Fossil water‐penny beetles (Coleoptera: Psephenidae: Eubrianacinae) from the Eocene of Europe,with remarks on their phylogenetic position and biogeography

Abstract: A species of water‐penny beetles is recorded from larval instars from the Middle Eocene Messel pit fossil site in Germany. This species clearly belongs to the psephenid subfamily Eubrianacinae, but its precise systematic affinities remain unclear. It is the second fossil species of this taxon recorded from Europe. The holotype of the first fossil species, Eubrianax vandeli Bertrand and Laurentiaux, 1963, is lost. The high number of fossil specimens from Messel allowed discussion of stratigraphic and spatial occurrence of the eubrianacine species in the Messel pit, but its larval ontogeny could not be unravelled. Because of the high number of fossils and their wide occurrence, it is inferred that the species from Messel gen. sp. 1 was an autochthonous faunal element of the Eocene Lake Messel, which might indicate that some parts of the former Lake Messel had a shore area with stones. The analysis of the phylogenetic position of both Eocene eubrianacine species showed that their phylogenetic placement cannot be resolved because preservational influences limit the evaluation of characters. The historical biogeography of Psephenidae and Eubrianacinae is analysed and discussed. The fossil record shows that psephenid beetles have fossil members occurring outside their current distribution range, so interpretations of their biogeography based only on extant members can be misleading.     

Synonymy and actual affinities of the putative Middle Eocene “New World vulture” Eocathartes Lambrecht , 1935 and “hornbill” Geiseloceros Lambrecht , 1935 (Aves, Ameghinornithidae)

The taxonomy and phylogenetic affinities of the putative “New World vulture”Eocathartes robustus,Lambrecht, 1935 and the “hornbill”Geiseloceros robustus Lambrecht, 1935 from the Middle Eocene of the Geisel Valley in Germany are revised. It is shown that the holotype specimens belong to a single individual, whose osteology closely resembles that ofStrigogyps sapea (Peters, 1987) from the Middle Eocene of Messel (Germany). The species is classified intoStrigogyps robustus (Lambrecht, 1935), n. comb., and provides further evidence for the great similarity between the Eocene avifaunas of the Geisel Valley and Messel.Strigogyps is a representative of the Ameghinornithidae whose phylogenetic affinities are uncertain; there is no fossil record of either Cathartidae (New World vultures) or Bucerotidae (hornbills) from the Geisel Valley.     

Miozäne Algenlaminite von Hausen in der Rhön

Laminated lake sediments of Early Miocene age are described from the Eisgraben near Hausen in the Rhön Mountains of central Germany. In composition they resemble the Middle Eocene Messel Oil Shale. Particularly remarkable are laminar concentrations ofTetraedron minimum (Chlorophyceae) remains. This is the third occurrence of these algae in the fossil record.     

Studies on Cenozoic crocodiles: 8.Bergisuchus dietrichbergi Kuhn (Sebecosuchia: Bergisuchidae n. fam.) from the Middle Eocene of Germany, some new systematic and biological conclusions

Bergisuchus dietrichbergi Kuhn is redescribed as a member of its own family, Bergisuchidae n. fam., within the Sebecosuchia. At present, two fragmentary specimens ofBergisuchus are known: the holotype ofBergisuchus, a partial rostrum, comes from the lower part of the Middle Eocene of the Messel pit, near Darmstadt; and a recently discovered fragmentary mandible from the middle part of the Middle Eocene of Geiseltal in an opencast coal-mining pit near Halle (Saale).Bergisuchus shows similarities with species ofSebecus which are known from the Paleocene to Miocene from South America. All palaeoecological and biomechanical data support the hypothesis ofBergisuchus as a small, mainly terrestrial crocodile, which did not build its nests near either of the fossil localities from which it has been collected.     

