This is an expanded version of the dataset of Shaw (1990b), which comprised 20 characters. We deleted his character 19 (metasomal shape elongate/compact) as the separation of its states break down when the fossils are included. His other characters are indicated in the list below. Most of the characters added are included to help resolve the relationships among the outgroup taxa and/or the fossil taxa not included by Shaw (1990b).
(0) below or level with ventral margin of eyes (Fig. 2A–F)
Figure 2. Head and antenna of Megalyridae: (A–C) anterior view; (D–F) lateral view. (A, G) Carminator affinis; (B) Neodinapsis peckorum; (C, D) Rigel chiliensis; (E) Dinapsis sp.; (F) Megalyra wagneri.
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(1) above ventral margin of eyes
State 1 is observed only in the Evanioidea and Trigonalidae included, as well as in Guyotemaimetsha.
A well-developed subantennal groove is observed on all extant and extinct Megalyridae included here, but not in Maimetshidae; among the outgroup taxa, it is observed only in Orussus, and its presence is not a ground-plan feature of Orussidae (Vilhelmsen, 2003b). Many species of Aulacidae, especially Pristaulacus spp. (but not P. strangaliae), also have well-developed subantennal grooves (Turrisi et al., 2009), but it is not a ground-plan trait of this family either.
3. Dorsal carina of subantennal groove (Shaw, 1990b: char. 4):
Inapplicable when the subantennal groove is absent. Shaw (1988) stated in the diagnosis of Carminator that ‘subantennal groove shallow, not bordered by carinae’; nonetheless, in Shaw (1990b)Carminator was scored as state 1. Carminator nooni appears to have a well-developed carina above the torulus, but not further posterior, and two other species described recently from Japan by Mita et al. (2007) show a distinct carina. Accordingly, this character has been scored polymorphic here (state 0 and 1).
4. Number of flagellomeres (ORDERED):
(0) more than 12 (excluding scape and pedicellus)
The antennae are broken in the holotype of Neodinapsis peckorum, and hence this taxon is scored as unknown. Twelve flagellomeres, in addition to being observed in Megalyridae, are observed in Pristaulacus (only in females, Turrisi et al., 2009) and Gasteruption.
(1) at least some flagellomeres compact (Fig. 2G)
Compact flagellomeres constitute an apomorphy of Dinapsis and Ettchellsia, but are present in many fossil megalyrids as well.
6. Median notch on anterior margin of clypeus:
The median notch is observed in Prodinapsis janzeni, P. minor, P. succinalis and Ukrainosa, as well as in Orthogonalys and Schlettererius among the outgroup taxa.
7. Mandible configuration (ORDERED):
(0) two or fewer mandibular teeth present
(1) three mandibular teeth present
(2) four or more mandibular teeth present (Fig. 2A)
State 2 is a potential apomorphy of Maimetshidae, but it is also observed in Megalava and Carminator (Shaw, 1988).
(0) symmetric, same number of teeth on mandibles
(1) asymmetric, different number of teeth on mandibles
The presence of asymmetric mandibles is a putative apomorphy for Maimetshidae and Trigonalidae.
(1) cuticular teeth around median ocellus present
The ocellar corona consists of a curved row or circlet of discrete teeth extending posteriorly between the median and lateral ocelli (see e.g. Vilhelmsen, 2003b). It is present only in Orussus and Schlettererius among the taxa included here.
State 0 was scored for Dinapsis in Shaw (1990), but a short crenulate sulcus is clearly visible immediately posterior to the median ocellus in at least D. oculohirta and D. hirtipes (see http://www.waspweb.org/Megalyroidea/Megalyridae/Dinapsini/Dinapsis). Similarly, a ‘longitudinal row of small foveae extending between dorsal ocelli’ was observed in Megalyra baltica by Poinar & Shaw (2007) and Perrichot (2009). Thus this character is scored as polymorphic for Dinapsis and Megalyra. The sulcus is present in most extinct Megalyridae, but absent from all extant members of the family except the Dinapsis spp. mentioned above.
