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.     

Fossil water striders from the Middle Eocene fossil sites of Eckfeld and Messel,Germany (Hemiptera,Gerromorpha)

Water striders (Hemiptera, Gerromorpha) are a very distinct ecological group specially adapted for life on the water surface. The present paper reports on four species of Gerromorpha from the Middle Eocene fossil sites of Eckfeld and Messel describingLutetiabates eckfeldensis n. gen. et n. sp.,Cylindrobates messelensis n. gen. et n. sp. (both Gerridae), and two nymphs most probably of the genusGerris. The record of two new members of Gerridae from the Paleogene as well as the hitherto known Gerromorpha from fossiliferous resins document a distinctly higher diversity of water striders within die European Paleogene than today. Lastly, comments are made on the fossil history as well as on the palaeobiological and palaeobiogeographical significance of the faunas.     

Exceptional three-dimensional preservation and coloration of an originally iridescent fossil feather from the Middle Eocene Messel Oil Shale

A feather from the Eocene Messel Formation, Germany, has been demonstrated to have been originally structurally colored by densely packed sheets of melanosomes similar to modern iridescent feathers exhibiting thin-film diffraction. The fossil itself currently exhibits a silvery sheen, but the mechanism for generating this optical effect was not fully understood. Here we use scanning electron microscopy, electron probe microanalysis, and dual-beam focused ion beam scanning electron microscopy to investigate the source of the silvery sheen that occurs in the apical feather barbules. Focused ion beam scanning electron microscopy provides a powerful tool for studying three-dimensionality of nanostructures in fossils. Use of the method reveals that the flattened apical barbules are preserved almost perfectly, including smooth structural melanosome sheets on the obverse surface of the fossil feather that are identical to those that cause iridescence in modern bird feathers. Most of each apical barbule is preserved beneath a thin layer of sediment. The silvery sheen is generated by incoherent light diffraction between this sediment layer and melanosomes and, although related to the original iridescence of the feather, is not a feature of the feather itself. The reddish and greenish hues frequently exhibited by fossil feathers from the Messel Formation appear to be due to precipitates on the surface of individual melanosomes.     

The implications of the numerical dating of the Messel fossil deposit (Eocene, Germany) for mammalian biochronology

Radiometric dating of volcaniclastic material from a borehole below the Middle Eocene Messel Formation leads to a revised numerical age of the Messel fossil deposit. Accordingly, the Grauvian/Geiseltalian boundary moves up from about 49 to 47.5 ma. The consequences of this change as well as the necessity of the Geiseltalian European Land Mammal Mega-Zone (ELMMZ)¹ for mammalian biochronology are discussed.     

Fossil ectomycorrhizae from the Middle Eocene

Fossil ectomycorrhizae were found recently among permineralized plant remains in the middle Eocene Princeton chert of British Columbia. The ectomycorrhizae are associated with roots of Pinus and have a Hartig net that extends to the endodermis, a pseudoparenchymatous mantle, and contiguous extramatrical hyphae that are simple-septate. The mycorrhizal rootlets lack root hairs and dichotomize repeatedly to form large, coralloid clusters. Reproductive structures are absent. Based on the morphological characteristics, and the identity of the host, the closely related basidiomycete genera Rhizopogon and Suillus are suggested as comparable extant mycorrhizal fungi. These exquisitely preserved specimens represent the first unequivocal occurrence of fossil ectomycorrhizae and demonstrate that such associations were well-established at least 50 million years ago.     

Cyanobacteria and seasonal death: A new taphonomic model for the Eocene Messel lake

Two new observations lead us to reconsider the taphonomy of Messel. First, several horses as well as turtles indicate death at a specific season although they come from different horizons. Second, specific structures in the Sediments are very similar to those found in Neumark-Nord. There they could be related to Cyanobacteria. Blooms of Cyanobacteria may poison die surface water. From actual observations it is known that animals drinking such poison water collapse immediately and die mostly near or within the water soon. This model might explain why birds and bats are so frequent throughout the entire profile of Messel. Both drink from the water surface during flight. In contrast to volcanic exhalations, assumed so far, such algae blooms occur repeatedly and always during early summer and autumn. An erratum to this article is available at .     

