Research Article |
Corresponding author: Diego N. Barbosa ( barbosa.laelius@gmail.com ) Academic editor: Ângelo Pinto
© 2022 Diego N. Barbosa, Marcel Gustavo Hermes, Anderson Lepeco.
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:
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We present the first phylogenetic hypothesis for Mesitiinae based on 112 morpho-structural characters and 61 species. The results did not support Argaman’s tribal classification for Mesitiinae, since no tribes were found to be monophyletic. Anaylax was found to be paraphyletic, and Gerbekas, Heterocoelia, Parvoculus, Pycnomesitius, Sulcomesitius, Zimankos were found to be polyphyletic. Two new genera are proposed and described: Hadesmesitius gen. nov. and Brachymesitius gen. nov.; Botoryan is considered as a junior synonym for Zimankos. Three species status are revalidated; and 11 species combinations were proposed, so that all genera are now monophyletic. The results indicate the thickness of integument in Mesitiinae could be related to their protection against their hosts.
Biogeography, monophyletic, morphofunctional features, new genera, paraphyletic, polyphyletic, tribal classification
The aculeate family Bethylidae includes nearly 3,000 species of parasitoid wasps, representing one of the most diverse lineages of Chrysidoidea (
The monophyly of the Mesitiinae was supported by the analyses in
The specimens used in this study were borrowed from the following collections, with curators in parentheses:
The images were obtained using a Leica MZ80 Stereomicroscope attached to a Leica DFC 495 video camera and captured with LEICA LAS (Leica Application Suite V3.6.0) by Leica Microsystems (Switzerland), using a dome illumination system described by
The terms applied to the structures follow
The ingroup is composed by males of 61 species (Table
Except for Australomesitius Barbosa & Azevedo, 2016, known only from the female, 17 out of the 18 genera currently included in the subfamily were sampled. The outgroup includes representatives of all extant subfamilies of Bethylidae (Table
Table
A total of 112 characters (Appendix
Taxa | Specimen | Zoogeographic region | Repository |
---|---|---|---|
Ingroup (Mesitiinae) | |||
Domonkosini | |||
Pilomesitius madagascarensis Móczár, 1970 | Allotype | Madagascar |
|
Zimankos alluaudi (Kieffer, 1913) | Allotype | Ethiopic |
|
Zimankos makoa Barbosa & Azevedo, 2012 | Holotype | Madagascar |
|
Zimankos pondo (Benoit, 1968) | Paratype | Ethiopic, Oriental |
|
Zimankos rieki Móczár, 1976 | Paratype | Oriental |
|
Zimankos szentivanyi Móczár, 1976 | Paratype | Oriental |
|
Zimankos vechti Móczár, 1979 | Paratype | Oriental |
|
Heterocoeliini | |||
Botoryan discolor (Nagy, 1968) | Holotype | Oriental |
|
Gerbekas carcelli (Westwood, 1874) | Voucher | Palaearctic |
|
Gerbekas laosensis Móczár, 1975 | Holotype | Oriental |
|
Heterocoelia cursor (Kieffer, 1906) | Allotype | Palaearctic |
|
Heterocoelia fischeri Móczár, 1971, jr. syn. of Pycnomesitius peringueyi (Kieffer, 1913) | Holotype | Ethiopic |
|
Heterocoelia halidaiella (Westwood, 1874) | Voucher | Palaearctic |
|
Heterocoelia halidaii (Westwood, 1874) | Voucher | Palaearctic |
|
Heterocoelia hungarica (Kieffer, 1906) | Voucher | Palaearctic |
|
Heterocoelia nikolskajae Móczár, 1984, jr. syn. of H. obscura (Kieffer, 1906) | Paratype | Palaearctic |
|
Heterocoelia obscurus (Kieffer, 1906) | Neótipo | Palaearctic |
|
Pycnomesitius benoiti (Móczár, 1970) | Paratype | Ethiopic |
|
Pycnomesitius desenpunctatus Móczár, 1971 | Allotype | Ethiopic | BMNH |
Pycnomesitius peringueyi (Kieffer, 1913) | Voucher | Ethiopic |
|
Sulcomesitius kosztarabi Móczár, 1984 | Paratype | Oriental |
|
Sulcomesitius nepalensis Móczár, 1986 | Paratype | Oriental |
|
Sulcomesitius punctaticollis (Fouts, 1930) | Holotype | Oriental |
|
Sulcomesitius sp.01 | Voucher | Oriental | QSBG |
Sulcomesitius thailandensis Móczár, 1977 | Paratype | Oriental |
|
Sulcomesitius wahisi Móczár, 1984 | Paratype | Oriental |
|
Mesitiini | |||
Anaylax betsileo Barbosa & Azevedo, 2012 | Holotype | Madagascar |
|
Anaylax mahafaly Barbosa & Azevedo, 2012 | Holotype | Madagascar |
