A new fossil subfamily of Bethylidae (Hymenoptera) from the Early Cretaceous Lebanese amber and its phylogenetic position

A New Bethylid Wasp in Lebanese Early Cretaceous Amber (Hymenoptera: Chrysidoidea), with Comments on other Mesozoic Taxa

“…Photomicrographs were taken with a Canon EOS 7D digital camera attached to an Infinity K-2 long-distance microscope lens, while line illustrations were prepared with the aid of a camera lucida attached to the aforementioned stereomicroscope. Morphological terminology and descriptions follow those generally used elsewhere in bethylid studies (e.g., Azevedo and Azar, 2012;Alencar and Azevedo, 2013;Ortega-Blanco and Engel, 2013;McKellar and Engel, 2014). The descriptions are provided in the context of providing fundamental character data for understanding biodiversity patterns and ultimately permitting the development and testing of evolutionary hypotheses (e.g., Grimaldi and Engel, 2007 group remains to be discovered); compound eyes well developed; ocelli present; mandibles short and thick; maxillary palpus with six palpomeres; labial palpus with four palpomeres (known only in Holopsenella).…”

“…The result of these investigations have been the elimination of past tribal classifications and the reduction of the number of subfamilies to Bethylinae, Scleroderminae, Pristocerinae, Mesitiinae, and Epyrinae. Azevedo and Azar (2012) added to this system a fifth subfamily, Lancepyrinae, based on an extinct genus in Lebanese amber. Interestingly, the Lancepyrinae were not basal in the preliminary phylogenetic analysis, suggesting that bethylids were already well established by the Barremian (Azevedo and Azar, 2012).…”

“…Azevedo and Azar (2012) added to this system a fifth subfamily, Lancepyrinae, based on an extinct genus in Lebanese amber. Interestingly, the Lancepyrinae were not basal in the preliminary phylogenetic analysis, suggesting that bethylids were already well established by the Barremian (Azevedo and Azar, 2012).…”

“…While numerous bethylid species are known from Paleogene and Neogene deposits (e.g., Azevedo and Azar, 2012;Barbosa et al, 2013), there are comparatively few published species from the Mesozoic although abundant unpublished material is known from New Jersey (Engel, unpublished data) and French ambers (Perrichot and Nel, 2008). Eleven species assigned to Bethylidae have been described from the Cretaceous, and in fossiliferous resins from the Barremian of Lebanon (Azevedo and Azar, 2012), Albian of Spain (Ortega-Blanco and , Albian-Cenomanian of Myanmar (Cockerell, 1917a(Cockerell, , 1917b(Cockerell, , 1920, Santonian of Taimyria (Evans, 1973), and Campanian of Canada Engel, 2012, 2014).…”

“…Eleven species assigned to Bethylidae have been described from the Cretaceous, and in fossiliferous resins from the Barremian of Lebanon (Azevedo and Azar, 2012), Albian of Spain (Ortega-Blanco and , Albian-Cenomanian of Myanmar (Cockerell, 1917a(Cockerell, , 1917b(Cockerell, , 1920, Santonian of Taimyria (Evans, 1973), and Campanian of Canada Engel, 2012, 2014). A 12th species, Cretabythus sibiricus Evans in Taimyrian amber, was assigned with hesitation to Scolebythidae (Evans, 1973: refer to Engel andEngel et al, 2013a, for discussion of Cretaceous scolebythids), but Carpenter (1986) considered the placement suspect and Rasnitsyn (1988) emphasized further details that indicate it to be a primitive bethylid, and we tend to concur with this conclusion (see below).…”