Legs orange or yellow. T1 black; T2—T4 orange with narrow, black end margins, T2 additionally with irregular brown markings; T5—T7 orange with black mark medially. Ovipositor sheaths brown. Head roundish, no details discernible. Antennae almost complete, with about 20 flagellomeres; most flagellomeres rather stout, median ones about 1. Pronotum moderately long with epicnemical carina extending at least to mid-height of pronotum. Sternaulus-area unclear. Propodeum with pleural carina, split where lateral longitudinal carina might be, and traces of probably middle part of basal transverse carina; broken off where posterior transverse carina might be; certainly with several delimited areas; submetapleural carina complete.
Pterostigma around 4. Hind wing 1Rs around 1. Fore and hind leg weakly indicated, hind leg seems rather slender, maybe with two spurs on hind tibia. Metasoma probably slightly compressed. T1 broadly attached but still diverging posteriorly, 1. T2—T4 wider than long, with hind margin shiny black. T5—T7 rather shorter. Last sternites visible, hypopygium short and straight. This genus cannot be placed in a single ichneumonid subfamily. However, most Tryphoninae and Banchinae do not have a transverse T2, and Banchinae only rarely have such long carinae on T1.
Since there is no extant genus in any of the three subfamilies that would match our fossil, also considering the very distinct colour pattern on the mesosoma, we define a new genus with uncertain subfamily placement. Mesoscutum with distinct colour pattern—ground colour black, orange-brown along extension of notauli on posterior half, sides of scutellum, postscutellum and possibly end of propodeum; very heavily punctured with very deep notauli meeting at mid-length.
Prescutellar groove with strong carinulae; axillar troughs of mesonotum and metanotum well-delimited, the former with transverse carinulae. T1 short and stout, evenly tapers towards front; with strong and evenly converging dorsal longitudinal carinae and strong and dense punctures on most of surface and diagonal rugae on sides of longitudinal carinae on basal half.
T2 and following tergites transverse and heavily punctured. Metasoma fusiform. Dedicated to a dear friend and one of the most enthusiastic and gifted biologists I TS have ever met. Like the fossil, Djordje Markovic is a remarkable specimen, seeking—and hopefully one day finding—his position in the scientific system. Holotype: SF MeI , female? Dorsal aspect of body with fore and hind wings, one complete antenna, parts of mid and hind legs; without ovipositor. Orange along extension of notauli on posterior half of mesoscutum, sides of scutellum, postscutellum and possibly end of propodeum.
Hind coxa, apex of tibia and tarsi black, hind tibia dark brown; hind femur, mid and front legs orange-brown. T1 black, remaining tergites orange-brown, T2 with four black lateral markings. Head with vertex distinctly punctured, gena short; occipital carina strong and evenly convex.
Antennae 1. Shape of scape and pedicel indistinct. Mesoscutum very heavily punctured on smooth and polished background; notauli very deep, parallel on anterior half of mesoscutum then either strongly converging around middle or extending parallel until end of mesoscutum; carinae along lateral margin uninterrupted from tegula to scutellum. Prescutellar groove with strong carinulae; Axillar troughs of mesonotum and metanotum well-delimited, the former with transverse carinulae. Propodeum quite short, heavily punctured, with at least basal portion of median longitudinal carinae and anterior transverse carina present; pleural carina probably present as well.
Wing veins somewhat distorted. Pterostigma around 3. Hind femur about 3. Hind tibia robust, becoming thicker towards apex, 4. Hind tarsomere very elongate and robust and hairy.
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Metasoma quite fusiform. T1 about 1. T2 and following tergites more than 2x wider than long, heavily punctured. No ovipositor visible; strong fusiform shape of metasoma suggests female. The Messel Pit is known as a locality with unique fossil preservation and many remarkable fossil findings [ 18 ]. Here we described the first ichneumonids from the locality, which unsurprisingly show good preservation that in most cases allows their confident placement in recent subfamilies and genera.
By far the largest number of described Eocene ichneumonid fossils have been placed in the subfamily Pimplinae [ 6 ], but the taxonomic position of many of them is actually uncertain and often based on superficial resemblance to extant representatives of the group [ 7 , 64 — 66 ]. The uncertainty partially comes from the plesiomorphic nature of many pimpline body and wing venation character states, such as the stout tergite 1 with dorsal longitudinal carinae and the quadrate areolet [ 51 , 67 ]. Even the subfamily placement of extant pimpline males can be difficult without having corresponding females [ 52 , 68 ].
Moreover, the suggested synapomorphies for the subfamily, e. We therefore consider the here described Scambus fossilobus as the first unequivocal record of the subfamily Pimplinae from the Eocene. The placement of our fossil in the subfamily is evidenced by the presence of lobed claws, a character state that supports the monophyly of at least a subset of genera in the subfamily Pimplinae [ 51 ] and has never been documented before in a fossil. Rhyssella vera represents the oldest unambiguous fossil of the subfamily Rhyssinae.
Three other rhyssine fossils have been described previously: Rhyssa antiqua Heer [ 69 ] from the Early Miocene locality Radoboj in Croatia, R.