Pumiliornis tessellatus Mayr, 1999 revisited — new data on the osteology and possible phylogenetic affinities of an enigmatic Middle Eocene bird

Although the avian speciesPumiliornis tessellatus Mayr, 1999 is known from two skeletons from the Middle Eocene of Messel in Germany, its phylogenetic affinities remained enigmatic. The new osteological data presented in this study document thatP. tessellatus had an at least semizygodactyl foot, with a very wide basal phalanx of the fourth toe, and lacked an ossified pons supratendineus on the distal tibiotarsus. Compared to the known zygodactyl and semizygodactyl birds, this tiny Middle Eocene species resembles the late Eocene/early Oligocene taxonEocuculus Chandler, 1999. Anew, tentatively referred wing ofEocuculus from the early Oligocene of France is described and compared withPumiliornis.      

Paleoecology of early middle Eocene bats from Messel,FRG. aspects of flight,feeding and echolocation

By their diversified flight apparatus Messel bats occupied specific flight niches similar to those of extant tropical bats. The small Palaeochiropteryx tupaiodon is considered to be most specialized for hunting close to the ground and for hovering inside dense vegetation. Contrarily, Hassianycteris spp. most likely were high and fast flyers in the open space.

The analysis of gut contents proves that Palaeochiropteryx spp. exclusively fed on small moths and caddis flies, i.e. slow and low flying insects. For P. tupaiodon this confirms the foraging strategy independently from wing morphology. Hassianycteris spp. preyed mainly on beetles or other insects with thick cuticules.

Inner ears of Messel microbats are less specialized compared to those of recent species. Especially P. tupaiodon shows no acoustical specialization with regard to its hunting habitat. Thus, we assume that during the early evolution of bats the development of different flight styles and wing shapes preceded acoustical refinements of the echolocation system.     

The early Eocene birds of the Messel fossil site: a 48 million‐year‐old bird community adds a temporal perspective to the evolution of tropical avifaunas

Birds play an important role in studies addressing the diversity and species richness of tropical ecosystems, but because of the poor avian fossil record in all extant tropical regions, a temporal perspective is mainly provided by divergence dates derived from calibrated molecular analyses. Tropical ecosystems were, however, widespread in the Northern Hemisphere during the early Cenozoic, and the early Eocene German fossil site Messel in particular has yielded a rich avian fossil record. The Messel avifauna is characterized by a considerable number of flightless birds, as well as a high diversity of aerial insectivores and the absence of large arboreal birds. With about 70 currently known species in 42 named genus‐level and at least 39 family‐level taxa, it approaches extant tropical biotas in terms of species richness and taxonomic diversity. With regard to its taxonomic composition and presumed ecological characteristics, the Messel avifauna is more similar to the Neotropics, Madagascar, and New Guinea than to tropical forests in continental Africa and Asia. Because the former regions were geographically isolated during most of the Cenozoic, their characteristics may be due to the absence of biotic factors, especially those related to the diversification of placental mammals, which impacted tropical avifaunas in Africa and Asia. The crown groups of most avian taxa that already existed in early Eocene forests are species‐poor. This does not support the hypothesis that the antiquity of tropical ecosystems is key to the diversity of tropical avifaunas, and suggests that high diversification rates may be of greater significance.     

Macrosiagon deuvei n. sp. (Coleoptera: Ripiphoridae) from the French Eocene amber

Macrosiagon deuvei n. sp., the second fossil representative of this extant genus of Ripiphoridae: Ripiphorinae: Macrosiagonini is described from the lowermost Eocene amber of Oise (France). The new species is compared with the extant species of the genus. Taxonomic position of other two fossil representatives of the family described from France by Perrichot et al. (2004) is discussed. The genus Paleoripiphorus Perrichot et al. 2004 is tentatively transferred from Ripiphorinae to Ripidiinae.     

Ein Archaeotrogon (Aves: Archaeotrogonidae) aus dem Mittel-Eoz?n der Grube Messel (Hessen, Deutschland)?