Setae on the eyes are present in Megalyra baltica, but absent in extant Megalyra spp.; hence, this taxon has been scored as polymorphic. The occurrence of setae varies within Megalyridae.
There is considerable variation in this character within Dinapsis; some species have the carina not fully developed, whereas others have two fully developed carina present, one behind the other (Fig. 2E). However, all Dinapsis spp. appear to have the carina at least partly developed, so Dinapsis has been scored state 1 for this character.
13. Occipital carina basally (Shaw, 1990b: char. 3) (UNORDERED):
(0) curving towards back of head (Fig. 2D)
(1) branching, anterior branch curving towards mandible (Fig. 2E)
State 1 is observed in all Megalyridae except Rigel and Megalyridia, which have state 0, and Carminator, which has state 2.
14. Pronotum anteromedially:
(0) well-developed, protruding
Figure 3. Mesosoma of Megalyridae: (A–C) dorsal view; (D–F) lateral view. (A) Megalyridia capensis; (B) Carminator affinis; (C) Dinapsis sp.; (D) Megalyra wagneri; (E) Ettchellsia sp.; (F) Rigel chiliensis.
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Having the pronotum reduced anteromedially is a ground-plan feature of Megalyridae, but is also observed in Gasteruption and Pristaulacus.
15. Pronotum posteromedially:
(0) high, well-developed posteriorly of transverse sulcus
The posteromedially reduced pronotum is a putative apomorphy of Evaniomorpha s.s. (reversed in Evania).
16. Anterior thoracic spiracle (ORDERED):
(0) concealed or not surrounded by sclerotized cuticle
(1) exposed, with posterior part free, not surrounded by ‘pronotal’ cuticle
(2) exposed, entirely surrounded by ‘pronotal’ cuticle (Fig. 3D–F)
State 1 is observed only in Megazar and is probably present in Megalava. Apart from these reversals, state 2 can be regarded as an apomorphy of Megalyridae and Ceraphronoidea.
17. Fore tibial apical stout spines (Shaw, 1990b: char. 14):
Figure 4. Leg features of Megalyridae, lateral view. (A, B) Carminator affinis, fore tibia and hind coxa; (C) Ettchellsia sp., hind leg; (D) Rigel chiliensis, hind tibia.
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These spines are present in all Megalyridae except Rigel, Megallica and two Prodinapsis spp.
18. Median mesoscutal sulcus:
(0) absent or weakly developed (Fig. 3B)
In Carminator, the sulcus is at most weakly visible (see Mita et al., 2007: 202). In the Megalyra minuta species group, the sulcus is reduced, so Megalyra has been scored polymorphic. However, the sulcus is probably present in the common ancestor of Megalyra (Shaw, 1990a), as it is present in all other Megalyridae except Carminator. The presence of this sulcus is probably a retained plesiomorphy, even if it is absent from most other Apocrita (Vilhelmsen et al., 2010).
19. Anterolateral transverse carina on mesoscutum:
Megalyra has a ‘small acute process' (Naumann, 1987: 219) or a ‘sharp spine’ (Poinar & Shaw, 2007: 67) on the anterolateral corners of the mesoscutum. However, the terminology ‘small projection’ should be preferred (Perrichot, 2009), and it is probably equivalent to the carina observed in most other megalyrids, so Megalyra is scored as state 1. The carina is situated more medially in Dinapsis, but is considered to be homologous here, and thus Dinapsis is scored 1. The presence of the carina is a putative apomorphy of Ceraphronoidea and Megalyridae, albeit with reversals within the latter.
According to Mita et al. (2007), ‘scratches’ of notauli are present in Carminator helios and C. japonicus. However, these structures are more likely to be the parapsides (see the following character), so Carminator has been scored state 0. Hence, the absence of notauli is probably a megalyrid ground-plan feature.
State 1 is observed only in Megazar and Megalava among the fossils. Parapsides are present in all extant Megalyridae except Megalyra. They are difficult to observe in many of the extant taxa, making it possible that they have been overlooked in some of the fossils.