|
Anaylax simplicitus Barbosa & Azevedo, 2011 | Holotype | Ethiopic |
|
Astromesitius indistintus Barbosa &Azevedo, 2011 | Holotype | Ethiopic |
|
Astromesitius minutissimus (Móczár, 1971) | Voucher | Ethiopic |
|
Astromesitius olavoi Barbosa & Azevedo, 2019 | Holotype | Oriental | QSBG |
Clytrovorus fuscicornis (Kieffer, 1906) | Holotype | Palaearctic |
|
Clytrovorus horvathi (Kieffer, 1906) | Allotype | Palaearctic |
|
Clytrovorus merina (Barbosa & Azevedo, 2012) | Holotype | Madagascar |
|
Clytrovorus zafimaniry Barbosa & Azevedo, 2012 | Holotype | Madagascar |
|
Incertosulcus capensis (Kieffer, 1911) | Allotype | Ethiopic |
|
Incertosulcus consimilis Móczár, 1970 | Paratype | Ethiopic |
|
Incertosulcus krombeini Móczár, 1970, jr. syn. of Parvoculus indicus Kieffer, 1???95 | Holotype | Palaearctic | BMNH |
Incertosulcus priesneri Móczár, 1978 | Holotype | Palaearctic |
|
Incertosulcus soikai Móczár, 1970 | Holotype | Palaearctic |
|
Incertosulcus vanharteni Barbosa & Azevedo, 2011 | Holotype | Ethiopic | CNCI |
Incertosulcus vietnamensis Móczár, 1977 | Paratype | Oriental |
|
Incertosulcus sp.01 | Voucher | Oriental | QSBG |
Itapayos antaimoro Barbosa & Azevedo, 2012 | Holotype | Madagascar |
|
Itapayos sp.01 | Voucher | Oriental | QSBG |
Mesitius granulata Móczár, 1984 | Holotype | Oriental |
|
Mesitius kiefferi Nagy, 1970 | Holotype | Palaearctic | ZMB |
Mesitius krombeini (Nagy, 1968) | Holotype | Oriental |
|
Mesitius paenepunctata (Benoit, 1968) | Holotype | Ethiopic |
|
Metrionotus alutaceus (Benoit, 1968) | Paratype | Ethiopic |
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Metrionotus brevispinosus (Benoit, 1968) | Paratype | Ethiopic |
|
Metrionotus rufohumerus Móczár, 1984 | Paratype | Oriental |
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Metrionotus wolfi Móczár, 1970 | Paratype | Ethiopic |
|
Parvoculus indicus (Kieffer, 1905) | Holotype | Oriental |
|
Triglenusini | |||
Bradepyris baleariensis Barbosa & Azevedo, 2015 | Voucher | Palaearctic |
|
Bradepyris dimorphus (Kieffer, 1911) | Allotype | Palaearctic | BMNH |
Bradepyris inermis (Kieffer, 1906) | Allotype | Palaearctic |
|
Bradepyris jordanicus Barbosa & Azevedo, 2015 | Voucher | Palaearctic |
|
Bradepyris proximus (Kieffer, 1906) | Holotype | Palaearctic |
|
Moczariella centenaria (Barbosa & Azevedo, 2014) | Holotype | Ethiopic |
|
Outgroup | |||
Bethylus cephalotes Forster, 1860 | Voucher | Palaearctic |
|
Goniozus legneri Gordh, 1982 | Voucher | Holarctic, Neotropical |
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Chlorepyris longifoveatus (Azevedo, 1999) | Paratype | Neotropical |
|
Epyris variatus Côrrea & Azevedo, 2002 | Paratype | Neotropical |
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Apenesia sahyadrica Azevedo & Waichert, 2006 | Paratype | Oriental |
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Disssomphalus cervoides Azevedo, 2003 | Paratype | Neotropical |
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Plastanoxus westwoodi (Kieffer, 1914) | Voucher | Holarctic, Neotropical |
|
Sclerodermus irradiatus (Lanes & Azevedo, 2004) | Paratype | Oriental |
|
The character matrix (Table S1) was produced using DELTA software (
The searches for the most parsimonious trees were carried out in TNT version 1.5 (Goloboff et al. 2016, using the Ratchet, Sectorial Searches and Tree-Fusing searching strategies (
It has been argued that results based on characters properly weighted are to be preferred over those with all characters equally weighted (Farris 1969,
Bayesian analyses were conducted in MRBAYES 3.2.7 (
The implied weighting analysis using k = 11,674805 resulted in one most parsimonious tree, with 675 steps, fit = 23,72727, consistency index (CI) = 0.19, and retention index (RI) = 0.61 (Figs
Characters and character states. A Metapectal-propodeal complex in dorsal view; B Metasoma lateral view; C Wings in dorsal view, red = nebulous cubital vein, blue = nebulous anal vein, green = subcostal vein; D Hind wing in dorsal view; E Hypopygium in ventral view; F. Male genitalia in ventral view, red = cuspis, blue = genital ring.