Zusammenfassung Ein Vogel aus dem Mittel-Eozän der Grube Messel (Hessen, Deutschland) wird als neue Gattung und Art der Archaeotrogonidae Mourer-Chauviré 1980 beschrieben (incertae sedis). Die Exemplare aus Messel wären die ersten artikulierten Skelette dieser Familie.Hassiavis laticauda n. gen. n. sp. unterscheidet sich vor allem im Bau des Coracoids von der GattungArchaeotrogon Milne-Edwards 1892. Besonders bemerkenswert ist die ausgezeichnete Erhaltung der Flügel- und Schwanzbefiederung einiger Exemplare. Einige Schwanzfedern des Holotypus zeigen eine Querbänderung, welche möglicherweise auf die ursprüngliche Pigmentierung dieser Federn zurückzuführen ist. Die phylogenetische Stellung der Archaeotrogonidae wird diskutiert. Zur Zeit sind keine Synapomorphien bekannt, welche diese Familie mit einer der bestehenden Ordnungen verbinden.

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An Archaeotrogon (Aves: Archaeotrogonidae) from the Middle Eocene of the Grube Messel (Hessen, Germany)?

Summary A bird from the Middle Eocene of the Grube Messel (Hessen, Deutschland) is described as a new genus and species of the Archaeotrogonidae Mourer-Chauviré 1980 (incertae sedis). The specimens from Messel would be the first articulated skeletons of this family.Hassiavis laticauda n. gen. n. sp. is distinguished from the genusArchaeotrogon Milne-Edwards 1892 in the morphology of the coracoid. The excellent preservation of the wing feathers and those of the tail in several of the specimens is exceptional. Some tail feathers of the holotype are barred, which might be traced back to the original pigmentation of these feathers. The phylogenetic position of the Archaeotrogonidae is discussed. At present no synapomorphies are known, which could set up a closer relationship between this family and one of the existing orders.

     

FOSSIL CERATOPHYLLUM (CERATOPHYLLACEAE) FROM THE TERTIARY OF NORTH AMERICA

Fossil fruits and a vegetative axis assignable to the extant genus Ceratophyllum are described from four North American Tertiary localities. Fossil fruits assignable to the extant species C. muricatum and C. echinatum are reported from the Eocene Green River and Claiborne formations, and the Miocene Esmerelda Formation, respectively. An extinct species, C. furcatispinum, is described from the Paleocene Fort Union Formation and represents the oldest published report of Ceratophyllum in the fossil record. The existence of extant angiosperm species in the Eocene is very unusual and may be attributable in this case to slow evolutionary rates and unusual evolutionary properties associated with hydrophily in the genus Ceratophyllum.     

Paroodectes feisti,der erste Miacide (Carnivora,Mammalia) aus dem Mittel-Eozän von Messel

From the Middle Eocene (Lower Lutetian) Oil Shale of Messel near Darmstadt (South-Hesse) the first completely preserved Miacid is described. The systematic position of the Miacidae is discussed before. A short review of the paleogeographic (North-America, Eurasia) and stratigraphic (L. Paleocene — U. Eocene, ? U. Oligocene) distribution is given; the fossil record of European species is more detailed. The odontological features of the Messel-Miacid are nearest to the new world genusOodectes from the Bridger Basin (Bridgerian). No direct relationship exists to European species. The structure of the postcranial skeleton shows great conformity with Northamerican Miacinae. Differences result from proportions skull-length/ stature and length of vertebral column/length of limbs. A well developed clavicle is extant. Scaphoid, lunar, and central are distinct. The humerometacarpal index is low (≤ 30). Metatarsals are elongated. Hand und foot are paraxone and adapted to plantigrade locomotion. An arboricole habit is assumed. The nameParoodectes feisti n. g. n. sp. is proposed. The species is considered as a member of the subfamily Miacinae. Besides morphologic features of the Viverridae and some Procyonidae, there are possibly certain relationships to the Primates.     