(0) meeting at inner angles for at least a short distance (Fig. 3A, C)
(1) separated by the anterior portion of inner axillar grooves or by large foveae (Fig. 3B)
Carminator has been scored polymorphic for this character. Regardless, the presence of large axillae abutting in the middle is apparently a ground-plan feature of Megalyridae, probably a retained plesiomorphy.
23. Mesocoxal articulations:
(0) short, articulation at base of coxa
(1) elongate, articulation displaced distally on coxa
State 1 is an apomorphy of Evaniomorpha s.s.
24. Mid-tibial spurs (ORDERED):
State 0 is so far only observed in Schlettererius (Stephanidae). State 1 is an apomorphy of Megalyridae, albeit with some fossil taxa reverting to state 2.
25. Posterior mesopleural margin (Shaw, 1990b: char. 6):
State 1 is observed in the extant taxa Carminator, Cryptalyra and Megalyra, and in the fossil taxa Megalava, Megazar and Rubes.
26. Row of foveae posterodorsally on mesopleuron:
(1) row of foveae extend along posterodorsal margin of mesopleuron, adjacent to boundary with metapleuron (Fig. 3D)
According to Shaw (1990b), state 0 is an apomorphy of Ettchellsia; however, foveae are also absent in many Dinapsis spp. (see www.waspweb.org, all Dinapsis spp. illustrated there where this character can be observed have state 0). Hence, Dinapsis has been scored polymorphic for this character. The foveae are absent also in Carminator, Cryptalyra and Megazar.
27. Metapleural pilosity:
(0) metapleuron mostly bare (Fig. 3F)
(1) metapleuron with well-developed pilosity (Fig. 3D, E)
State 0 is a putative apomorphy of Megalyridae, albeit with reversals in Ettchellsia, Megalyra and Neodinapsis.
28. Longitudinal carina laterally on hind coxa (Shaw, 1990b: char. 15):
(0) absent or not visible in lateral view (Fig. 4B)
Some taxa (e.g. Carminator) have a carina developed at the dorsal margin of the hind coxa. This is difficult to observe in lateral view if the hind coxa is closely appressed to the metasoma. Accordingly, only taxa (Dinapsis, Ettchellsia) in which the carina is situated on the outer side of the coxa and thus clearly visibly laterally have been assigned state 1. Within Dinapsis, the extent of the carina varies considerably, but it is always present. The presence of the carina is hence an apomorphy for Dinapsis and Ettchellsia.
29. Hind coxal sculpture (Shaw, 1990b: char. 16) (UNORDERED):
This character is highly variable within Dinapsis, which has been scored polymorphic for all three states. It also displays considerable variation across Megalyridae, making it of limited phylogenetic value.
30. Comb-like spines along inner margin of hind femur and tibia:
State 1 is observed only in Megazar and Megalava; the spines may extend to the hind trochanter in some specimens.
Erect setae are observed in Carminator, Cryptalyra, Dinapsis, Ettchellsia and Megalyra, as well as in Megaspilus.
A single hind tibial spur is observed only in Cryptalyra and Megalyra, but probably evolved independently in these two taxa.
33. Relative length, hind tarsomeres 2–4:
(0) elongate, combined length at least 1.5× of tarsomere 5
State 1 is a putative apomorphy for Dinapsis and Ettchellsia.
34. Tarsal pulvilli (= plantulae):
(1) present, at least in females
The presence of tarsal pulvilli, a rare occurrence among Apocrita (Schulmeister, 2003), is a putative apomorphy of Maimetshidae and Trigonalidae.
35. Subapical projections on the tarsal claws:
(0) absent or weakly developed
Subapical projections are present on the tarsal claws in Evania, Orthogonalys, Pristaulacus and Guyotemaimetsha.
The absence of this vein is an apomorphy of Carminator and Cryptalyra.
37. FW, RS apically (ORDERED):
(1) weakly developed, nebulous or infumate (Fig. 5A, D)
(2) well developed, tubular (Fig. 5C)
Following (Shaw, 1990a), Megalyra has been scored polymorphic (states 0, 1 and 2). This character is highly variable throughout Megalyridae and difficult to interpret.