Twenty-four characters were found as synapomorphies for the subfamily, 14 of them are exclusive transformations for Mesitiinae as listed below:
Ch. 5:1 malar space projected (Fig.
Ch. 23:0 contour of eye protruding (Fig.
Ch. 28:1 anterior depression of occiput present (Fig.
Ch. 31:0 ventral half of mesoccipital carina angled - (Fig.
Ch. 41:0 notauli of mesoscutum convergent posteriorly (Fig.
(Ch. 49:1 metapostnotal depression present (Fig.
Ch. 50:1 connection between central depression and triangular lateral depression of metapectal-propodeal disc (Fig.
Ch. 72:1 space between tegula and mesoscutum present (Fig.
Ch. 77:1 prestigmal abscissa of radial 1 of forewing present (Fig.
New nomina, nomenclatural acts and changes in combination in this study.
Original spelling/status | Current spelling/status | Spelling status |
— | — | Hadesmesitius gen. nov. |
— | — | Brachymesitius gen. nov. |
Anaylax simplicitus Barbosa & Azevedo, 2011 | Anaylax simplicitus Barbosa & Azevedo, 2011 | Hadesmesitius simplicitus (Barbosa & Azevedo, 2011) comb. nov. |
Incertosulcus krombeini Móczár, 1970 | Incertosulcus krombeini Móczár, 1970 | Brachymesitius krombeini (Móczár, 1970) comb. nov. |
Heterocoelia fischeri Moczar, 1971 | Pycnomesitius fischeri (Móczár, 1971) jun. syn. of P. peringueyi (Kieffer, 1913) | Gerbekas fischeri (Móczár, 1971) stat. rev. et comb. nov. |
Heterocoelia nikolskajae Móczár, 1984 | Heterocoelia nikolskajae Móczár, 1984 jun. syn. of H. obscura (Móczár, 1984) | Gerbekas nikolskajae (Móczár, 1984) stat. rev. et comb. nov. |
Sulcomesitius nepalensis Móczár, 1986 | Sulcomesitius nepalensis Móczár, 1986 | Metrionotus nepalensis (Móczár, 1986) comb. nov. |
Sulcomesitius wahisi Móczár, 1984 | Sulcomesitius wahisi Móczár, 1984 | Pycnomesitius wahisi Móczár, 1984 comb. nov. |
Heterocoelia laoensis Móczár, 1975 | Gerbekas laoensis (Móczár, 1975) | Sulcomesitius laoensis (Móczár, 1975) comb. nov. |
Botoryan Argaman, 2003 | Botoryan Argaman, 2003 | Botoryan Argaman, 2003 syn. nov. of Zimankos Argaman, 2003 |
Mesitius discolor Nagy, 1968 | Botoryan discolor (Nagy, 1968) | Zimankos discolor (Nagy, 1968) comb. nov. |
Zimakos makoa Barbosa & Azevedo, 2012 | Zimakos makoa Barbosa & Azevedo, 2012 | Pilomesitius makoa (Barbosa & Azevedo, 2012) comb. nov. |
Mesitius krombeini Nagy, 1968 | Mesitius krombeini Nagy, 1968 | Zimankos krombeini (Nagy, 1968) comb. nov. |
Ch. 85:1 constriction between metasomal sternite I and II present (Fig.
Ch. 87:1 lateral ventral lap metasomal tergum I present (Fig.
Ch. 89:1 metasomal segment II longer than others (Fig.
Ch. 101:1 projection of genital ring present (Fig.
Ch. 105:1 fusion between gonostipes and basivolsela present (Fig.
The other 10 characters states found as synapomorphies are not exclusive for Mesitiinae, but contributed to define the subfamily:
Ch. 7:1 orientation of malar space parallel (Fig.
Ch. 8:1 inner keel of mandible present (Fig.
Ch. 14:1 torulus and median clypeal carina fused (Fig.
Ch. 20:0 flagellomeres 1-11 slender (Fig.
Ch. 21:0 eye small (Fig.
Ch. 26:2 anterior ocellus crossing supra-ocular line (Fig.
Ch. 37:0 anterior margin of propleuron angled (Fig.
Ch. 81:0 hind wing with three distal hamuli (Fig.
Ch. 94:1 hypopygium as long as wide (Fig.
Ch. 106:1 cuspis with two arms (Fig.
Bayesian analyses largely corroborated the backbone of the relationships retrieved in parsimony (Figs S2, S3). Results from unpartitioned and partitioned analyses differed. In both analyses the genus Anaylax was not recovered as monophyletic, with Anaylax simplicitus being recovered as a distinct lineage relative to other species of the genus included in the present account. The unpartitioned analysis recovered Incertosulcus krombeini as sister group to Metrionotus, while in the analysis using partitioning by homoplasy score it was recovered nested within Metrionotus. However, the posterior probability of the clade Metrionotus + I. krombeini was very low in both cases (i.e., < 0.42). Both analyses also recovered different taxa as the sister group to all other mesitiine lineages: Moczariella centenaria in the unpartitioned analysis and the genus Bradepyris in the partitioned analysis.
