Corresponding author: Diego Nunes Barbosa (
We present the first phylogenetic hypothesis for
Barbosa DN, Hermes MG, Lepeco A (2022) Phylogeny of Mesitiinae (Hymenoptera: Bethylidae): assessing their classification, character evolution and diversification. Arthropod Systematics & Phylogeny 80: 603–625.
The aculeate family
The monophyly of the
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
Table
A total of 112 characters (Appendix
Specimens included in the phylogenetic analysis
Taxa | Specimen | Zoogeographic region | Repository |
---|---|---|---|
|
|||
|
|||
Allotype | Madagascar |
|
|
Allotype | Ethiopic |
|
|
Holotype | Madagascar |
|
|
Paratype | Ethiopic, Oriental |
|
|
Paratype | Oriental |
|
|
Paratype | Oriental |
|
|
Paratype | Oriental |
|
|
|
|||
Holotype | Oriental |
|
|
Voucher | Palaearctic |
|
|
Holotype | Oriental |
|
|
Allotype | Palaearctic |
|
|
Holotype | Ethiopic |
|
|
Voucher | Palaearctic |
|
|
Voucher | Palaearctic |
|
|
Voucher | Palaearctic |
|
|
Paratype | Palaearctic |
|
|
Neótipo | Palaearctic |
|
|
Paratype | Ethiopic |
|
|
Allotype | Ethiopic | BMNH | |
Voucher | Ethiopic |
|
|
Paratype | Oriental |
|
|
Paratype | Oriental |
|
|
Holotype | Oriental |
|
|
Voucher | Oriental |
|
|
Paratype | Oriental |
|
|
Paratype | Oriental |
|
|
|
|||
Holotype | Madagascar |
|
|
Holotype | Madagascar |
|
|
Holotype | Ethiopic |
|
|
Holotype | Ethiopic |
|
|
Voucher | Ethiopic |
|
|
Holotype | Oriental |
|
|
Holotype | Palaearctic |
|
|
Allotype | Palaearctic |
|
|
Holotype | Madagascar |
|
|
Holotype | Madagascar |
|
|
Allotype | Ethiopic |
|
|
Paratype | Ethiopic |
|
|
Holotype | Palaearctic | BMNH | |
Holotype | Palaearctic |
|
|
Holotype | Palaearctic |
|
|
Holotype | Ethiopic | CNCI | |
Paratype | Oriental |
|
|
Voucher | Oriental |
|
|
Holotype | Madagascar |
|
|
Voucher | Oriental |
|
|
Holotype | Oriental |
|
|
Holotype | Palaearctic | ZMB | |
Holotype | Oriental |
|
|
Holotype | Ethiopic |
|
|
Paratype | Ethiopic |
|
|
Paratype | Ethiopic |
|
|
Paratype | Oriental |
|
|
Paratype | Ethiopic |
|
|
Holotype | Oriental |
|
|
|
|||
Voucher | Palaearctic |
|
|
Allotype | Palaearctic | BMNH | |
Allotype | Palaearctic |
|
|
Voucher | Palaearctic |
|
|
Holotype | Palaearctic |
|
|
Holotype | Ethiopic |
|
|
|
|||
Voucher | Palaearctic |
|
|
Voucher | Holarctic, Neotropical |
|
|
Paratype | Neotropical |
|
|
Paratype | Neotropical |
|
|
Paratype | Oriental |
|
|
Paratype | Neotropical |
|
|
Voucher | Holarctic, Neotropical |
|
|
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 (
Characters and character states.
Characters and character states.
Part of the cladogram obtained with parsimony under implied weighting (
Twenty-four characters were found as synapomorphies for the subfamily, 14 of them are exclusive transformations for
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.
Part of the cladogram obtained with parsimony under implied weighting (
New nomina, nomenclatural acts and changes in combination in this study.
|
|
|
— | — |
|
— | — |
|
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
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
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
Part of the cladogram obtained with parsimony under implied weighting (
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
Part of the cladogram obtained with parsimony under implied weighting (
The name
United Arab Emirates.
Only the type species
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
This genus shares similarities with
The name
Iraq.
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:
The topologies obtained allowed the identification of morphological characters which potentially played important roles during the diversification of
From the 19 genera proposed for
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
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
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,
Based on this pattern, it may be that the early diversification of
The present study is the most comprehensive cladistic treatment focusing on
Cladogram showing the relationships among genera of
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
#1. Host/
0.
1.
#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
1. as long as
2. shorter than
#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
Table S2
Figure S1
Figure S2
Figure S3