From hell's heart I stab at thee! A determined approach towards a monophyletic Pteromalidae and reclassification of Chalcidoidea (Hymenoptera)
Author
Burks, Roger
https://orcid.org/0000-0003-3032-7939
Department of Entomology, University of California Riverside, Riverside, CA, USA
burks.roger@gmail.com
Author
Mitroiu, Mircea-Dan
https://orcid.org/0000-0003-1368-7721
Faculty of Biology, Alexandru Ioan Cuza University, Iasi, Romania
Author
Fusu, Lucian
https://orcid.org/0000-0003-0819-026X
Faculty of Biology, Alexandru Ioan Cuza University, Iasi, Romania
Author
Heraty, John M.
https://orcid.org/0000-0002-9246-5651
Department of Entomology, University of California Riverside, Riverside, CA, USA
Author
Jansta, Petr
https://orcid.org/0000-0001-6409-3603
Department of Zoology, Faculty of Science, Charles University, Prague, Czech Republic & Department of Entomology, State Museum of Natural History, Stuttgart, Germany
Author
Heydon, Steve
Bohart Museum of Entomology, University of California, Davis, CA, 95616, USA
Author
Papilloud, Natalie Dale-Skey
https://orcid.org/0000-0001-7582-0386
Insects Division, Natural History Museum, London, UK
Author
Peters, Ralph S.
Zoologisches Forschungsmuseum Alexander Koenig, Leibniz Institute for the Analysis of Biodiversity Change, Bonn, Germany
Author
Tselikh, Ekaterina V.
https://orcid.org/0000-0002-9184-043X
Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia
Author
Woolley, James B.
Department of Entomology, Texas A & M University, College Station, TX, USA
Author
van Noort, Simon
https://orcid.org/0000-0001-6930-9741
Research and Exhibitions Department, South African Museum, Iziko Museums of South Africa, PO Box 61, Cape Town 8000 South Africa & Department of Biological Sciences, University of Cape Town, Private Bag, Rondebosch, 7701, South Africa
Author
Baur, Hannes
https://orcid.org/0000-0003-1360-3487
Department of Invertebrates, Natural History Museum Bern, Bern, Switzerland & Institute of Ecology and Evolution, University of Bern, Bern, Switzerland
Author
Cruaud, Astrid
https://orcid.org/0000-0001-8932-4199
CBGP, INRAE, CIRAD, IRD, Montpellier SupAgro, University of Montpellier, Montpellier, France
Author
Darling, Christopher
Department of Natural History, Royal Ontario Museum, Toronto, ON, M 5 S 2 C 6, Canada & Department of Ecology and Evolutionary Biology, University of Toronto, Toronto, ON, M 5 S 1 A 1, Canada
Author
Haas, Michael
https://orcid.org/0000-0001-6869-6698
Department of Entomology, State Museum of Natural History, Stuttgart, Germany
Author
Hanson, Paul
Escuela de Biologia, Universidad de Costa Rica, San Pedro de Montes de Oca, San Jose 11501 - 2060, Costa Rica
Author
Krogmann, Lars
https://orcid.org/0000-0002-3724-1735
Department of Entomology, State Museum of Natural History, Stuttgart, Germany & Institute of Biology, Biological Systematics (190 n) University of Hohenheim, Stuttgart, Germany
Author
Rasplus, Jean-Yves
https://orcid.org/0000-0001-8614-6665
CBGP, INRAE, CIRAD, IRD, Montpellier SupAgro, University of Montpellier, Montpellier, France
text
Journal of Hymenoptera Research
2022
2022-12-20
94
13
88
http://dx.doi.org/10.3897/jhr.94.94263
journal article
http://dx.doi.org/10.3897/jhr.94.94263
1314-2607-94-13
6CB807239A47403FABEC9AF8AE7F417F
ADCFB8021287566FB2D7E8A8711D5CAE
Eusandalinae Fusu, new subfamily
Type genus.
Eusandalum
Ratzeburg, 1852.
Diagnosis.