The interpretation of topologies obtained allowed us to propose 17 nomenclatural changes: two new genera, one genus synonymy, three revalidations in species status, and 11 new specific combinations (Figs
Anaylax simplicitus Barbosa & Azevedo, 2011 by original designation.
The length of first flagellomere shorter than pedicel (#18:2), ventral half of occipital carina absent (#30:0), mesoscutellum touching metapectal-propodeal disc (#47:1), propodeal spiracle circular (#61:1), distance between distal hamuli and first hamuli more separated than others (#82:1), ventral arm of paramere of genitalia S-shaped (#104:1) are autapomorphies of Hadesmesitius. This genus has similarity with Anaylax and Clytrovorus, because they have the head, dorsal pronotal area and mesoscutum coriaceous, the median pronotal line and median mesonotal sulcus absent, and the posterior propodeal projection absent. Other characteristics also help to distinguish Hadesmesitius from Anaylax and Clytrovorus, as follows: hypopygium longer than wide and with filamentary branches, similar to Pilomesitius, Pycnomesitius, Sulcomesitius, and Zimankos; hind wing with distance between distal hamuli and first hamuli more separated than others, similar to Zimankos; and the ventral arm of paramere of genitalia S-shaped is shared with Gerbekas and Heterocoelia. Based on comparisons with the other Mesitiinae genera and mainly on its monophyly, we introduce Hadesmesitius as a new genus for Mesitiinae.
Wings subhyaline. Head: As long as wide; malar space shorter than VOL, parallel; clypeus with median lobe quadrate, median clypeal carina arched; antenna with pubescence sparse and short; pedicel fusiform, first flagellomere shorter than pedicel, flagellomeres long; eye small; frons not foveolate, with frontal carina; ocelli small; anterior ocellus posterior to supra-ocular line; dorsal half of occipital carina low, ventral half of occipital carina absent. Pronotum: Dorsal pronotal area shorter than wide, coriaceous, with humeral angle rounded, side slightly incurved, anterior margin outcurved, posterior margin straight, median pronotal line absent; mesoscutum coriaceous, median mesonotal sulcus absent, notaulus narrow; mesoscutellum touching metapectal-propodeal disc; metapectal-propodeal disc as long as its half width, metapostnotal median carina incomplete, without longitudinal ridge between metapostnotal median carina and metapostnotal-propodeal carina, posterior propodeal projection absent; spiracle shape circular; propodeal declivity coriaceous and ecarinate; lateral surface of metapectal-propodeal complex coriaceous, without carinae. Wings: Hind wing with first hamuli more separated than others. Metasoma: Dorsal and ventral region of terga III–VI polished, with sparse setae at posterior margin; hypopygium bilobate, spiculum as long as half of hypopygium, with filamentary and long branch, longer than wide, lateral margin parallel, corner angulate. Genitalia: With harpe dorsal arm shorter than ventral arm, ‘S’-shaped, and with basal margin narrow, ventral arm of harpe wide apically; cuspis with distinct arms; aedeagus slender, with apex posterior to harpe apex, apical margin rounded, lateral of margin of basal portion slightly outcurved.
The name Hadesmesitius, masculine, is a combination of the “Hades”, the Greek mythology god that has a forked weapon with the same shape of the hypopygium in this genus, which is diagnostic for the group, and the name “Mesitius”, the type genus of Mesitiinae.
United Arab Emirates.
Only the type species Anaylax simplicitus Barbosa & Azevedo, 2011 in its current combination Hadesmesitius simplicitus (Barbosa & Azevedo, 2011) comb. nov.
Incertosulcus krombeini Móczár, 1970 by original designation.
The malar space convergent (#7:0), apex of median clypeal carina [in profile] inclined (#11:2), eye very small (#21:2), pubescence of eye absent (#22:0), posterior propodeal projection wide (#64:0), number of distal hamuli of hind wing four (#81:1), hypopygium wider than long (#94:2), anterolateral hypopygial apodeme present (#100:1) were found to be autapomorphies for Brachymesitius. The type species was first described as a species of Incertosulcus. The genus was characterized by the presence or absence of a long posterior propodeal projection, making the identification dubious.
This genus shares similarities with Anaylax, Clytrovorus, and Hadesmesitius which have the head, dorsal pronotal area and mesoscutum coriaceous, and the median pronotal line and median mesonotal sulcus absent. Other characteristics distinguish Brachymesitius from Anaylax, Clytrovorus, and Hadesmesitius, including eyes very small, similar to Bradepyris and Moczariella; the anterolateral hypopygial apodeme similar to Mesitius; and the hind wing with four distal hamuli is shared with Zimankos; the presence of frontal carina and the propodeal declivity and lateral surface of metapectal-propodeal complex areolate are exclusive for Brachymesitius as diagnostic characteristics.