Highly resolved early Eocene food webs show development of modern trophic structure after the end-Cretaceous extinction

Generalities of food web structure have been identified for extant ecosystems. However, the trophic organization of ancient ecosystems is unresolved, as prior studies of fossil webs have been limited by low-resolution, high-uncertainty data. We compiled highly resolved, well-documented feeding interaction data for 700 taxa from the 48 million-year-old latest early Eocene Messel Shale, which contains a species assemblage that developed after an interval of protracted environmental and biotal change during and following the end-Cretaceous extinction. We compared the network structure of Messel lake and forest food webs to extant webs using analyses that account for scale dependence of structure with diversity and complexity. The Messel lake web, with 94 taxa, displays unambiguous similarities in structure to extant webs. While the Messel forest web, with 630 taxa, displays differences compared to extant webs, they appear to result from high diversity and resolution of insect–plant interactions, rather than substantive differences in structure. The evidence presented here suggests that modern trophic organization developed along with the modern Messel biota during an 18 Myr interval of dramatic post-extinction change. Our study also has methodological implications, as the Messel forest web analysis highlights limitations of current food web data and models.     

Protoresinacarus brevipedis gen. n., sp. n. from Early Cretaceous Burmese amber: the first fossil record of mites of the Family Resinacaridae (Acari: Heterostigmata: Pyemotoidea)

A new genus and species of mites, Protoresinacarus brevipedis gen. n., sp. n. (Acari: Heterostigmata: Pyemotoidea), is described from Early Cretaceous Burmese amber. This represents the first fossil record of a member of the family Resinacaridae. It is represented by 21 phoretic females adjacent to an adult mantidfly (Neuroptera: Mantispidae). This is the first record of phoresy of pyemotid mites on members of the insect order Neuroptera. The fossil mites differ from extant members of the family in possessing distinctly shorter legs I, which do not reach beyond the apex of the gnathosoma, and by the long setae v ₁, v ₂ and c ₂.     

The first New World occurrence of the Eocene birdPlesiocathartes (Aves: ?Leptosomidae)

Two new species of the Eocene avian taxonPlesiocathartes (Aves: ?Leptosomidae) are described. They are similar in size and overall appearance to other members of the genus, as well as to the extant Madagascan cuckoo-rollerLeptosomus discolor. So far, putative fossil Leptosomidae have been found only in the Eocene European de-posits of Messel, Geiseltal, Quercy, and London Clay. Both new species are from the Lower Eocene Green River Formation (USA) and thus the first New World representatives of this taxon.     

Paleorhodococcus dominicanus n. gen., n sp. (Actinobacteria) in a faecal droplet of Triatoma dominicana (Hemiptera: Reduviidae: Triatominae) in Dominican amber

Paleorhodococcus dominicanus n. gen., n sp. (Actinobacteria) is described from a faecal droplet of Triatoma dominicana (Hemiptera: Reduviidae: Triatominae) in Dominican amber. The fossil can be distinguished from the species of the closely related extant genus Rhodococcus Zopf 1891 by its spherical–ellipsoidal cocci forming substrate filaments with elementary branching, the clustering of coccoidal elements, the short filaments bearing reduced side branches and its occurrence in a faecal droplet of the extinct triatomine bug, P. dominicanus. This is the first fossil record of an Actinobacter, which shows that these organisms formed symbiotic associations with insects by the mid-Tertiary.     

First substantial Middle Eocene record of the Lithornithidae (Aves): A postcranial skeleton from Messel (Germany)

A postcranial skeleton of a representative of the palaeognathous Lithornithidae (Aves) is described from the Middle Eocene of Messel in Germany. The specimen is slightly smaller than Lithornis plebius from which it, however, differs in limb bone proportions. It constitutes the latest fossil record of the Lithornithidae in Europe, whose only other Middle Eocene record is a fragmentary tibiotarsus from North America.     

New trace fossils of termites (Insecta: Isoptera) from the late Eocene‐early Miocene of Egypt,and the reconstruction of ancient isopteran social behavior

Three new ichnogenera and five new ichnospecies are described for new trace fossils of termitaria (including associated gallery systems) of subterranean termites from upper Eocene through lower Miocene rocks of northern Egypt. All but two ichnospecies (Krausichnus trompitus, ichnogen, and ichnosp. nov., and A’, altus, ichnosp. nov.) show varying degrees of affinity to nests of the extant subterranean termite species Sphaerotermes sphaerothorax (Termitidae, Macro‐termitinae).

