A remarkable new ‘gruiform’ bird from the Middle Eocene of Messel (Hessen, Germany)

Salmila robusta n. gen., n. sp. (Aves: ‘Gruiformes’) is described from the Middle Eocene of Messel (Hessen, Germany). The new species is assumed to have had a predominantly terrestrial habit. It shares derived features of the carpometacarpus with the recent Psophiidae and some taxa of the Cariamae. In several features, however, it distinctly differs from other ‘gruiform’ birds (e. g. in the larger proximal end of the humerus and in the shape of the margo caudalis of the sternum). AlthoughSalmila robusta has been tentatively assigned to the Cariamae in this study, its exact systematic position deserves further investigations. The rather short tarsometatarsus of the new species resembles that of the ‘Messel-phorusrhacid’Aenigmavis sapea Peters 1987.     

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.      

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.     

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.     

New specimens of the Middle Eocene fossil mousebird Selmes absurdipes Peters 1999

Described here are an excellently preserved new specimen of the Middle Eocene mousebird Selmes absurdipes from the type locality Messel (Hessen, Germany) and first records of this species from the Geiseltal (Sachsen-Anhalt, Germany). These specimens allow the recognition of several previously unknown osteological features, and thus more detailed comparisons with other fossil and recent Coliiformes. Selmes absurdipes, which previously has been known only from a single poorly preserved skeleton, most closely resembles the Lower Eocene species Chascacocolius oscitans in its overall morphology. The species shares a derived character with each of the two coliiform families, the extinct Sandcoleidae and the Coliidae, and at present it cannot be assigned to either of these families with certainty.     

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.     

Gymnoascus verticillatus smith

Summary The type specimen ofGymnoascus verticillatus Smith was studied and is described as the type of a new, monotypic genus,Actinodendron. The new genus is characterized by the formation of brown to dark colored ascocarps bearing brown appendages with verticels of short branchlets. Ascospores ofA. verticillatum (Smith)Orr &Kuehn are pale brown and almost discoid. Additional specimens of this species are reported.     

The first juvenile specimen of Eolacerta (Squamata: Eolacertidae) from the early–middle Eocene of the Messel Pit (Germany)

We describe the first juvenile specimen of an eolacertid lizard. The material comes from one of the most important Eocene localities, the Messel Pit in Germany. The new specimen provides unique information on the early ontogeny of Eolacerta, the largest known lizard from Messel, with a maximum snout-vent length greater than 30 cm and a mass approaching 1 kg. The specimen described here, with a SVL of 11.3 cm and an estimated mass of 21 g, can be allocated more precisely to Eolacerta robusta based on the co-occurrence and the combination of following features: (1) the nasal process of premaxilla is long; (2) position of lacrimal (being more anteriorly located in Stefanikia siderea); (3) the postorbital process of jugal is broad; (4) the ratio of the anterior and posterior region of the frontal between the sulcus interfacialis; (5) a mid-parietal constriction of the parietal table is present; (6) the interparietal shield broadens anteriorly; (7) the transverse sulcus is straight anteriorly. The incipient character of the parietal constriction and the slightly lower number of maxillary teeth (28 vs. 30–32 in adults) are consistent with a juvenile animal. Very important is evidence for the presence of pterygoid dentition (pointed teeth arranged in a single line) and the absence of palatine dentition. Ceratobranchial I is observed for the first time for this species, and its shape and length are very similar to those of Lacertidae. There are 27 presacral vertebrae in the juvenile, as in adults. In the juvenile specimen, ventral keel on the centrum is present in all vertebrae. The ventromedial portion of the ischium is well preserved here and gives information on the exact shape of this portion, at least in juvenile form. The scalation, as far as it is preserved, is similar to that of Stefanikia, except that the rectangular subdigital scales are longer in comparison with their width, and therefore have a broader appearance.     