Wings : hyaline. Head: As long as wide; malar space as long as VOL, convergent; clypeus with median lobe rounded, median clypeal carina inclined; antenna with pubescence sparse and short; pedicel cylindrical, first flagellomere as long as pedicel, flagellomeres short; eye very small, without pubescence; frons foveolate, with frontal carina; ocelli very small; anterior ocellus crossing supra-ocular line; dorsal and ventral half of occipital carina low. Pronotum: Dorsal pronotal area shorter than wide, foveolate, with humeral angle rounded, side straight, anterior margin outcurved, posterior margin incurved, median pronotal line absent; mesoscutum coriaceous, median mesonotal sulcus absent, notaulus present and narrow; mesoscutellum not touching the metapectal-propodeal disc; metapectal-propodeal disc as long as its half width, metapostnotal median carina complete, with longitudinal ridge between metapostnotal median carina and metapostnotal-propodeal carina, posterior propodeal projection very short and thick; spiracle shape elliptical; propodeal declivity areolate, with median and lateral carinae; lateral surface of metapectal-propodeal complex areolate, without carinae. Wings: Hind wing with four distal hamuli. Metasoma dorsal and ventral region of terga III–VI polished; hypopygium bilobate, spiculum short, with lobate and short branch, wider than long, lateral margins convergent, with lateral anterior projection.
The name Brachymesitius, masculine, is a combination of the names “brachy”, from the Greek “short”, and refers to the reduced size of structures, such as eye size, flagellomeres, ocelli, length of dorsal pronotal area, posterior propodeal projection, and hypopygium, which are diagnostic for the group, and the name “Mesitius”, the type genus of Mesitiinae.
Iraq.
Incertosulcus krombeini Móczár, 1970, now Brachymesitius krombeini (Móczár, 1970) stat. rev. et comb. nov., removed from the synonym of Parvoculus indicus Kieffer, 1905.
This genus is characterized by the antenna with flagellomeres wide with pubescence dense and short, the forewing with nebulous Cu vein, the hypopygium with wide spiculum and branches lobate, and the male genitalia with parameres S-shaped (
This genus is characterized by the head as long as wide, the anteromesoscutum without median mesonotal line, the posterior propodeal projection short, and the metasomal tergum II densely punctured (
This genus is characterized by the malar space as long as vertex-ocular line, convergent anteriorly, in front view, the anteromesoscutum with median mesonotal line well impressed, the forewing with nebulous Cu and A veins, the hypopygium with branches lobate and long, and the male genitalia with dorsal paramere S-shaped, ventral paramere narrower than dorsal (
This genus is characterized by the malar space with sides parallel and as long as vertex-ocular line, the antennal pubescence dense and mid-sized (about 0.5 × flagellomeral width), the dorsal pronotal area with humeral angle projected, the hind wing with four hamuli, and the hypopygium with wide spiculum and long branches (
Although
Among genera represented by more than one terminal in parsimony analyses, five were found to be monophyletic: Astromesitius Barbosa & Azevedo, 2019, Bradepyris, Clytrovorus, Itapayos, and Mesitius; three were found to be paraphyletic: Botoryan, Metrionotus, and Pilomesitius; and eight were found to be polyphyletic: Anaylax, Gerbekas, Heterocoelia, Incertosulcus, Parvoculus, Pycnomesitius, Sulcomesitius, and Zimankos. Additionally, the paraphyly of Metrionotus and Pilomesitius and the polyphyly of Gerbekas, Heterocoelia, Pycnomesitius, Sulcomesitius, and Zimankos required new combinations (see below). Botoryan was found forming a clade nested within Zimankos and therefore we treat it as junior synonym of the latter. Anaylax simplicitus Barbosa & Azevedo, 2011 and Incertosulcus krombeini Móczár, 1970 were both recovered as distinct lineages, not clustering with other species of their respective genera. Mesitius krombeini was retrieved as closely related to the clade formed by Zimankos + Botoryan. Sulcomesitius nepalensis Móczár, 1986 clustered with species of Metrionotus. Gerbekas laosensis Móczár, 1975 was recovered as related to four species of Sulcomesitius.
The topologies obtained allowed the identification of morphological characters which potentially played important roles during the diversification of Mesitiinae, including sculpture of frons (#24); sculpture of dorsal pronotal area (#33); presence of median pronotal line (#36); presence of posterior propodeal projection (#63); length of hypopygium (#94); shape of posterior hypopygeal margin (#96); and length of hypopygium branches (#98). These characters are unique to the subfamily and allow us to hypothesize about their evolution. These hypotheses are largely based on convergent characteristics shared between Mesitiinae and Chrysidinae (Chrysididae) (Argaman, 2003).
From the 19 genera proposed for Mesitiinae, 15 exhibit roughly sculptured frons (character #24, state 1) and pronotal area (character #33, state 1), with foveolate patterns (Figs
Cryptocephalini and Clytrini (Cryptocephalinae) leaf beetles have close association with ant nests. The larval stages remain in the ant nest in a positive interaction. The cocoon brought by the beetle mother is carried by the ants into the nest to complete its development (
The median pronotal line (Character #36, state 1) characteristic of many mesitiine lineages (Fig.
