellow and without darker areas on the metasomal terga.
This species can be distinguished from
This species is named after the dipterist Dr Mike Irwin, who collected the type series. The epithet is a noun in the genitive case.
Holotype female:
The holotype is in perfect condition as are the paratypes.
Within the group of species with a netrion present, the occipital carina almost touching the orbital carina, and having a sculptured gena,
This species is named after Ms Dorothy Jackson, who collected
Holotype female:
The holotype is in perfect condition as are the paratypes.
This species is named after the East New Britain word ‘kiau’ (in Kuanua language) which means ‘egg’, and refers to the stage of the host parasitized by
Holotype female:
The holotype is in perfect condition.
Along with
This species is named after the great French zoologist Jean-Baptiste Lamarck. The epithet is a noun in the genitive case.
Holotype female:
The holotype specimen is in good condition.
This species can be easily separated from all other
Holotype female:
The holotype is in good condition except that the right hind wing and right antenna are detached from body and glued to the point.
This species can be distinguished from all other
This species is named after the island from which the species was collected, Madagascar. The epithet is used as an adjective.
Holotype female:
The holotype and paratype are in perfect condition.
This species is very similar to
28 females, 1 male:
We were unable to obtain any of the type series, and so our interpretation of
When taking all of the available material into account some variation is evident in the females of this species: the antennal clava can be completely yellow or the distal half of the clava slightly darker, and the vertex and mesoscutum vary from completely yellow to slightly honey yellow in color. In addition, the specimens: OSUC 239209, OSUC 239211– 239213, OSUC 238767, OSUC 238803-238805, OSUC 248896, OSUC 250858, OSUC 261831, OSUC 261852, OSUC 268732, OSUC 321806 and OSUC 321894 differ from the remaining material by having the body completely yellow, but otherwise they match
Thisspecies belongs to a group that has short, smooth notauli, a well-defined netrion, central keel present, and sculptured antennal scrobe always with transverse costae in the background. Within this group
Holotype female:
The holotype is in perfect condition.
Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: upper 1/3 granulose, lower 1/3 with transverse foveae, otherwise smooth. Netrion: present, smooth, well developed, sub-obovate. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: weakly rugulose mixed with weak granulae. Sculpture of mesoscutellum: granulose. Mesoscutellar profile: elevated, anterior margin higher than posterior. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: densely, finely rugulose. Shape of propodeal anterior spine: elongate, narrow, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: weakly coriaceous. Sculpture of upper 1/4 of mesopleuron: densely longitudinally costate across half width. Metapleural sculpture: mainly smooth, lower third sparsely longitudinally carinate.
This species is very similar to
The name of this species is an arbitrary combination of letters and is used as a noun in apposition.
Holotype female:
The holotype is in perfect condition. The differences between
This is the only species of
This species is named after our friend and colleague, the chalcid specialist and great insect collector Dr John Noyes from the Natural History Museum, London. The epithet is a noun in the genitive case.
Holotype female:
The holotype has the left legs separate and glued to the point. Otherwise it is in good condition. The paratype in perfect condition: it has the metasoma completely honey yellow and the remainder of the body, including the legs, yellow.
This species is named after the trichogrammatid specialist Dr John Pinto, formerly of the University of California, Riverside. The epithet is a noun in the genitive case.
Specimens from South Australia have been reared from eggs of
Holotype female:
The holotype is in perfect condition; the paratypes are for the most part in very good condition except for OSUC 239063 and OSUC 239074 which have the heads detached from body and glued to the point, and OSUC 239046 which is missing the metasoma. Some females have a different color pattern in that the whole body is dark brown with the anterior edge of T1 (except the horn), T2 and the legs yellow.
This species can be distinguished from all other
This species is named after the dipteran expert Dr Evert Schlinger, collector of this species. The epithet is a noun in the genitive case.
Holotype female:
The holotype is in perfect condition.
This is the only known species to have a furrow at the lateral areas of the antennal scrobes in combination with a completely dark brown body.
This species is named after our friend and colleague, hymenopterist Dr Mike Sharkey from the University of Kentucky. The epithet is a noun in the genitive case.
Holotype female:
The holotype is in perfect condition except for the left antenna which is detached from the body but glued to the point; the paratype specimens are in good condition.