38. FW, RS configuration, when present (ORDERED):
(0) arched towards wing tip (Fig. 5D)
(2) arched towards stigma (Fig. 5C)
The taxa having this vein reduced (see previous character) have been scored inapplicable for this character, making it difficult to optimize unambiguously.
(1) present, branching from Rs + M (Fig. 5C)
The anterior part of this vein seems to be weakly developed in Rigel (Fig. 5A); this taxon was assigned state 0. This vein is apparently absent from the ground plan of Megalyridae, but present in Dinapsis, Ettchellsia and Neodinapsis as well as in most of the Cretaceous fossil Megalyridae and Maimetshidae.
(0) anterior, submarginal cell less than twice as large as discal cell (Fig. 5A, C)
(1) posterior, submarginal cell at least twice as large as discal cell (Perrichot, 2009, fig. 15.2)
State 1 is observed in Cretodinapsis, Megazar and Maimetshidae.
(0) absent or spectral, colourless (Fig. 5A–C)
(1) tubular or infumate (coloured) (Fig. 5D)
Following (Shaw, 1990a), Megalyra has been scored polymorphic. Otherwise, the vein is at most weakly developed in all Megalyridae except Cretodinapsis, Megalava and Megazar, as well as in Maimetshidae.
(0) absent distally of cu-a (Fig. 5B)
(1) at least darkened line present distally of cu-a (Fig. 5D)
This character is variable across Megalyridae.
43. FW, infumate banding pattern (Shaw, 1990b: char. 11):
The presence of infumate bands in the wing or infumation of the entire wing is a putative apomorphy of Dinapsis and Ettchellsia. However, among Megalyridae it is also observed in Megalyra and Megalyridia.
(0) absent or reduced, at most small swelling at RS junction (Fig. 5B–D, F)
(1) thick and broad beyond Rs (Fig. 5A)
Shaw (1990b) scored the pterostigma as absent in Carminator. Because of this variation within the genus, which was the only one to be scored absent by Shaw (1990b), we have recoded this character in only two states. Given this definition, a well-developed pterostigma is present only in Rigel and Cretodinapsis, as well as in Maimetshidae and all the outgroup taxa.
Figure 6. (A) Strict consensus of three trees of fit 9.46429 computed by implied-weights analyses with k = 5, only extant taxa included. (B) Strict consensus of 7399 trees of length 187 steps computed by equal-weights analyses, all taxa included. (C) Strict consensus of three trees of fit 15.38203 computed by implied-weights analyses with k = 5, all taxa included.
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(3) long, approaching wing margin (Fig. 5D)
According to Shaw (1990a), most Megalyra spp. have Rs very long in the hind wing and those that do not probably have it secondarily reduced; hence, Megalyra has been scored state 3. This character is highly variable within Megalyridae, state 2 probably being the ground-plan state.
46. Propodeal sculpture (Shaw, 1990b: char. 7) (UNORDERED):
According to Rasnitsyn & Brothers (2009), state 1 is the condition in all Maimetshidae except Guyotemaimetsha, which is state 2. Megalyra is scored polymorphic (states 1 and 2) following Shaw (1990a). This character varies considerably across Megalyridae.
(0) posteroventral, not far removed from metacoxal foramina
(1) anterodorsal, at least halfway between propodeal antecostal sulcus and metacoxal foramina
State 1 is an apomorphy of Evanioidea.
48. Ovipositor length (Shaw, 1990b: char. 20) (ORDERED):
(0) more than 2× body length (Fig. S1a)
(1) 0.75–1.25× body length (Figs S1b; S2a, b)
(2) less than 0.75× body length (Figs S1c; S2c, d)
Dinapsis was scored state 1 according to the present definition of the states, not state 2 as in Shaw (1990b). Many Megalyridae have state 2, only Megalyra has state 0, but state 1 is probably the ground-plan condition.