Among bethylids, the posterior projections on the propodeum (Fig.
On the other hand, the musculature could indicate another adaptation associated with this structure. The muscle T1-S/T2 has its origin at the posterior corner of the metapectal-propodeal complex and inserts at anterior margin of second metasomal segment. We dissected some Mesitiinae specimens with this projection and observed that the T1-S/T2 muscle has its origin inside the projection, thus increasing the anchorage insertion point of this muscle and giving it more strength. Additionally, this muscle is related to the sternal torsion of the metasoma (
All mesitiine wasps have the second metasomal segment longer than the others, an exclusive feature for the subfamily (
There are several shapes of hypopygium exclusive to Mesitiinae, which are described by seven characters in the present analyses (Characters #94 to #100).
Clade A (Fig.
Muscles located at the base of male genitalia are responsible for movements such as protraction as well as copulation, being inserted at the anterior region of the hypopygium, including the spiculum and anterolateral apodeme. Therefore, the contraction and relaxation between the genitalia base and the spiculum provides the movement of genitalia structures. Thus, the shape and size of the spiculum have direct association with insertion of muscles in the genitalia, affecting the kind and potential of its movements that is, with more and diversified muscle insertions, structures will be capable of performing more complex movements.
The modification of the length of the hypopygial branches is associated with the deformation of the hypopygium, which results from the contraction of the muscles. More muscles inserted at a longer spiculum promote a higher degree of hypopygium deformation, hence the long branches are associated with long median indentation, providing the hypopygium with an area of deformation, giving the structure more flexibility. This is also associated with a wide spiculum. On the other hand, short branches are associated with a simple acute spiculum, since the muscle contraction provides less deformation to the hypopygium, without the need of a deformation area.
The hypopygium shape is associated with muscle insertion and hence it could provide specific functions and adaptations for each genus.
Presently, Mesitiinae are known from warm regions of the Old World, encompassing all of its four zoogeographical regions: Afrotropical (including Madagascar), Australian, Oriental, and Palearctic (Fig.
Based on this pattern, it may be that the early diversification of Mesitiinae occurred in the Palearctic region, with lineages progressively occupying the adjacent Oriental and Afrotropical regions and, later, Australia and Madagascar. Occupation of Madagascar likely occurred several times independently within the subfamily, while only a single lineage was able to reach Australia. In the future, a dated phylogeny of the family allied to the exploration of its relationships among other bethylid lineages and discoveries regarding fossil history may provide valuable evidence for a detailed approach on biogeography and diversification of Mesitiinae.
The present study is the most comprehensive cladistic treatment focusing on Mesitiinae tribal classification and character evolution, and the first to treat a large representative group and more accurate classification for each tribe.
Triglenusini were recovered as paraphyletic, and Domonkosini, Heterocoeliini and Mesitiini as polyphyletic, showing the classification in
Cladogram showing the relationships among genera of Mesitiinae (according to the taxonomic treatment adopted herein), branching support, and their geographic distributions. Colored squares indicate presence on the respective regions: AU – Australia; OR – Oriental; PL – Palearctic; AF – Afrotropical; MD – Madagascar. Position of Australomesitius is inferred based on characters mentioned in the main text. 1 Moczariella centenaria; 2 Metrionotus yarrowi; 3 Incertosulcus priesneri; 4 Sulcomesitius bicolor; 5 Gerbekas fischeri; 6 Zimankos krombeini.
DNB. planned, prepared, and designed the study. MH. and AL. supervised the study. DNB. performed the photography. DNB and MH. Performed the cladistic analyzes, AL. performed the Bayesian analyzes. DNB. described and recognized the new taxa. DNB. wrote the first draft of the manuscript. DNB., MH. and AL. discussed the results and revised the manuscript. All authors have read and approved the final version of the manuscript.
The authors have declared that no competing interests exist.