Holotype female,
The holotype of
The following color variations occur in this species: the body ranges from yellow with the tips of the propodeal spines and T1 horn dark brown to mostly dark brown with the posterior area of the mesoscutum and mesoscutellum light yellow. The occipital carina of the specimens OSUC 239180 and OSUC 239173 is present but very fine which renders it difficult to see; in all other specimens it is well developed. The sculpture also varies between the anterior propodeal spines, from being completely smooth to completely punctate in some specimens with darker color (i.e. OSUC 239165, 265167, 239170–239172). The sculpture of the mesal portion of T2 varies from in the development of the costae. In any case there is always coriaceous background sculpture.
The species
This species is named after Veronica Valerio, sister of the first author. The epithet is a noun in the genitive case.
Holotype female:
The holotype has all but one leg missing, but otherwise is in perfect condition; the paratypes are in good condition. The color of the metasoma varies from completely dark brown (matching the color of the head and mesosoma) to being conspicuously lighter than the head and mesosoma. The color of the antenna may vary from yellow to brown.
Along with
This species is named after the great English naturalist Alfred Russel Wallace: long live his legacy! The epithet is a noun in the genitive case.
Holotype female:
The holotype generally is in good condition: the left front and hind legs are glued to the point, and the right hind leg is covered in glue. The metasomal color pattern varies between specimens from almost completely yellow (i.e. OSUC 238004) to being mostly honey yellow with whitish areas on T3 and dark brown T4–T6 (i.e. OSUC 238001). In some specimens (i.e. OSUC 239020) the upper area of the T1 horn is dark brown with the remainder of the metasoma yellow, and the mesoscutum (which is normally light yellow) being a honey yellow color mesally.
This species is named after our friend and colleague, the microgastrine systematist Dr Jim Whitfield at the University of Illinois. The epithet is a noun in the genitive case.
Holotype female:
Holotype specimen is in perfect condition. Most of paratype specimens are in good condition.
This species can be separated from all
This species is named after the Australian entomologist and invertebrate photographer Paul Zborowski. The epithet is a noun in the genitive case.
Holotype female:
The holotype is in perfect condition as are the paratypes. In some specimens (e.g. OSUC 237956) the area between the propodeal anterior spines has some dark honey yellow color instead of the typical yellow.
This species can be separated from all the species which have the occipital carina almost touching the orbital carina by the absence of the S2 anterior carina, the posterior medial area of the mesoscutellum not depressed, and the complete absence of the lagrimal. The closest species is
This species is named after the anime character ‘Invader Zim’, in reference to the invasion of the spider egg sacs that occurs when
Holotype female:
We would like to thank Luciana Musetti (at OSUC) for her critical help with the management of specimens and support during the project; to Roger Burks (at OSUC) for his help in scoring characters for some Australian species; to Danielle Strander (The University of Adelaide) for her illustrations of the T1 horns, and to Sara Hemly, Joe Cora, and all the students at OSUC for their help with managing/ handling of specimens. Also thanks to everyone involved in the
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antenna |
|
antennal scrobe |
|
anterior ocellus |
|
area, areas |
|
body |
|
carina, carinae |
|
central keel |
|
compound eyes |
|
costae |
|
costulae |
|
coxae |
|
depression |
|
edge |
|
eyes |
|
face |
|
fore wing |
|
frons |
|
furrow, furrows |
|
gena |
|
head |
|
lateral ocellus, lateral ocelli |
|
lateral propodeal areas |
|
legs |
|
malar space |
|
mandible |
|
margins |
|
mesepimeron |
|
mesepisternum |
|
mesonotum |
|
mesoscutellum |
|
mesoscutum |
|
mesoscutum length |
|
mesosoma |
|
metapleuron |
|
metasoma |
|
netrion |
|
notaulus |
|
occipital carina |
|
occiput |
|
ocellus, ocelli |
|
orbital carina |
|
patch |
|
plate |
|
posterior ocellus |
|
pronotum |
|
propleuron |
|
propodeum |
|
ridges |
|
scrobe |
|
sculpture |
|
seta |
|
spines |