Tachypteron franzeni n. gen., n. sp., earliest emballonurid bat from the Middle Eocene of Messel (Mammalia, Chiroptera)

A new genus and species of emballonurid microchiropteran,Tachypteron franzeni, is described from the early Middle Eocene (MP 11) of Grube Messel, near Darmstadt, Germany. The holotype is extraordinarily well-preserved. It is the first unequivocal representative of an extant clade among Messel bats and the oldest unequivocal record of Emballonuridae. The flight apparatus ofT. franzeni is highly specialized for a rapid and constant flight style. The proportions of the strikingly narrow wing, the outline of the flight membranes and external ear, the morphology of the postcranial skeleton, and the relative cochlea size ofT. franzeni and extantTaphozous species are almost identical, while the dentition ofT. franzeni is distinguished by more plesiomorphic features.Tachypteron and the fairly diverse Paleogene record of emballonurids from France, consisting mainly of isolated teeth, document the earliest radiation known from an extant bat family and suggest a rather intense pre-Middle Eocene diversification of emballonurids.     

Non-marine Teredolites from the Middle Eocene of southern England

Middle Eocene, non-marine sediments from southern England contain examples of Teredolites borings in two contrasting palaeoenvironmental settings, viz.: (A) as in situ borings in an allochthonous lignite in an abandoned river channel and (B) as bored logs in cross-bedded fluvial sandstones of probable point-bar origin. The lignite is 0.30 m thick, of which the upper 0.20 m is intensely bored. Rounded pebbles of ?charcoal at its base also show small borings. A log in a fluvial sandstone shows densely-packed. radial club-shaped borings. now filled with sandstone. The wood substrate has subsequently been oxidized away. To our knowledge, this is the first detailed account of Teredolites from an ancient freshwater setting.     

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 Developmental Origins of Mosaic Evolution in the Primate Limb Skeleton

The central hypothesis of this paper is that basic properties of vertebrate limb development bias the generation of phenotypic variation in certain directions, and that these biases establish focal units, or regions, of evolutionary change within the primate hand and foot. These focal units include (1) a preaxial domain (digit I, hallux or pollex, metapodial and proximal phalanx), (2) a postaxial domain (metapodials and phalanges of digits II?CV), and (3) a digit tip domain (terminal phalanges and nails/claws of rays I?CV). The existence of these focal units therefore provides a mechanistic basis for mosaic evolution within the hand and foot, and can be applied to make specific predictions about which features of the limb skeleton are most likely to be altered in primate adaptive radiations over time. Examination of the early primate fossil record provides support for this model, and suggests that the existence of variational tendencies in limb development has played a major role in guiding the origin and evolution of primate skeletal form.     

Morphology and Function of the Vertebral Column in Remingtonocetus domandaensis (Mammalia, Cetacea) from the Middle Eocene Domanda Formation of Pakistan

The archaeocete family Remingtonocetidae is a group of early cetaceans known from the Eocene of India and Pakistan. Previous studies of remingtonocetids focused primarily on cranial anatomy due to a paucity of well-preserved postcranial material. Here we describe the morphology of the known vertebral column in Remingtonocetus domandaensis based largely on a single well-preserved partial skeleton recovered from the upper Domanda Formation of Pakistan. The specimen preserves most of the precaudal vertebral column in articulation and includes seven complete cervical vertebrae, ten partial to complete thoracic vertebrae, six complete lumbar vertebrae, and the first three sacral vertebrae. Cervical centra are long and possess robust, imbricating transverse processes that stabilized the head and neck. Lumbar vertebrae allowed for limited flexibility and probably served primarily to stabilize the lumbar column during forceful retraction of the hind limbs. Vertebral evidence, taken together with pelvic and femoral morphology, is most consistent with interpretation of Remingtonocetus domandaensis as an animal that swam primarily by powerful movement of its hind limbs rather than dorsoventral undulation of its body axis.     

Taxonomic description of in situ bee pollen from the middle Eocene of Germany

The middle Eocene Messel and Eckfeld localities are renowned for their excellently preserved faunas and diverse floras. Here we describe for the first time pollen from insect-pollinated plants found in situ on well-preserved ancient bees using light and scanning electron microscopy. There have been 140 pollen types reported from Messel and 162 pollen types from Eckfeld. Here we document 23 pollen types, six from Messel and 18 from Eckfeld (one is shared). The taxa reported here are all pollinated by insects and mostly not recovered in the previously studied dispersed fossil pollen records. Typically, a single or two pollen types are found on each fossil bee specimen, the maximum number of distinct pollen types on a single individual is five. Only five of the 23 pollen types obtained are angiosperms of unknown affinity, the remainder cover a broad taxonomic range of angiosperm trees and include members of several major clades: monocots (1 pollen type), fabids (7), malvids (4), asterids (5) and other core eudicots (1). Seven types each can be assigned to individual genera or infrafamilial clades. Since bees visit only flowers in the relative vicinity of their habitat, the recovered pollen provides a unique insight into the autochthonous palaeo-flora. The coexistence of taxa such as Decodon, Elaeocarpus, Mortoniodendron and other Tilioideae, Mastixoideae, Olax, Pouteria and Nyssa confirms current views that diverse, thermophilic forests thrived at the Messel and Eckfeld localities, probably under a warm subtropical, fully humid climate. Our study calls for increased attention to pollen found in situ on pollen-harvesting insects such as bees, which can provide new insights on insect-pollinated plants and complement even detailed palaeo-palynological knowledge obtained mostly from pollen of wind-pollinated plants in the dispersed pollen record of sediments. In the case of Elaeocarpus, Mortoniodendron, Olax and Pouteria the pollen collected by the middle Eocene bees represent the earliest unambiguous records of their respective genera.     

Middle Eocene primate tarsals from China: implications for haplorhine evolution

We describe tarsal remains of primates recovered from the Middle Eocene (approximately 45 mya) Shanghuang fissures in southern Jiangsu Province, China. These tarsals document the existence of four higher-level taxa of haplorhine primates and at least two adapid species. The meager and poorly preserved adapid material exhibits some similarities to European adapines like Adapis. The haplorhine primates are divided into two major groups: a "prosimian group" consisting of Tarsiidae and an unnamed group that is anatomically similar to Omomyidae; and an "anthropoid group" consisting of Eosimiidae and an unnamed group of protoanthropoids. The anthropoid tarsals are morphologically transitional between omomyids (or primitive haplorhines) and extant telanthropoids, providing the first postcranial evidence for primates which bridge the prosimian-anthropoid gap. All of the haplorhines are extremely small (most are between 50-100 g), and the deposits contain the smallest euprimates ever documented. The uniqueness of this fauna is further highlighted by the fact that no modern primate community contains as many tiny primates as does the fauna from Shanghuang.     

Anthracobunids from the Middle Eocene of India and Pakistan Are Stem Perissodactyls

Anthracobunidae is an Eocene family of large mammals from south Asia that is commonly considered to be part of the radiation that gave rise to elephants (proboscideans) and sea cows (sirenians). We describe a new collection of anthracobunid fossils from Middle Eocene rocks of Indo-Pakistan that more than doubles the number of known anthracobunid fossils and challenges their putative relationships, instead implying that they are stem perissodactyls. Cranial, dental, and postcranial elements allow a revision of species and the recognition of a new anthracobunid genus. Analyses of stable isotopes and long bone geometry together suggest that most anthracobunids fed on land, but spent a considerable amount of time near water. This new evidence expands our understanding of stem perissodactyl diversity and sheds new light on perissodactyl origins.     

New species of Primozygodactylus from Messel and the ecomorphology and evolutionary significance of early Eocene zygodactylid birds (Aves,Zygodactylidae)

Zygodactylids (Zygodactylidae) are the extinct sister taxon of passerines (Passeriformes) and among the more abundant small arboreal birds in the early Eocene German fossil site Messel. Four species of the taxon Primozygodactylus have previously been identified and here two new species are described. In addition, new fossils of the poorly known P. eunjooae are reported. The fossils corroborate the presence of two very long central tail feathers for Primozygodactylus, and the feathering of the taxon corresponds with that of extant birds foraging in scrub and undergrowth. Species diversity of zygodactylids falls short of that of passerines, the most species-rich extant avian clade, and in bill shapes they also do not reach the diversity seen in extant Passeriformes. Because zygodactylids closely resemble passerines in skeletal morphology, the evolutionary success of passerines is unlikely to be due to particular skeletal features. Passerines and zygodactylids coexisted in Europe from the early Oligocene to the middle Miocene, and both groups probably differed in ecological characteristics. The zygodactyl foot of zygodactylids may have represented an adaptation for clinging to tree trunks, and if nesting in tree cavities, zygodactylids may have succumbed to competition for safe nesting places with avian or mammalian competitors.

http://zoobank.org/urn:lsid:zoobank.org:pub:7F142141-7E1F-4568-89BF-E2363D128C36     

The deciduous dentition and dental replacement in the Eocene bat Palaeochiropteryx tupaiodon from Messel: The primitive condition and beginning of specialization of milk teeth among Chiroptera

The deciduous dentition and tooth replacement pattern of Palaeochiropteryx tupaiodon from the early Middle Eocene of Messel, near Frankfurt, Germany, are described. Ontogenetic states include fetuses to subadults. The posterior portion of the deciduous dentition (dP3-4) still shows the primitive eutherian condition of molarization, while the anterior part (dI-dC) was already engaged in the evolution of the highly derived condition found in living bats for clinging to the mother's fur. A styliform and sharp anterior dentition is considered a prerequisite in earliest chiropteran evolution. The greatly modified milk teeth of all living bats developed in different clades by parallel evolution under high selective pressure. The tiny and, at initial stages, poorly calcified teeth are substantiated by a newly developed microradiographic technique which is described in detail.