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The fossils originally assigned to the family Hydrophilidae (Coleoptera, Polyphaga) from the late Miocene locality of Öhningen (southern Germany) are revised. Nine hydrophilid species are recognized, most of them representing the tribe Hydrophilini. Five species are reliably assigned to genera: Hydrochara noachica (Heer, 1847), n. comb. (=Hydrophil...
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Context 1
... braunii was described according to the single specimen deposited in ''Carlsruher Sammlung.'' We were not able to localize any historical collection of fossil insects in Karlsruhe (Germany), and therefore consider the holotype as lost. In order to fix the identity of the species, we therefore designate the specimen identified as H. braunii by O. Heer and illustrated by Heer (1862, pl. 4, fig. 14) as the ...
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... presence of clear frontal sutures arising closely before eyes clearly assigns the following fossils to the Hydro- philidae, which is also supported by the preserved morphology as it corresponds to hydrophilids in all aspects. Within Hydro- philidae, the large body size, absence of sutural stria and longitudinally carinate prosternum clearly assign the species into the Hydrophilini. Within Hydrophilini, the presence of the wide emargination of the anterior margin of the clypeus excludes the genera Sternolophus, Tropisternus, Protistolophus, and the simply carinate prosternum excludes Tropisternus and Hydro- philus. Hence, the above characters would assign the fossils to the genera Hydrobiomorpha or Brownephilus that may be only distinguished by the presence/absence of a pair of pits on labrum and the morphology of the male genitalia. The labral pits are actually clearly present at least in one specimen (Fig. 3.11) and seem to be present also in specimen ETHZ NO-Z51.3-39-4:4- Pl.I147b (holotype of H. heeri n. sp.), which clearly indicates that the genus Hydrobiomorpha is really present in¨Ohningenin¨ in¨Ohningen deposits. The preservation of other fossils with excised clypeus is rather poor in the labral region, causing that the presence or absence of the pits cannot be evaluated. In absence of additional contradict- ing characters, we consider also these remaining fossils to belong FIGURE 1-Scatter plots of body measurements of the examined specimens assigned to the medium-sized Hydrophilini (for measurements of isolated elytra see Table 2, for large-sized Hydrophilini see Fig. 4). 1, plot based on the proportions of the pronotum; 2, plot based on the proportions of the ...
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... of species in¨Ohningenin¨ in¨Ohningen deposits.-Heer (1847, 1862) diagnosed his species largely by the proportions of elytra and the shape of lateral margin on the pronotum. The latter character is only preserved in few exceptionally preserved specimens, and several more complete specimens were moreover lost. Examina- tion of all available fossils showed that characters on which the species-level taxonomy of Hydrophilus is based are not preserved. Hence, we used the morphometric data of the elytra to estimate the number of morphospecies present in¨Ohningenin¨ in¨Ohningen deposits. Elytral dimensions of¨Ohningenof¨ of¨Ohningen specimens are highly variable (39.6% for elytral length and 41.0% for elytral width). The maximal intraspecific variation of body dimensions reaches 30% for modern species of Hydrophilus, which suggests that at least two distinct species were probably present in¨Ohningenin¨ in¨Ohningen. Two narrowly separated groups are also seen on the scatter plot of elytral dimensions (Fig. 4). Due to the absence of additional data we use the gap between both groups to formally delimit two morphospecies, each of them not overlapping 30% variation of Hydrophilus knorrii, no material studied. The specimen should be deposited in ''Carlsruher Sammlung'' but we were not able to localize any historical collection of fossil insects in Karlsruhe (Germany). For that reason, we consider the only known specimen as lost. Proposed synonymy is based on the figure and the measurements given in the original description. Fig. 4 of the original description]. Paralectotypes, piece only (ETHZ NO-Z51.3-39-4:4-Pl.I143); incomplete specimen with partly preserved parts of head, pronotum and anterior legs, placed in a wooden box labeled ''Pl. I. 143'' [corresponds to plate 4, fig. 2 of the original description]; piece and counterpiece (ETHZ NO-Z51.3-39-4:4- Pl.I144), partly preserved pair of posterior legs, larger piece broken in two parts, both labeled ''Hydrophilus / spectabilis H. 144'' [corresponds with plate IV, Fig. 7
fig. 15 of the original ...