|
tegula |
|
terga |
|
torular triangle |
|
vein |
|
vertex |
|
wing vein |
|
wing, wings |
|
Characters and their states used in the phylogenetic analysis of
1. Lower head shape in anterior view
0. moderately short and strongly narrowed towards mouth, head appearing short and broad [101]
1. elongate and broad at mouth, head appearing elongate and somewhat thin [77]
2. Lagrimal
0. absent [35]
1. present, large [36]
2. present, small
3. Compound eye size in relation to head height (in anterior view)
0. approximately 1/2 or more [75]
1. approximately 1/3 or less [219]
4. Antennal scrobe sculpture
0. smooth or nearly so [87]
1. with granulate, coriaceous or rugulose sculpture [171, 273]
2. with confuses sinuate transverse costate sculpture throughout [225]
3. upper half with semicircular fine ridges, remainder smooth
4. almost completely cover with semicircular fine ridges
5. covered with vertical carinae
5. Surface of torular triangle
0. slightly bulging [117]
1. flat [287]
6. Central keel on frons
0. absent [193]
1. present (partially or throughout antennal scrobe height) [75]
7. Mandibular teeth shape
0. all subequal in length
1. middle tooth clearly longer than upper and lower teeth
2. upper and middle teeth subequal in length, lower tooth conspicuously smaller than upper two
8. Malar space sculpturing
0. smooth or mainly smooth, without fan-like sculpture
1. only with conspicuous fan-like sculpture
2. completely cover by sinuate, transverse, fine ridges
3. almost completely cover with semicircular, fine ridges from antennal scrobe
4. completely cover by granulate sculpture, without fan-like sculpture
5. with sparse, fine, dorsoventral carinae, weak granulae in background
6. weakly rugulose throughout, without fan-like striae
7. with sparse, short, fan-like striae mixed with weak coriaceous sculpture
8. with weak rugulose sculpture mixed with granulate sculpture
9. Surface of medial area of vertex
0. slightly depressed [43]
1. slightly round [39]
2. conspicuously depressed
10. Compound eye setosity
0. not evident, if observable at 60× magnification then very short and sparse [285]
1. clearly long and dense [10]
2. short and dense
11. Lateral ocellar size
0. smaller than the combined length of the 3 adjacent ommatidia of the compound eye [290]
1. larger than the combined length of the 3 adjacent ommatidia of the compound eye [15]
12. Distance between lateral ocellus and occipital carina
0. less or equal to 0.49× of ocellar diameter [11]
1. between 0.5× and 1.2× of ocellar diameter [31]
2. larger than 1.5× of ocellar diameter [29]
13. Distance between lateral ocellus and compound eye
0. 0.33× ocellar diameter or less [15]
1. approximately 0.5× ocellar diameter
2. approximately 0.66× ocellar diameter
3. 1× or more ocellar diameter or more
14. Surface of vertex
0. evenly rounded [29]
1. conspicuously bent medially [43]
15. Distance between occipital carina to orbital carina
0. well separated by distance at least 2× width of occipital carina [32]
1. separated by distance slightly greater than width of occipital carina [216]
2. contiguous or nearly so, separated by distance subequal to width of occipital carina [290]
16. Shape of midupper area of occipital carina
0. well-defined and sharply bent down, “M” shaped [245]
1. well-defined and slightly bent medially [182]
2. well-defined and slightly curved upwards [2]
3. carina virtually absent, not well-defined
17. Netrion
1. present, well-defined [20]
2. represented only by sculpture at lateral pronotal areas [19]
18. Notauli
0. completely absent [182]
1. present as a smooth furrow [188]
2. completely obscured due to dense longitudinally costate sculpture
3. present as a furrow with transverse elements along their length [284]
19. Mesoscutum sculpture
0. smooth or mainly smooth [242]
1. covered with broad longitudinal sculpture
2. covered with just one type of sculpture other than longitudinal or granulate sculpture [100, 128]
3. anterior area with sculpture different to remainder of mesoscutum [146]
4. covered with granulate sculpture [104]
20. Mesoscutellum sculpture
0. completely smooth
1. mainly smooth with row(s) of punctures laterally
2. mainly smooth with sculpture other than row(s) of punctures laterally
3. covered with sinuate longitudinally costate sculpture [260]
4. covered with granulate or rugulose sculpture [74, 278]