Antennal flagellum with 9 flagellomeres, clava undivided. Eyes ventrally divergent. Clypeus with truncate apical margin. Labrum subquadrate, exposed. Subforaminal bridge with median area flanked by elongate posterior tentorial attachments; postgenal bridge externally separates the lower tentorial bridge from the hypostomal carina; postgenal groove and postgenal lamina absent. Anterolateral mesoscutal corners projecting shoulder-like on either side of the pronotum (a feature shared with
Calosotinae
sensu stricto
). Notauli superficial and convergent, and except for
Archaeopelma
, ending about halfway across mesoscutum (Fig.
112
). Axilla approximated (
Archaeopelma
Gibson and
Paraeusandalum
Gibson) or widely separated (
Eusandalum
Ratzeburg,
Licrooides
Gibson and
Pentacladia
Westwood) medially. Axillular groove or carina absent. Frenum absent. Acropleuron expanded and forming the largest surface of the mesopleuron, either comparatively small and not reaching metapleuron and metacoxa (
Archaeopelma
,
Licrooides
) or occupying most of the visible part of the mesopleuron and extending to the metapleuron (the other three genera). All legs with 5 tarsomeres; protibial spur stout and curved; basitarsal comb longitudinal; ventral membranous area anterior to mesocoxal attachment present; mesotibial spur stout except only slightly thickened in
Archaeopelma
. Fore leg with protibial dorsal spicules (except in
Licrooides
). Mesotarsus almost never with row of pegs along both sides, the exception being
Licrooides
: with spine-like setae on both antero- and posteroventral margins (
Archaeopelma
) or a row of pegs on posteroventral margin and a row of spine-like setae along anteroventral margin (remaining three genera). In
Licrooides
there are robust spines on both margins that almost appear as pegs. Metasoma in females with separate Mt8 and Mt9, hence without syntergum (
Eusandalum
) or with Mt8 and Mt9 fused but delimited by a transverse suture between cerci (
Archaeopelma
) or below each cercus (remaining genera; Fig.
113
). Sexual dimorphism reduced, limited mainly to primary sexual features and antennal structure.
Discussion.
Eusandalinae
new subfamily were treated until now as part of
Calosotinae
, however in next-generation molecular analyses (Cruaud et al., submitted) they were never recovered as monophyletic with the other
Calosotinae
. Instead, a reduced group of
Calosotinae
that includes
Balcha
Walker,
Calosota
Curtis and
Tanythorax
Gibson (
Calosotinae
sensu stricto
) are sister to
Heydenia
(
Heydeniidae
) in all final analyses and closer to
Eupelminae
than to
Eusandalinae
.
Eusandalinae
are part of the same large clade containing also the
Eupelminae
and
Calosotinae
, but the three
Eupelmidae
subfamilies never form a monophyletic group since the clade also includes
Ditropinotellinae
,
Heydeniidae
and
Solenurinae
;
Eusandalinae
are the basal group. Beside the three genera included in these molecular analyses (Cruaud et al., submitted) we also include in
Eusandalinae
Archaeopelma
and
Licrooides
based on a UCE analysis with a larger sampling (unpublished data). A possibly biphyletic
Calosotinae
sensu lato
was also recognised by
Gibson (1989)
, however with
Licrooides
hypothesized as closer to
Calosota
and allied genera and not to
Eusandalum
, and
Archaeopelma
as the most basal lineage of all
Calosotinae
. From the morphologically close
Calosotinae
sensu stricto
, the
Eusandalinae
differ mainly in having an undivided clava, convergent notauli, scutellum without axillular groove or carina, mesotarsus almost never with two fully developed rows of pegs and a metasoma with incompletely fused or independent Mt8 and Mt9. In
Calosotinae
: clava with three clavomeres, notauli present as paramedially parallel lines, axillular groove present and continuing seamlessly with the scuto-scutellar suture and as a result scutellum with a carinated groove extending from base to apex, mesotarsus with a row of pegs on either side, and Mt8 and Mt9 completely fused to form the syntergum. However, all the characteristic features of
Eusandalinae
are either plesiomorphic or homoplastic. A thorough comparative analysis of all these characters can be found in
Gibson (1989)
.