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... of species in¨Ohningenin¨ in¨Ohningen deposits.-Heer (1847, 1862) diagnosed his species largely by the proportions of elytra and the shape of lateral margin on the pronotum. The latter character is only preserved in few exceptionally preserved specimens, and several more complete specimens were moreover lost. Examina- tion of all available fossils showed that characters on which the species-level taxonomy of Hydrophilus is based are not preserved. Hence, we used the morphometric data of the elytra to estimate the number of morphospecies present in¨Ohningenin¨ in¨Ohningen deposits. Elytral dimensions of¨Ohningenof¨ of¨Ohningen specimens are highly variable (39.6% for elytral length and 41.0% for elytral width). The maximal intraspecific variation of body dimensions reaches 30% for modern species of Hydrophilus, which suggests that at least two distinct species were probably present in¨Ohningenin¨ in¨Ohningen. Two narrowly separated groups are also seen on the scatter plot of elytral dimensions (Fig. 4). Due to the absence of additional data we use the gap between both groups to formally delimit two morphospecies, each of them not overlapping 30% variation of Hydrophilus knorrii, no material studied. The specimen should be deposited in ''Carlsruher Sammlung'' but we were not able to localize any historical collection of fossil insects in Karlsruhe (Germany). For that reason, we consider the only known specimen as lost. Proposed synonymy is based on the figure and the measurements given in the original description. Fig. 4 of the original description]. Paralectotypes, piece only (ETHZ NO-Z51.3-39-4:4-Pl.I143); incomplete specimen with partly preserved parts of head, pronotum and anterior legs, placed in a wooden box labeled ''Pl. I. 143'' [corresponds to plate 4, fig. 2 of the original description]; piece and counterpiece (ETHZ NO-Z51.3-39-4:4- Pl.I144), partly preserved pair of posterior legs, larger piece broken in two parts, both labeled ''Hydrophilus / spectabilis H. 144'' [corresponds with plate IV, Fig. 7
fig. 15 of the original ...
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... knorrii was described according to a single isolated elytron deposited in ''Carlsruher Sammlung'' ( Fig. 5.7). Since we were not able to localize any collection of fossils insects in Karlsruhe, the specimen seems to be lost and we based our results solely on the data provided in the original description. When the measurements of the elytron listed in the original description were transferred to millimeters and included into the scatter plot (Fig. 4), it seems to fall into the size variation of H. spectabilis. For that reason, we synonymize here H. knorrii with H. ...
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Citations
... The discovery is based on a reinterpretation of various insect groups from the famous Öhningen Lagerstätte, following on prior reviews of Bibionidae and Hydrophilidae (Fikáček and Schmied 2013;Skartveit and Pika 2014). ...
Clubtail dragonfly Lindenia heeri sp. nov., is described and figured as the first fossil representative of the extant lindeniine genus Lindenia, based on a finely preserved forewing from the mid-Miocene lacustrine maar of Öhningen, Germany. The
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... Hypotheses on the timing of the evolution of the Hydrophilidae have also been proposed and explored (Bloom et al. 2014, Toussaint andShort 2018) but the number of fossils documenting the past diversity remains limited. Most Mesozoic and Cenozoic fossils of the family are preserved as compressions (e.g., Fikáček et al. 2010b, 2010a, 2014, 2017, Fikáček and Schmied 2013. Amber inclusions are very rare, representing only five species from Cretaceous Burmese amber (Fikáček et al. 2017), Eocene Baltic amber (Arriaga-Varela et al. 2021b) and Miocene Dominican amber (Fikáček and Engel 2011). ...
We describe the first definite fossil of the water scavenger beetle subfamily Enochrinae (Coleoptera: Hydrophilidae): Cymbiodyta samueli sp. n. from the Eocene Baltic amber from the Lithuanian coast. The new species is extremely similar and likely closely related to the only European species, C. marginella and confirms the European occurrence of the genus since the Eocene. A reanalysis of the historical biogeography of the genus, including the fossil taxon, revealed a wide Euro-American distribution of the ancestor of all modern species of the genus, corresponding to the position of landmasses and existing land connections between North America and Europe in the Late Cretaceous. The biogeographic reconstructions and the fossil both suggest that European Cymbiodyta is an ancient relict lineage which used to be more diverse in the past but survived until today in a single species C. marginella.
... The Middle Miocene continental deposits of Öhningen ("Oeningen" in early literature) in south-western Germany are famous for the diversity of their fossil flora and insect fauna (Hantke 1954, SelmeieR 1990, FiKáčeK & schMied 2013. It is likewise famous for the fossil giant salamander Andrias scheuchzeri (Holl, 1831), which was first described as a human fossil from before the Biblical deluge. ...