5. anterior area covered with small, dense rugula, otherwise with coarse, spaced rugulae [284]
6. covered with coriaceous sculpture [212]
21. Mesoscutellum shape
0. flat, conspicuously wider than long, midposterior area not depressed [212]
1. globose to very globose
2. flat, conspicuously wider than long, midposterior area depressed
22. Mesoscutellum posterior edge (in dorsal view)
0. lobed medially, ‘U’ shaped
1. bent medially, ‘W’ shaped
2. more or less straight or slightly curved [212]
3. conspicuously curved
23. Lateral propodeal areas
1. only with transverse ridges (dense or not)
2. with confused coriaceous sculpture but no transverse ridges present
3. only with longitudinal ridges
4. upper half with longitudinal ridges, remainder covered with granulate sculpture
5. with confused areolate-foveate sculpture
6. with rugulose sculpture
24. Propodeal anterior spines shape (in lateral view)
0. short, small, tips round
1. elongate, of approximately uniform width throughout, and tips round
2. lobate at apex which is much wider than base
3. spines absent
4. short, broad, tips acute
5. elongate, of approximately uniform width throughout, and tips acute
25. Inter-propodeal anterior spine area (in dorsal view)
0. smooth or nearly so
1. transversely costate
2. with longitudinally costate sculpture
3. with few lateral semi-transverse ridges
4. rugulose throughout
26. Fore wing veins
0. at most with basal area of subcostal vein tubular, otherwise veins absent
1. all veins present and tubular
27. Shape of wings
0. slightly convex, with slight constriction at 2/5 of its length [274]
1. very convex in lateral view, with conspicuous constriction at 2/5 of its length
2. flat, not convex, without obvious constriction at 2/5 of its length
28. Stigmal vein (r-rs)
0. absent
1. very short, less than 2.5× length of marginalis [239, 293]
2. elongate, more than 2.6× length of marginalis [227, 295]
29. Postmarginal vein (R1)
0. absent [294]
1. present, minute
2. present, short
30. Shape of T1 horn (posterior view)
0. elongate and narrow
1. short and broad [156]
2. short and narrow
3. elongate and broad
4. not applicable because of the absence of horn
31. T1 horn posterior lateral carinae
0. absent
1. present but not conspicuously cristate
2. present, conspicuously cristate
3. not applicable because of the absence of horn
32. Sculpturing on apical area of T1 horn (dorsal view)
0. absent (smooth)
1. with dense longitudinal carinae throughout
2. mostly smooth with few longitudinal carinae
3. not applicable because of the absence of horn
33. Sculpturing on posterior surface of T1 horn (posterior view)
0. absent (smooth)
1. mostly covered with well-defined longitudinal carinae
2. mostly smooth with few longitudinal carinae
3. with transverse ridges throughout
4. not applicable because of the absence of horn
34. Shape of T1 horn in cross section
0. subcircular or dorsoventrally compressed
1. laterally compressed
2. not applicable because of the absence of horn
35. T2 lateral carinae
0. absent
1. present but not well developed
2. conspicuously present
36. T2 length vs. length of T3
0. T2 longer than T3
1. T2 equal to T3 or virtually so [88]
2. T2 shorter than T3 [82]
37. Ovipositor shape
0. straight or curved into T1 horn [1B]
1. ovipositor bent over inside T1 horn (i.e. U-shaped) [1C]
2. T1 horn absent so ovipositor straight and shorter than metasoma [1A]
38. T1 shape
1. T1 narrow and elongate, approximately as wide as long or longer than wide [88]
2. T1 short and broad, conspicuously wider than long [76]
39. S2 anterior carinae
0. absent [68]
1. present, but inconspicuous, uninterrupted [67]
2. clearly evident, uninterrupted [69]
3. clearly evident, interrupted medially [65]
40. Metasoma shape
0. short and compact
1. elongate, not particularly constricted at T1
2. elongate, strongly constricted at T1 [72]
41. Number of segments in antenna
0. with 11–12 segments
1. with 8–9 segments [315]
Matrix character states used for the phylogenetic analysis of
Character 00000000011111111112222222222333333333344
number 12345678901234567890123456789012345678901
Idris_floris 11001101101010222024014321220020000202111
Idris_sp1 00010121110000221024002301211101300202011
Idris_sp2 00010101110100222024023321222101100102011
Idris_sp3 00011101120100122044023321220433420121201
C_axfordi 00011001101210122033021100000200210112220
C_copelandi 00021002101110022213121210100221212111220
C_diazae 00031001100130022002021100100210011111020