A small, “wader-like” bird from the Early Eocene of Messel (Germany)

A small bird with a shorebird-like beak and a very long hind toe is described from the Early Eocene of Messel in Germany. Vanolimicola longihallucis, gen. et sp. nov. is one of the few candidate species for a “wading bird” from Messel, that is, a bird that may have foraged along the shoreline of the ancient lake. A few features indicate a relationship to charadriiform birds and Jacanidae (jacanas) in particular, but charadriiform affinities are only weakly supported. In the preserved skeletal elements, V. longihallucis also resembles the taxon Songzia from the Early Eocene of China, which is considered to be a representative of the Ralloidea (rails and allies). A potential biogeographic significance of the new Messel fossil is strengthened by similar fossils from the North American Green River Formation.     

A note onSaissetia privigna [Hem.: Coccidae] in Pakistan and the breeding of its natural enemies

During a survey for the parasites and predators ofSaissetia privigna De Lotto, 4 additional species of natural enemies, namelyAnysis saissetiae (Ashm.),Metaphycus citricola Annecke & Mynhardt,Eublemma sp. andPullus coccidivora Ayyar were recorded in Pakistan.Aneristus ceroplastae How. andEncyrtus lecaniorum Mayr, which are known to be important parasites of this scale, were bred onS. privigna andPulvinaria sp. cultured on fruits ofCucurbita maxima in the laboratory.A. ceroplastae was introduced into France and proved useful in controllingCeroplastes rusci (L.) on citrus.

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Résumé Au cours d'une campagne de recherche de parasites et prédateurs deSaissetia privigna De Lotto au Pakistan, on a trouvé 4 espèces supplémentaires d'ennemis naturels;Anysis saissetiae (Ashm.),Metaphycus citricola Annecke & Mynhardt,Eublemma sp. etPullus coccidivora Ayyar.Aneristus ceroplastae How. etEncyrtus lecaniorum Mayr, connus comme parasites importants de cette cochenille, ont été élevés surS. privigna etPulvinaria sp. multipliés en laboratoire sur des fruits deCucurbita maxima. A. ceroplastae a été introduit en France et s'est révélé efficace pour la lutte contreCeroplastes rusci (L.) sur citrus.

     

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.

     

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.     

Phaneropsolus macacae (Premvati, 1958) Saoud, 1964 from the Taiwanese monkey,Macaca cyclopis (Swinhoe, 1862)

In November, 1968, a single specimen ofPhaneropsolus macacae (Premvati, 1958)Saoud, 1964 was collected from the small intestine of a female Taiwanese monkey,Macaca cyclopis (Swinhoe, 1862). The Taiwanese monkey was added as a new host for this trematode.     

Mallomonas robusta (Chrysophyceae,Synuraceae): Electron microscopical study of silica armour

Mallomonas robusta (Matvienko)Péterfi & Momeu (Mallomonopsis robusta) has been investigated by transmission electron microscopy and is redescribed in detail. Its silica armour, consisting of scales and bristles, exhibits a very peculiar fine structural pattern. Differences and possible relationships with other species are discussed. The Romanian material is designated as the nomenclatural type (neotype).     

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.     

Exceptionally preserved plant parenchyma in the digestive tract indicates a herbivorous diet in the Middle Eocene bird <Emphasis Type="Italic">Strigogyps sapea</Emphasis> (Ameghinornithidae)

The holotype skeleton of the avian species Strigogyps sapea from the Middle Eocene of Messel preserves remains of the digestive tract that have not been studied so far. The crop content of the specimen includes numerous densely packed, tiny spherules containing large cavities. These peculiar structures match well with previously described fossil plant parenchyma cells with starch grain moulds. We hypothesize that the spherule layer represents plant parenchyma ingested by the bird, and that the spherules themselves are casts of cytoplasm with storage organelles. Our study provides a further example of exceptional soft tissue preservation in Messel vertebrates and shows that Strigogyps was at least facultatively herbivorous. A herbivorous diet of Strigogyps is also indicated by the fact that the crop content includes seed remains, and it constitutes further evidence against previously hypothesized close affinities between the taxon and the carnivorous Phorusrhacidae.     