We thank the curators of the museums listed for providing the material required for this study, Géllert Púskás and Sándor Csósz for hosting DNB at
Character list
#1. Host/
0. Lepidoptera/
1. Coleoptera/
#2. Sex dimorphism/
0. absent/
1. present/
#3. Length of head/
0. longer than wide/
1. as long as wide/
2. wider than long/
#4. Shape of head [in profile]/
0. globoid [in lateral view]/
1. narrow [in lateral view]/
#5. Layout of malar space/
0. not projected/
1. projected/
#6. Length of malar space/
0. longer than VOL/
1. as long as VOL/
2. shorter than VOL/
#7. Orientation of malar space/
0. convergent anteriorly/
1. parallel/
#8. Presence of inner keel of mandible/
0. absent/
1. present/
#9. Delimitation of median lobe of clypeus/
0. not delimitated/
1. delimitated/
#10. Presence of clypeal lateral lobe/
0. absent/
1. present/
#11. Shape of apex of median clypeal carina [in profile]/
0. arched/
1. straight/
2. inclined/
#12. Shape of apex of median clypeal carina [in dorsal view]/
0. spoon-like shaped/
1. line shaped/
#13. Height of median clypeal carina/
0. below torulus/
1. above torulus/
#14. Fusion between torulus and median carina of clypeus/
0. not fused/
1. fused/
#15. Density of pubescence of antenna/
0. sparse/
1. dense/
#16. Length of pubescence of antenna/
0. short/
1. long/
2. medium/
#17. Shape of flagellomeres of antenna/
0. cylindrical shape/
1. caliciform/
#18. Length of first flagellomere of antenna [in relation to pedicel]/
0. as long as pedicel/
1. longer than pedicel/
2. shorter than pedicel/
#19. Length of flagellomeres 1–11 of antenna/
0. short/
1. long/
#20. Width of flagellomeres 1–11 of antenna/
0. slender/
1. strong/
#21. Size of eye/
0. small/
1. large/
2. very small/
#22. Pubescence of eye/
0. absent/
1. present/
#23. Contour of eye/
0. protruding/
1. in same level of head/
#24. Sculpture of frons/
0. not foveolate/
1. foveolate/
#25. Density of sculpture of frons/
0. densely foveolate/
1. sparsely foveolate/
#26. Location of anterior ocellus of ocellar triangle/
0. placed above imaginary top line of eyes/
1. placed below imaginary top line of eyes/
2. placed at imaginary mid line of eyes/
#27. Shape of hypostomal carina/
0. angled/
1. straight/
2. rounded/
#28. Anterior depression of occiput/
0. absent/
1. present/
#29. Presence of dorsal half of occipital carina/
0. absent/
1. present/
#30. Presence of ventral half of occipital carina/
0. absent/
1. present/present, but so weakly
#31. Shape of ventral half of postoccipital carina/
0. angled/
1. rounded/
#32. Length of pronotal disc/
0. shorter than wide/
1. as long as wide/
2. longer than wide/
#33. Sculpture of pronotal disc/
0. not foveolate/
1. foveolate/
#34. Density of sculpture of pronotal disc/
0. sparse/
1. dense/
#35. Presence of projection of corner of pronotal disc/
0. absent/
1. present/
#36. Presence of longitudinal pronotal furrow/
0. absent or indistinct/
1. present or distinct/
#37. Angulation anterior margin of propleuron/
0. angled/
1. straight/
2. concave/
#38. Length of mesoscutum/
0. shorter than scutellum/
1. as long as scutellum/
2. longer than scutellum/
#39. Presence of longitudinal furrow of mesoscutum/
0. absent/
1. present/
#40. Presence of notaulus of mesoscutum/
0. absent/
1. present/
#41. Orientation of notauli of mesoscutum/
0. convergent posteriorly/
1. parallel/
#42. Impression of notaulus of mesoscutum/
0. weakly impressed/
1. well impressed/
#43. Presence of parapsidal furrow of mesoscutum/
0. absent/
1. present/
#44. Presence of transcutal articulation/
0. inconspicuous/
1. conspicuous/
#45. Impression of scutoscutellar sulcus/
0. inconspicuous/
1. conspicuous/
#46. Presence of connection between scutellar groove and axilla/
0. absent/
1. present/
#47. Extension of scutellum/
0. not touching propodeal disc/
1. touching propodeal disc/
#48. Length of propodeal disc/
0. shorter than half width of propodeal disc/
1. as long as half width of propodeal disc/
2. longer than half width of propodeal disc/
#49. Metapostnotal depression/
0. absent/
1. present/
#50. Connection between central depression and triangular lateral depression of propodeal disc/
0. absent/
1. present/
#51. Presence of median carina of propodeal disc/
0. absent/
1. present/
#52. Extension of median carina of propodeal disc/
0. incomplete/
1. complete/
#53. Fusion between sublateral and inner discal carina of propodeal disc/
0. absent/
1. present/
#54. Presence of inner discal carina of propodeal disc/
0. absent/
1. present/
#55. Extension of inner discal carina of propodeal disc/
0. incomplete/
1. complete/
#56. Orientation of inner discal carina of propodeal disc/
0. parallel with median carina/
1. not parallel with median carina/
#57. Presence of sublateral carina of propodeal disc/
0. absent/
1. present/
#58. Presence of lateral carina of propodeal disc/
0. absent/
1. present/
#59. Presence of posterior carina of propodeal disc/
0. absent/
1. present/