C_harteni 00011004100110021000021000000210011112020
C_jenningsi 00001001100200002100021130100200110112120
C_leblanci 00021000100110022211121210100131112112220
C_lucianae 00021000100200022211121210100220012112220
C_normani 00041003100130022212121210100221212111220
C_sallyae 00001001100211122100021020000200210112120
C_tessae 00001001101100122132021100000210011112?20
C_tullyae 00001001101130112100021020000200110112120
C_watshami 00021002100120022211121210100222212111220
C_bouceki 00001101100200112132021010000200210112020
Od_africanus 1000011[14]111100[01]21134026101020101111112320
Od_aldrovan 12010118111100021124021101020201212211220
Od_anningae 10000117110100121124026101021101212111320
Od_australiensis 00001101100200[01]22244020021010000110212120
Od_baeri 10010117110100021124026101020101211211020
Od_berryae 00000101100200022244023021010000210111120
Od_bosei 10000118111100021124021101020201212111220
Od_cardaleae 10020114111100021144026141020101312212220
Od_darwini 10021115111100021134021401020101212112220
Od_dayi 10001117011102021124026101020201212111220
Od_gallowayi 00000101100200012344021001010000110212120
Od_gentingen 11001111111000021134023111021001111212220
Od_guineaensis 10021116111100021124006121021001111112320
Od_hackeri 10011116111?0?1211230210?1020101312112320
Od_harveyi_sp3 00000101100200012130021021010000210210120
Od_heratyi 10011118111000111023026121021001212212020
Od_heydoni 10011114111100021144021101021001212212220
Od_irwini 10010118111100011044023401020101210211020
Od_jacksonae 10000117111100221144026501020001111111320
Od_kiau 10021116113100021124026001020001212211020
Od_lamarki 10000014111100021124023101020101212212220
Od_longiceps 100010141111000211260234[03]1020101011212220
Od_madagasc 10001110110100121126026401020202211211220
Od_markadicus 100[01]0117111100121144021101020201212111220
Od_mayri 10020116111100021144026101020101212112220
Od_mot 12000114111100021124026101020201212112220
Od_noyesi_sp2 101111171111000213440264[02]1020101111212220
Od_pintoi 10000110100200021120021401010000210211120
Od_schlingeri 10011114111100111024023001020101010212020
Od_sharkeyi 10021111111100021124026101020101011212220
Od_spinosus 10011116110100021123026141020101112212220
Od_veroae 10011114111100221034026401020101010211020
Od_wallacei 10011017110200021144021101020001111212320
Od_whitfieldi 10020116111100[01]21124026101020101212212220
Od_zborowskii 00020101100200012325021121010001110112120
Od_zimi 00000111110100221134021401022201111110220
Od_flaviss 10000111111100022132023131020101012211220
Od_szaboi 10050111111100022124026131020201012111220
Od_sp4 10000111220101002344211001221302012212220
Od_sp5 10100111110200022123021001221101012211220
Od_sp6 10000110120100022132021101010001011211220
Od_sp7 10000111121100221130021541011001312111220
Habitats associated with
2° forest
among undergrowth
behind vegetable garden
burnt pine
bush
campground; pond
canopy
casuarina forest
clearing mostly on scraps wood; nr. edge clearing
closed forest
coastal dunes; vegetation; sandy area
creek
creekbed
dry evergreen
dry evergreen forest
dry forest
dry gallery woodland
dry scleophyll forest
dry sclerophyll
dry sclerophyll gully
dry wash; open area
dump; open forest
eucalypt &
eucalypt litter; grazing land
evergreen forest
flooded low forest
forest
forest edge
forest gap
forest w/ few large trees & good general cover
gallery forest
grape vines; corrugated cardboard
grass; roadside; floodplain
grassland
high forest
hill; evergreen forest
hilltop; road
indigenous forest margin
mangrove swamp
medium forest
midelevation oak forest
midelevation secondary forest
mixed deciduous
mixed deciduous forest
moist evergreen
open forest
open sclerophyll gully
orange leaf
pasture
pine lands
pine treefarm
rainforest
rainforest gully
rainforest margin
rainforest; litter
rastojo forest
relict indigenous forest
riverine woodland
sand; low altitude dense humid forest
sclerophyll,
scrub; pine
seasonally swampy woodland next to grassland
second growth bush
secondary rainforest
shore; gully
small undisturbed riparian forest; valley
spiny forest thicket
stream
trail
tropical dry forest
tropical dry forest on tsingy
tropical dry forest; tsingy
tropical forest; botanical garden
tropical rainforest
tuart forest
TV aerial
under bark
under gum bark
wet forest
wet grazing floor
woodland
woodland next to grassland
Darwin core taxa list. (doi:
Darwin core specimen data. (doi: