New fossil ants of the subfamily Myrmeciinae (Hymenoptera, Formicidae) from Germany

Two new fossil members of the subfamily Myrmeciinae are described: Archimyrmex wedmannae, sp. nov (Grube Messel, Germany, Middle Eocene) and Prionomyrmex wappleri, sp. nov. (Rott, Germany, Late Oligocene). Members of the genus Archimyrmex were known earlier from Eocene deposits of the United States and Argentina, and members of the genus Prionomyrmex were known from Late Eocene Baltic amber.     

Elucidating the affinities and habitat of ancient, widespread Cyperaceae: Volkeria messelensis gen. et sp. nov., a fossil mapanioid sedge from the Eocene of Europe

The sedges (family Cyperaceae) are an economically and ecologically important monocot group dating back at least to the Paleocene. While modern genera are mostly unknown before the Oligocene, several extinct taxa are recognized as the earliest sedges. Their affinities have been unclear until now, because they are found as isolated, often abraded fruits or endocarps. Exceptionally preserved sedge fossils from the Middle Eocene of Messel, Germany yield more characters for identification. Fossil cyperacean infructescences with in situ pollen are recognized for the first time and show features of the early-divergent mapanioid sedges. Pollen resembles that of tribe Hypolytreae. Comparisons with extant taxa suggest the closest affinities with Hypolytrum and Mapania. However, the Messel fossils represent a distinct taxon, Volkeria messelensis gen. et sp. nov. Without the additional characters of infructescence and pollen, the Messel fruits would have been placed in the extinct genus Caricoidea, a typical Eocene sedge that was widespread across Eurasia. Similarities of fruit structure suggest that Caricoidea was also a mapanioid sedge. Mapanioid sedges are found today in tropical wet forests and swamps, a distribution suggesting that early sedges occupied a similar habitat, unlike many modern sedges, and were not precursors to open grassland vegetation.     

New spiders in Upper Cretaceous amber from New Jersey in the American Museum of Natural History (Arthropoda: Araneae)

The oldest described fossils of the extant spider family Araneidae (Araneinae; gen. et sp. indet.), the extant genus Orchestina (Oonopidae; O. sp. indet.) and the new fossil genus Palaeosegestria (Segestriidae; P. lutzzii gen. et sp. nov.) are presented from Upper Cretaceous amber of New Jersey. The known fossil range of the extant family Araneidae is extended approximately 50 myr from the previously oldest described araneid from the Middle Eocene oil shales of the Messel pit in Hesse, Germany. The fossil range of the extant genus Orchestina is also extended 50 myr from the previously oldest described specimen in Eocene Baltic amber.     

New trogons from the early Tertiary of Germany

A new trogon (Aves, Trogoniformes), Primotrogon? pumilio sp. nov., is described from the Middle Eocene (49 Ma) of Messel in Germany. It is the first articulated skeleton of an Eocene trogon, of which only very few fragmentary remains were hitherto known. It is also the earliest complete skeleton of a trogon, exceeding previous finds (Primotrogon wintersteini Mayr 1999) by at least 15 million years. An isolated wing of a trogon is further identified from the early Oligocene (about 30–34 Ma) of the fossil site Frauenweiler in Germany. Primotrogon? pumilio and all other sufficiently well‐preserved early Tertiary trogons appear to be stem group representatives of the Trogoniformes, corresponding with molecular clock data indicating a mid‐Tertiary origin of crown group Trogoniformes.     

EOCENE RECORDS OF BEE FLIES (INSECTA, DIPTERA, BOMBYLIIDAE, COMPTOSIA): THEIR PALAEOBIOGEOGRAPHIC IMPLICATIONS AND REMARKS ON THE EVOLUTIONARY HISTORY OF BOMBYLIIDS

Abstract:  The first fossil records of Holarctic representatives of the genus Comptosia Macquart, 1840 from the middle Eocene Messel Pit, Germany, and the upper Eocene of Florissant, USA, are reported. The fossil from Messel, Comptosia pria sp. nov., is represented by a well-preserved wing, displaying characteristic wing venation, remains of the second wing and elements of the thorax. The Florissant fossil, C. miranda comb. nov., is preserved almost completely with both wings and most parts of the body visible. Extant members of the genus Comptosia are currently found only in Australia, and close relatives are known from southern South America. Previously this group of genera has been thought to be of Gondwanan origin, but these Eocene fossil representatives of Comptosia from the Holarctic region argue against this interpretation, and suggest that Comptosia and its relatives were more widespread in both Northern and Southern hemispheres during the Palaeogene. The fossil record of bee flies suggests a major radiation of bombyliids in the late Cretaceous or early Cenozoic. This assumption is supported by the comparatively young geological age of many bombyliid host taxa. A key factor driving the specialization of many bombyliid larvae on ground-dwelling hosts may have been avoidance of competition with tachinid flies and parasitic hymenopterans.     

Diversity of Menispermaceae from the Paleocene and Eocene of South China

We present here the earliest known Asian fossil records of the Menispermaceae based on fossil fruits from Paleocene and Eocene localities in South China. A new genus and species, Paleoorbicarpum parvum sp. nov., and two new species of Stephania Loureiro, S. ornamenta sp. nov. and S. geniculata sp. nov., are recognized from Paleocene deposits of the Sanshui Basin, Guangdong, and a new occurrence of the widespread Eocene species Stephania auriformis (Hollick) Han & Manchester is recognized from the Maoming Basin, Guangdong. The Paleocene Stephania specimens described here represent the earliest fossil endocarp record of the Menispermaceae in eastern Asia. This discovery shows that the moonseed family had arrived in tropical and humid South China by at least the middle Paleocene, which provides important evidence for the origin and phytogeographic history of the family.     

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.     

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.     

LUTETIACADER, A PUZZLING NEW GENUS OF CANTACADERID LACE BUGS (HETEROPTERA: TINGIDAE) FROM THE MIDDLE EOCENE MESSEL MAAR, GERMANY

Abstract:  A new genus of cantacaderid lace bugs, Lutetiacader gen. et sp. nov., is described from the lower Middle Eocene of Messel Quarry, Germany. It clearly belongs to the cantacaderid lineage of Tingidae, and is the second known fossil genus of the tribe Cantacaderini (after Paleocader Froeschner). It appears to be most similar to extant species of the genera Nectocader Drake (Brazil) and Teratocader Drake (Malay Peninsula). Lutetiacader occupies a central position with respect to the geographical gap between the ranges of these two extant genera. Its biogeographical significance is discussed.     

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.     

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.     

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.     

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.     

Late Eocene leaves of Nageia (section Dammaroideae) from Maoming Basin,South China and their implications on phytogeography

A new species of Nageia, Nageia maomingensis sp. nov., is described from the Late Eocene of Maoming Basin, Guangdong Province, South China. According to its cuticular characteristics, the present fossil species is assigned to N. sect. Dammaroideae, showing the most resemblance to the extant species N. motleyi (Parl.) de Laub. At present, this section is mainly distributed in the Pacific islands region from South Asia to New Guinea. The discovery of N. maomingensis demonstrates that plants of this section inhabited South China at least by the Eocene. The distribution and living habitat of extant N. sect. Dammaroideae imply that South China was warmer during the Late Eocene than today, and may represent one of the centers for early diversification of Nageia and its section Dammaroideae during the Eocene. In addition, we postulate that plants of sect. Dammaroideae migrated southward in response to climate cooling after the Eocene.     

Establishing a fossil record for the perianthless Piperales: Saururus tuckerae sp. nov. (Saururaceae) from the Middle Eocene Princeton Chert

Investigations of small permineralized flowers from the Middle Eocene Princeton Chert, British Columbia, Canada have revealed that they represent an extinct species of Saururus. Over 100 flowers and one partial inflorescence were studied, and numerous minute perianthless flowers are borne in an indeterminate raceme. Each flower is subtended by a bract, and flowers and bracts are borne at the end of a common stalk. Five stamens are basally adnate to the carpels. Pollen is frequently found in situ in the anthers. Examined under SEM and TEM, pollen grains are minute (6-11 μm), monosulcate, boat-shaped-elliptic, with punctate sculpturing and a granulate aperture membrane. The gynoecium is composed of four basally connate, lobed carpels with recurved styles and a single ovule per carpel. Flower structure and pollen are indicative of Saururaceae (Piperales), and in phylogenetic analyses using morphological characters, the fossils are sister to extant Saururus. The fossil flowers are described here as Saururus tuckerae sp. nov. These fossil specimens add to the otherwise sparse fossil record of Piperales, represent the oldest fossils of Saururaceae as well as the first North American fossil specimens of this family, and provide the first evidence of saururaceous pollen in the fossil record.     

Anatomy and development of fruits of Lauraceae from the Middle Eocene Princeton Chert

Investigations of the Middle Eocene Princeton Chert reveal evidence for the connection of lauraceous flowers to fruits through a developmental series. Youngest fruits are found with attached floral remnants. Later stages show receptacle enlargement, fruit wall thickening, and the development of abundant sclereid clusters. Mature fruits are borne on a shallow receptacle and have an endocarp palisade layer of radially elongate cells with stellate outlines, an inner mesocarp layer of radiately arranged sclereid clusters, and a fleshy outer mesocarp layer containing numerous idioblasts with contents. Each mature fruit bears a single seed retaining the outer integument with an innermost radially elongate transfusion cell layer. Mature seeds contain a cellular embryo bearing idioblasts. Fruits are distinguishable from previously described anatomically preserved fossil taxa. This study represents the only documented developmental reconstruction of fossil fruits of Lauraceae and that self-pruning evolved prior to the Eocene. Anatomical modifications over the developmental sequence indicate that different stages of maturity preserved together, may be erroneously identified as several taxa at a fossil locality. Fossil morphotypes typically underestimate species number, but this study suggests that the number of inferred species based on fruit types may be inflated for Lauraceae, potentially exaggerating the tropical interpretation of the paleoenvironment.     

CAESALPINIA SUBGENUS MEZONEURON (LEGUMINOSAE,CAESALPINIOIDEAE) FROM THE TERTIARY OF NORTH AMERICA

Legume fruits from the Eocene of Tennessee and Wyoming and the Miocene of Idaho are described and assigned to Caesalpinia subgenus Mezoneuron (Caesalpinioideae), an extant Paleotropical taxon that does not occur in North or South America today. Morphological and anatomical details of the fruits are used in evaluating their systematic relationships. The features of the fossil fruits are accommodated only within this extant subgenus. These fossils represent the only reliable known occurrence of C. subgenus Mezoneuron in the paleobotanical record. These fossils suggest that subgenus Mezoneuron was distinct from subgenus Caesalpinia by the Middle Eocene. Further, they document the widespread occurrence of this currently Paleotropical group for at least 30 million years in North America.     

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.     

湖北丹江口早始新世梳趾鼠类一新属及伴生哺乳动物化石

描述了湖北省丹江口市均县镇一始新世化石点的3种哺乳动物化石,其中包括梳趾鼠类一新属种：林氏汉南鼠（新属、新种）Hannanomys lini gen. et sp. nov.。根据林氏汉南鼠与其他梳趾鼠类性状对比及该地点动物群组合特点推测其时代为早始新世晚期,与产于玉皇顶组中段的王家营大尖地方动物群的时代相当。该化石点是李官桥盆地中汉江南岸早始新世化石地点的首次发现,它的认定对在此地区寻找古新统/始新统界线有重要意义。     

EOCENE FLORAL EVIDENCE OF LAURACEAE: CORROBORATION OF THE NORTH AMERICAN MEGAFOSSIL RECORD

Numerous megafossils of Lauraceae have been reported from the early Tertiary of North America, but the subfamilial affinities are usually not well understood due to the great morphological variability found in extant taxa. The flowers of Androglandula tennessensis gen. et sp. nov. Taylor, from the Middle Eocene Claiborne Formation, are six-parted, pedicellate, bracteate, and have stamens with paired basal staminal glands. The flowers have ethereal oil cells and paracytic stomates throughout. The fossil species has affinities with the subtribe Cinnamomineae, and this supports suggestions that the Middle Eocene climate of the southeastern U.S. was subtropical. The existence of this fossil, and reports of the subtribe from the Eocene of Europe, indicate a South American-North American-European-southeast Asian paleodistribution suggesting that extinction in North America and Europe was the cause of the tribe's current disjunct distribution.