Herein, we revise an extensive set of mollusc fossils from the Upper Freshwater Molasse deposits of Öhningen palaeolake (SW Germany; Middle Miocene, MN7). Based on material housed in paleontological collections in Europe and North America, we present the first thorough systematic account of the phylum from this historic locality. A total of ten species were identified from Öhningen: three freshwater gastropods (Lymnaea dilatata Noulet, 1854, Gyraulus cf. applanatus (Thomä), Planorbarius mantelli (Dunker)), two terrestrial gastropods (Granaria cf. schuebleri (Klein), Palaeotachea sylvestrina (Schlotheim)), and five bivalves (Anodonta splendens Goldfuss, A. lavateri (Münster), Pseudunio flabellatus (Goldfuss), “Dreissena” sp., Pisidium escheri (Mayer-Eymar, 1865)). Three freshwater and five terrestrial gastropod genera mentioned in older literature could not be found in the available material. Ours is the first report of a Dreissenidae from Öhningen
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The Elateridae (click-beetles) are the largest family in Elateroidea; however, their relationships, systematics and classification remain unclear. Our understanding of the origin, evolution, palaeodiversity and palaeobiogeography of Elateridae, as well as reconstruction of a reliable time-calibrated phylogeny for the group, are hampered by the lack of detailed knowledge of their fossil record. In this study, we summarize the current knowledge on all described fossil species in Elateridae, including their type material, geographic origin, age, bibliography and remarks on their systematic placement. Altogether, 261 fossil species classified in 99 genera and nine subfamilies are currently listed in this family. The Mesozoic click-beetle diversity includes 143 species, with most of them described from the Jurassic Karatau, and 118 described species are known from the Cenozoic deposits, mainly from the Eocene North American Florissant Formation and European Baltic amber. Available data on the described past diversity of Elateridae suggest that almost all fossil lineages in this group are in urgent need of revision and numerous Mesozoic species might belong to different families. Our study is intended to serve as a comprehensive basis for all subsequent research focused on the click-beetle fossil record.
... A few described fossil species, Perotis bruckmanni Heer, 1862; P. lavateri Heer, 1847; Capnodis antiqua Heer, 1847; C. puncticollis Heer, 1847 andC. spectabilis Heer, 1862, originated from the locality of Öhningen (South Germany) with the age assigned to the Late Miocene (Selmeier, 1990;Lutz, 1997;Fikáček & Schmied, 2013). ...
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... Still, this is not the first instance of alydids being confused with predatory heteropterans of Cimicomorpha (see Jordan, 1969;Kerzhner, 1981;Popov, 2007). Likewise, it is not the first instance of early studied material from Öhningen requiring a revised classification (e.g., Fikáček & Schmied, 2013). Thus, this study is another example that highlights the need for more contemporary treatment and/or reassessment of previously described fossil insect taxa, particularly those described in, and untouched since, the 19 th century. ...
It is no secret that fossilization can be a fickle process, being strongly influenced by a number of abiotic (e.g., habitat, oxygen level, isolation from other organisms, deposition, land disturbance, type of fossilization) and biotic (e.g., skeletal type, size of organism, life stage, population size) factors (e.g., Zherikhin, 2002). It also is a tremendously slow process, providing an extensive window of time for those forces to exert their influence. The result is a highly variable state of preservation from fossil to fossil. Some are extraordinarily true-to-life depictions of what the organism actually looked like millions of years ago (Li et al., 2013; Swanson et al., in press). Others, disarticulated and/or subject to geological pressures and upheavals, can fossilize less completely, with the resulting “depiction” difficult to interpret. More than occasionally, low preservational quality negatively affects taxonomic resolution, resulting in the misidentification or erroneous placement of fossil taxa (e.g., Popov, 2007; Fikáček & Schmied, 2013).
... More than 3,000 extant species have been described (Hansen 1999a;Short 2018), while approximately 180 extinct species have been mentioned in the literature. Detailed studies of specimens from some deposits revealed that a large percentage of the fossil records attributed to Hydrophilidae is based on misidentifications (e.g., Fikáček and Schmied 2013;Fikáček et al. 2014). The majority of described fossil water scavenger beetles are compression fossils from the Mesozoic or Eocene to Miocene deposits (e.g., Fikáček and Schmied 2013), and therefore many crucial morphological characters are not clearly visible. ...
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ABSTRACT
Water scavenger beetles (Coleoptera: Hydrophilidae) are a broadly distributed group with high extant diversity. However, reliable records from Baltic amber (Eocene) are scarce and limited to two undescribed species previously reported in the literature. Here we study these two specimens plus four additional ones.
All specimens were sub-optimal in terms of preservation and visibility of their morphology using traditional light microscopy. Opaque bubbles, cracks in the amber pieces and contracted appendages obscured most of the crucial characters, preventing their formal description and making them prone to misidentification. Here we used X-ray micro-computed tomography (μCT) to reconstruct the morphology of the embedded specimens and described three new species: Anacaena morla sp. nov., Crenitis profechuyi sp. nov. and Helochares fog sp. nov. Micro-CT reconstructions allowed us to visualise the morphology of the specimens to an extent where it was possible to hypothesise possible phylogenetic relations of the new taxa as well as the biogeographic implications of these relationships.
... Contemporary revisions of old fossil descriptions usually tend to reduce the number of species described in a given taxon. For example, a recent revision of hydrophilid beetles described by Heer (1862) from the Ö hningen locality in Germany, known for its rich insect deposits (Selmeier, 1990), revealed numerous inaccurate family and genus attributions (Fikácek & Schmied, 2013). Heer (1862) Paulian, 1976) from the localities in Lake Victoria (Paulian, 1976) and Ö hningen, Germany (Heer, 1862) can be reliably considered as scarabaeines, but their generic placement will require a separate investigation of the specimens. ...
Despite the increasing rate of systematic research on scarabaeine dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae), their fossil record has remained largely unrevised. In this paper, we review all 33 named scarabaeine fossils and describe two new species from Dominican amber (
Canthochilum alleni
sp.n.
,
Canthochilum philipsivieorum
sp.n.
). We provide a catalogue of all fossil Scarabaeinae and evaluate their assignment to this subfamily, based primarily on the original descriptions but also, where possible, by examining the type specimens. We suggest that only 21 fossil taxa can be reliably assigned to the Scarabaeinae, while the remaining 14 should be treated as doubtful Scarabaeinae. The doubtful scarabaeines include the two oldest dung beetle fossils known from the Cretaceous and we suggest excluding them from any assessments of the minimum age of scarabaeine dung beetles. The earliest reliably described scarabaeine fossil appears to be
Lobateuchus parisii
, known from Oise amber (France), which shifts the minimum age of the Scarabaeinae to the Eocene (53 Ma). We scored the best-preserved fossils, namely
Lobateuchus
and the two
Canthochilum
species described herein, into the character matrix used in a recent morphology-based study of dung beetles, and then inferred their phylogenetic relationships with Bayesian and parsimony methods. All analyses yielded consistent phylogenies where the two fossil
Canthochilum
are placed in a clade with the extant species of
Canthochilum
, and
Lobateuchus
is recovered in a clade with the extant genera
Ateuchus
and
Aphengium
. Additionally, we evaluated the distribution of dung beetle fossils in the light of current global dung beetle phylogenetic hypotheses, geological time and biogeography. The presence of only extant genera in the late Oligocene and all later records suggests that the main present-day dung beetle lineages had already been established by the late Oligocene–mid Miocene.
... Fossils have been excavated here since the 18th century and the site has yielded a very rich material of fossil animals and plants, roughly 900 species of animals and 500 species of plants have been described from the site to date (Ungricht & Pika-Biolzi, undated). The insect fauna of Öhningen is famous as one of the richest fossil insect faunas known, though it is likely that the number of species names published is an over-estimate of the number of recognisable species from the site (Fikácek & Schmied 2013). The Öhningen insect fossils were mainly described by Heer in four monographs (Heer 1847(Heer , 1849(Heer , 1853(Heer , 1862. ...
revised. Three species have been named, but not published, most likely by Oswald Heer, these are formally
described here: Bibio crassinervis sp.n., Dilophus pumilio sp.n. and Plecia macilenta sp.n. The
genus Lithosomyia Carpenter, 1986 ( = Mesomyia Pongracz, 1928, nec Macquart, 1850) is synonym
ized with Bibio Geoffroy, 1762. Bibio elongatus Heer, 1849 is not recognizable to genus and is
reclassified as Bibionidae, incertae sedis. Bibio obsoletus Heer, 1849 is recognizable as a Bibio species
but entirely devoid of characters useful for species identification, and is placed as Bibio, incertae
sedis. Bibio oblongus Heer, 1849 is synonymized with Bibio brevis Heer, 1849, Protomyia speciosa
Heer, 1864 is synonymised with Protomyia amoena Heer, 1849 and moved to the genus Penthetria,
and Protomyia affinis Heer, 1849 is synonymised with Plecia jucunda (Heer, 1849). A key is presented
to the recognisable species of Bibionidae from Öhningen. The material from Öhningen consists
almost exclusively of female specimens, the problems this raises are discussed.