The taxonomic status ofGraecopithecus

In 1944 a single mandibular specimen was found in Pyrgos, southern Greece.von Koenigswald (1972) described this specimen and allocated it to a new taxon,Graecopithecus freybergi.de Bonis and colleagues later described new hominid specimens from northern Greece and allocated them to the new taxon,Ouranopithecus macedoniensis.Martin andAndrews (1984), however, argued that there was no reason for maintaining a generic or specific distinction between the northern and southern Greek fossil populations as suggested byde Bonis andMelentis. This paper reexamines the taxonomic status of the southern and northern Greek hominids in light of recent fossil discoveries and concludes that there is no evidence for recognizing two hominine genera from the Miocene of Greece. There is evidence, however, to suggest that the Macedonian hominid is specifically distinct from the southern Greek hominid. Thus, becauseGraecopithecus has taxonomic priority overOuranopithecus, it is concluded that the Macedonian specimens should be allocated to a separate species ofGraecopithecus.     

Taxonomic significance of biométric characters and the consequences for classification and biostratigraphy, exemplified through moussoneliform daonellas (Daonella, Bivalvia; Triassic)

The middle Triassic bivalve genusDaonella Mojsisovics, 1874 would be an ideal guide fossil (worldwide distribution, high evolutionary rates); however, the classification of daonellas is difficult. Most daonellas lack distinct characters and have a wide range of morphologic variation. A new taxonomic attempt was necessary to get a coherentDaonella classification. The new taxonomic concept proposed herein consists of four steps. (1) Biometric characters are defined unambiguously, based on landmarks and pseudolandmarks. (2) The taxonomic significance of biometric characters is analyzed. Several characters depend significantly on growth stage of specimens or on the lithology of fossil localities. In addition, the intercorrelations between biometric characters are studied to exclude weighting in some morphologic features. The study of the taxonomic significance of biometric characters shows, how important a careful selection of taxonomic characters is. (3) Most daonellas are not determinable by univariate or bivariate methods. The principle component analysis (PCA) is applied to detect groups of similar holomorphs in multivariate space. (4) The interpreted groups from PCA are applied to a canonical discriminant function analysis. This analysis helps in deciding whether two groups are sufficiently distinct to be separated as different species. This new taxonomic concept was applied to moussoneliform daonellas of the middle Triassic from the Southern Alps. Three different forms were used for the study herein:Daonella moussoni (Mérian, 1853) from the Varenna Limestone (Perledo-Varenna Formation, Grigna, Italy),D. aff.moussoni (Mérian)Rieber, 1969 from the lower Meride Limestone (Monte San Giorgio, Switzerland) andD. pseudomoussoni Rieber, 1969 from the Grenzbitumenzone (Monte San Giorgio, Switzerland). The new classification shows thatD. pseudomoussoni is a separate species and distinguishable fromD. moussoni andD. aff.moussoni, which belong to the same species. A coherent systematics ofDaonella becomes possible on the basis of newly defined taxonomic characters. Daonellas can now be used as relevant guide fossils for the middle Triassic. The presented example demonstrates a biostratigraphic correlation between the lower Meride Limestone at Monte San Giorgio and the Varenna Limestone at Grigna. The stratigraphic distribution ofD. moussoni can be assigned to thegredleri Zone.     

New specimens of the early Eocene stem group galliform Paraortygoides (Gallinuloididae), with comments on the evolution of a crop in the stem lineage of Galliformes

Two new specimens of the fossil stem group galliform Paraortygoides messelensis Mayr 2000 (Gallinuloididae) are described from the Middle Eocene of Messel in Germany, including a complete skeleton in which the hitherto unknown skull of this species is preserved. The shorter and more protruding crista deltopectoralis of the humerus, also for the first time visible in one of the new specimens, shows gallinuloidids to be the sister taxon of all other, extinct and extant, galliform birds. Gallinuloidids distinctly differ from modern Galliformes in several other plesiomorphic osteological features, mainly of the pectoral girdle, of which the absence of a spina interna on the sternum is here reported for the first time. It is assumed that major evolutionary transformations in the stem lineage of Galliformes are related to the evolution of a large crop, which appears to have been absent in gallinuloidids. The vegetarian food component of gallinuloidids thus probably mainly consisted of soft plant matter rather than coarse material such as seeds.