#60. Extension of posterior carina of propodeal disc/
0. incomplete medially/
1. complete/
#61. Shape of propodeal spiracle/
0. elliptical/
1. circular/
#62. Location of propodeal spiracle/
0. placed at dorsal surface of propodeum/
1. placed at lateral surface of propodeum/
#63. Presence of posterior spine of propodeum/
0. absent/
1. present/
#64. Width of posterior spine of propodeum/
0. thick/
1. slender/
#65. Presence of median carina of declivity of propodeum/
0. absent/
1. present/
#66. Presence of lateral carina of declivity of propodeum/
0. absent/
1. present/
#67. Presence of superior carina of side of propodeum/
0. absent/
1. present/
#68. Mesopleuron foveae distinction/
0. absent/
1. present/
#69. Presence of posterior carina of side of propodeum/
0. absent/
1. present/
#70. Fusion between subtegular fovea and episternal furrow of mesopleuron/
0. not fused/
1. fused/
#71. Presence of transverse furrow of mesopleuron/
0. absent/
1. present/
#72. Presence of diastema between tegula and mesoscutum/
0. absent/
1. present/
#73. Presence of costal vein of forewing/
0. absent/
1. present/
#74. Shape of transverse median vein of forewing/
0. bi-angulate/
1. rounded/
#75. Presence of nebulous cubital vein of forewing/
0. absent/
1. present/
#76. Presence of nebulous anal vein of forewing/
0. absent/
1. present/
#77. Presence of distal fusion among costal and subcostal vein before stigma of forewing/
0. absent/
1. present/
#78. Presence of proximal hamuli of hind wing/
0. absent/
1. present/
#79. Number of proximal hamuli of hind wing/
0. one/
1. two/
2. three/
3. six/
#80. Presence of distal hamuli of hind wing/
0. absent/
1. present/
#81. Number of distal hamuli of hind wing/
0. three/
1. four/
2. one/
3. five/
#82. Distance between distal hamuli of hind wing/
0. separated each other by uniform space/
1. first hamuli more separated than others/
#83. Presence of dorsal process of hind coxa/
0. absent/
1. present/
#84. Shape of tarsal claw/
0. one tooth/
1. two teeth/
2. three teeth/
#85. Presence of constriction between tergum I and tergum II of metasoma/
0. absent/
1. present/
#86. Presence of ventral sculpture at tergum I of metasoma/
0. absent/
1. present/
#87. Presence of lateral ventral lap of tergum I of metasoma/
0. absent/
1. present/
#88. Presence of dorsal setae at tergum I of metasoma/
0. absent/
1. present/
#89. Length of tergum II of metasoma/
0. as long as others/
1. longer than others/
#90. Type of dorsal texture of tergum II of metasoma/
0. polished/
1. coriaceous/
#91. Presence of dorsal sculpture of tergum II of metasoma/
0. absent/
1. present/
#92. Presence of ventral sculpture of tergum II of metasoma/
0. absent/
1. present/
#93. Type of ventral texture of tergum II of metasoma/
0. polished/
1. coriaceous/
#94. Length of hypopygium/
0. longer than wide/
1. as long as wide/
2. wider than long/
#95. Width of median anterior process of hypopygium/
0. acute/
1. wide/
#96. Shape of posterior margin of hypopygium/
0. simple/
1. bilobed
#97. Shape of branches of posterior margin of hypopygium/
0. lobose/
1. filamentary/
#98. Length of branches of posterior margin of hypopygium/
0. short/
1. long/
#99. Orientation of lateral margin of hypopygium/
0. parallel/
1. convergent/
#100. Presence of lateral anterior projection of hypopygium/
0. absent/
1. present/
#101. Presence of projection of genital ring/
0. absent/
1. present/
#102. Number of paramere arms of genitalia/
0. one/
1. two/
#103. Shape of dorsal arm of paramere of genitalia/
0. “S” shaped/
1. club-shaped/
2. filamentary shaped/
#104. Shape of ventral arm of paramere of genitalia/
0. club-shaped/
1. “S” shaped/
#105. Presence of fusion between basiparamere and basivolsella of genitalia/
0. absent/
1. present/
#106. Number of arms of cuspis of genitalia/
0. one/
1. two/
#107. Arms of cuspis of genitalia/
0. distinct/
1. hardly distinct/
#108. Width of aedeagus of genitalia/
0. slender/
1. wide/
#109. Length of aedeagus of genitalia/
0. not reaching paramere apex/
1. surpassing paramere apex/
2. aligned with paramere apex/
#110. Apex shape of aedeagus of genitalia/
0. rounded/
1. truncate/
2. angled/
#111. Presence of apical sickle process of aedeagus of genitalia/
0. absent/
1. present/
#112. Shape of lateral margin of aedeagus basal portion of genitalia/
0. convex/
1. straight/
Table S1
Data type: .xlsx
Explanation note: Matrix of coded state characters related to e ach taxon.
Table S2
Data type: .xlsx
Explanation note: Mesitiinae genera, their number of species and type species of in its original combination.
Figure S1
Data type: .pdf
Explanation note: Symmetric resampling of cladistic analysis tree.
Figure S2
Data type: .pdf
Explanation note: Bayesian tre e based on partitioned parameters, indicating posterior probably for each node.
Figure S3
Data type: .pdf
Explanation note: Bayesian tree based on unpartitioned parameters, indicating posterior probably for each node.