Termitichnus qatranii (Bown, 1982) is divided into two ichnospecies, the previously named T. qatranii, and a more generalized form, T. simplicidens, ichnosp. nov. Vondrichnus obovatus, ichnogen. and ichnosp. nov., is named for simple, possibly macrotermitine nests with oblate form, and Fleaglellius pagodus, ichnogen. and ichnosp. nov., records a nest form similar to V. obovatus, but one in which successive vertical growth by chamber apposition has produced a subterranean, tower‐like structure, reminiscent of that built by extant, epigeous Cubitermes. Krausichnus trompitus, ichnogen. and ichnosp. nov., and A’, altus, ichnosp. nov., record two unique nest architectures probably produced by unknown but related species of humivorous termites. The nest architecture expressed by Krausichnus is of wholly unknown affinity and is only distantly related in form to the ichnofossils of other termite nests known from the Tertiary of Egypt.

Study of the architecture of the nests of these ancient termites reveals details important in reconstructing the phylogeny of termite nests and documents two novel blueprints for chamber expansion and society budding in what were probably primitive Macrotermi‐tinae. It also indicates that at least two constructed edifices, earlier known only as epigeous manifestations of extant termite species, were almost certainly first developed by species living underground.

The extant Macrotermitinae are a subfamily of termites originally believed to have evolved in the post‐Eocene of the Ethiopian biogeo‐graphic region. We offer fossil evidence of four distinct, possibly macrotermitine structures from upper Eocene rocks. This evidence suggests that this termite subfamily (or at least their peculiar mode of nest construction) might have evolved considerably earlier, as it is already well established and exhibits several variants by the late Eocene in Egypt.     

FRUITS AND SEEDS OF TRIBE GORDONIEAE (THEACEAE) FROM THE EOCENE OF NORTH AMERICA

A new genus and species Gordoniopsis polysperma and two new species of Gordonia (Gordonieae, Camellioideae, Theaceae) are described based on fossil fruit and seed remains. These specimens are part of a large flora consisting of various plant organs from the middle Eocene Claiborne Formation in western Kentucky and Tennessee. Gordoniopsis is a five-valved loculicidally dehiscent capsule similar to capsules of Gordonia but differing in having unwinged seeds and a greater number of seeds per locule. The two Gordonia species are among the earliest unequivocal records of the genus and two of only four fossil Gordonia species known with in situ seeds. Two extinct genera, Gordoniopsis and Andrewsiocarpon, and the extant genus Gordonia in the tribe Gordonieae are known from the middle Eocene Claiborne flora, suggesting an early radiation within the tribe. Based on a survey of Recent fruits and seeds we concur with Keng's proposal to merge Laplacea with Gordonia.     

Inflorescence bracts of fossil and extant Tilia in North America,Europe, and Asia: patterns of morphologic divergence and biogeographic history

The enlarged inflorescence bract diagnostic of extant Tilia has an extensive Tertiary fossil record in the Northern Hemisphere. Diversity of bract morphology, and the extent of adnation between peduncle and bract, is reviewed for fossil and extant species of Tilia. An extinct type of bract with an orbicular outline and palmate venation is documented by the fossil species Tilia circularis (Chaney) comb. nov. from the early Oligocene of Oregon and is designated Type A. Living species of the genus have elongate bracts with predominately pinnate venation that are borne in two basic configurations: Type B, with the peduncle fused only to the extreme base of the bract lamina, as in extant Tilia endochrysea Hand.-Mzt. of southern China; and Type C with the peduncle fused medially along the basal one-third of the bract lamina, as in most extant species. Bracts of Type B were widely distributed in the Tertiary of western North America (late Eocene to Miocene) and Europe (early Miocene to Pliocene), while those of Type C are known in the fossil condition only from the middle and late Tertiary of Asia and Pliocene of Europe. The bracts of T. circularis, like those of type B, are borne on relatively long stalks and have the peduncle fused only at the extreme base. The fossil record supports recognition of the following characters as apomorphic in Tilia bract evolution: bracts sessile, peduncle adnate to the upper surface of the bract, and pinnate bract venation.