However, there are a couple of genera in the related subfamilies Pimplinae and Poemeninae that can have very similar wings. Moreover, the wing venation pattern in R. III, Fig 18 in Heer [ 69 ]. In the case of R. However, Brues [ 70 ] also reported an areolated propodeum in the fossil, which does not occur in extant Rhyssinae. In any case, this fossil is from the latest Eocene and thus considerably younger than the here described R.
The newly described Trigonator macrocheirus is the first certain labenine fossil and the only one from Eurasia Fig 9. The only previously described fossil of this subfamily, Albertocryptus dossenus , shows some untypical characters when compared to recent representatives of the subfamily, such as small size, an elongate scape and short antenna with only 19 flagellomeres, a very wide and short pterostigma, and a strongly humped T1 in the shape of a petiole even though a similar state occurs in the genus Labium in the tribe Groteini.
Therefore, even though they placed the fossil in the recent tribe Poecilocryptini, McKellar et al. In contrast, T. Labenine today show a primarily Gondwanan distribution, but they are absent from African, India and Madagascar and present in the southern parts of North America. Stars indicate localities of the labenine fossil record, which suggest a more global distribution of the subfamily in the past; the approximate ages of the fossils are given in brackets.
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But it supports, together with the McKellar et al. However, the oldest clear ichneumonid fossils from the extinct subfamily Palaeoichneumoninae are only between and Ma old Early Cretaceous [ 4 , 8 ], and up until now, there is no fossil evidence that any of the extant subfamilies was already present during that period. The available age estimates for the family Ichneumonidae, based on phylogenetic methods, are around Ma [ 75 ] and Ma [ 76 ], but they stem from dating studies where ichneumonids were not a focal group and are thus rather unprecise due to low taxon sampling.
Therefore, a dated phylogeny of the family with age estimates for the subfamilies is essential to resolve the timing of the labenine radiation. Alternatively, the diversification of Labeninae occurred during the Early Cretaceous and the group dispersed either via existing land bridges [ 77 — 79 ], as suggested for many insect groups [ 80 — 84 ], or by a long-distance dispersal.
Ichneumonid wasps are today and have probably always been very mobile organisms [ 85 — 88 ] with comparatively low levels of endemism [ 1 , 2 , 89 , 90 ]. The dispersal abilities of Labeninae thus might be high enough that trans-oceanic dispersal is a real option, which would already have erased any signal of past distributional patterns, especially those as old as Cretaceous vicariance events.
Nevertheless, a more complete fossil record of labenine is needed to infer the dispersal pathways. A link could possibly be made between Labeninae and the extinct, Late Cretaceous subfamily Labenopimplinae. Labenopimpline are a morphologically heterogeneous assemblage of ichneumonid fossil genera with mixed characteristics also found in modern Labeninae and Pimplinae [ 12 , 91 , 92 ].
They have been described from Russia [ 12 , 92 ], Canada [ 13 ] and Botswana [ 91 ], which reflects their wide distribution range during the Late Cretaceous see checklist in Li et al. If the subfamily were indeed an ancestral lineage to Labeninae as suggested by Kopylov [ 12 ], the hypothesis of an initial wide distribution of Labeninae linked to high dispersal abilities would be supported.
However, the two subfamilies clearly differ in the shape of T1 and the attachment point of the metasoma, and their relationships remain rather unclear; thus any conclusion would be speculative. Xanthopimpla is the largest genus of the subfamily Pimplinae, with a wide distribution in the tropics and subtropics Fig 10 [ 2 , 57 , 58 ].
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The species richness is highest in the Indo-Australian and Oriental regions, while a few species can be found in Africa and in the Neotropics [ 57 , 58 ]. Because of its current distribution, the genus is considered to indicate a warm-temperate and subtropical climate [ 59 ]. The two newly described Xanthopimpla species support this observation. In the Eocene, the climate around former Lake Messel was warm and humid.
The vegetation around the former Lake Messel seems to be have been a multilevel canopy forest with a large tropical-paratropical component, and large part of the fauna consisted of subtropical and tropical elements [ 25 , 34 ]. Xanthopimpla is nowadays widely distributed in tropical and subtropical areas of the world.
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Striped area indicates uncertain occurrence of the genus in France, while stars indicate localities of the fossil record; approximate age of the fossils is given in brackets. The taxonomic placement of incompletely preserved fossils can be very challenging due to a lack or uncertain interpretation of visible synapomorphies [ 10 ]. However, fossils from the Messel Pit often show remarkable preservation with many morphological details retained [ 11 , 12 ]. Although this was the case in the newly described ichneumonid fossils, we were not able to firmly place two of them.
Both Polyhelictes bipolarus and Mesornatus markovici show character state combinations that occur in more than one extant ichneumonid subfamily. This issue has already been reported for ichneumonid fossils and is mostly attributed to the high rates of homoplasy observed in these parasitoids [ 7 , 13 ]. A phylogenetic analysis with combined morphological data from extant and fossil ichneumonids could possibly resolve taxonomic position of such taxa—especially of P. The unique fossil preservation in the Eocene Messel Pit allows us to firmly place several ichneumonid fossils in extant subfamilies and genera.
As a result, we report the oldest unequivocal representatives of the subfamilies Pimplinae, Rhyssinae and Labeninae and of the genus Xanthopimpla which was up till now only known from the latest Eocene.