New carnivoraforms from the early Eocene of Europe and their bearing on the evolution of the Carnivoraformes

Two new mammalian carnivoraform species, Uintacyon hookeri sp. nov. and Quercygale smithi sp. nov., are described from the early Eocene of Europe. U. hookeri sp. nov. is recorded in Mutigny (MP8 + 9, PE IV), Avenay (MP8 + 9, PE V), Brasles, Condé‐en‐Brie (MP8 + 9) and Cuis (MP 10), while Q. smithi sp. nov. comes from Mutigny and Mancy (MP10). Because the two species are not recorded in earliest Eocene localities such as Dormaal and Le Quesnoy (MP7, PE I), it is proposed that they dispersed after the main phase of the Mammal Dispersal Event. U. hookeri sp. nov. supports the existence of terrestrial connections with North America, while Q. smithi sp. nov. implies possible faunal exchange with Asia. This evidence for the evolution of the Carnivoraformes supports: (1) a rapid decrease in their diversity after the Mammal Dispersal Event; and (2) the existence of a mammal turnover event in Europe during the early Eocene. The discovery of a new species of Quercygale, which is generally considered as the closest carnivoraform to the crown‐group Carnivora, shows that the genus had already lost the M₃ by the early Eocene and supports an important, but very poorly known, radiation of the carnivoraforms at least during the earliest early Eocene.     

New highlights about the enigmatic marine snake Palaeophis maghrebianus (Palaeophiidae; Palaeophiinae) from the Ypresian (Lower Eocene) phosphates of Morocco

Abstract: Palaeophis maghrebianus belongs to the Palaeophiinae (Palaeophiidae). This snake subfamily is relatively poorly known, and it is mainly represented by disarticulated vertebrae and ribs and by a few vertebral segments. Its intracolumnar variability remains also poorly understood. The discovery of new isolated vertebrae and vertebral segments of Palaeophis maghrebianus in the Ypresian (Lower Eocene) Phosphates of Morocco enables us to provide a more detailed diagnosis of this species and to describe its intracolumnar variability. Moreover, the new material reveals that this species could reach gigantic size being, with Palaeophis colossaeus, one of the two longer palaeophiids. The microanatomical and histological analysis of some vertebrae illustrating diverse positions along the vertebral column reveals the presence of osteosclerosis, especially in the anterior and mid‐precloacal regions. The occurrence of this osseous specialization implies a role in buoyancy and body trim control in this taxon, which is considered a shallow marine dweller based on its anatomical features and geological data. Palaeophis maghrebianus also displays a dense vascular network suggesting a growth speed, and thus a metabolic rate, much higher than in the biggest extant snakes.     

New fossil Hyaenodonta (Mammalia,Placentalia) from the Ypresian and Lutetian of France and the evolution of the Proviverrinae in southern Europe

The proviverrines from the Ypresian (MP7–MP10) and Lutetian (MP11–MP14) are represented mainly by species recorded in the northern and central parts of Europe (Paris Basin, Belgian Basin, Germany, Switzerland). Here, we describe fossils from southern France: Saint‐Papoul (MP8 + 9; Aude) and Aigues‐Vives 2 (?MP13; Aude). One dentary with secant molars from Saint‐Papoul represents a new genus and species, Preregidens langebadrae. This taxon is possibly present in Avenay (France), the MP8 + 9 reference locality. One of the three dentaries discovered in Aigues‐Vives 2 belongs to the hypercarnivorous Oxyaenoides schlosseri, previously represented by only two isolated lower molars. This dentary appears to be the most derived of the proviverrines. This species is possibly present in Saint‐Martin‐de‐Londres (France), a locality that is considered to be close to the MP13 reference level. The two other dentaries from Aigues‐Vives 2 support the presence of Eurotherium theriodis and provide the first possible evidence of sexual dimorphism in a proviverrine species. A phylogenetic analysis of the proviverrines is performed to resolve the phylogenetic position of the three taxa. This identifies a close relationship between the new genus (Preregidens) and Oxyaenoides. The new fossils allow the age of Saint‐Papoul and Aigues‐Vives 2 to be refined: the first locality is considered to be close in age to Avenay (Ypresian; France), while the second one seems to be close to Egerkingen γ (Lutetian; Switzerland), which is considered to be possibly close in age to the MP13 reference level. Finally, the presence of O. schlosseri and E. theriodis in the southern part of France is compatible with the hypothesis that the mammals involved in the first intra‐Eocene turnover migrated northwards.     

New material of Diacodexis (Mammalia,Artiodactyla) from the early Eocene of Southern Europe

Diacodexeidae are the first representatives of Artiodactyla in the fossil record. Their first occurrence is at the very base of the Ypresian (earliest Eocene, 56.0 Ma) with Diacodexis, a genus well diversified during the early Eocene in Europe, especially during the MP7–MP8 + 9 interval. However, most of European species are documented by scarce material, retrieved from single localities. In this work, we describe new Diacodexis material from ～MP7 and ～MP8 + 9 localities of Southern Europe, including material of D. antunesi from Silveirinha, considered as the most primitive European Diacodexis species, and material from three localities from Southern France (Fordones, Palette, and La Borie). The new material documents Diacodexis premolar morphology and deciduous dentition which bear potentially important phylogenetic information, as well as astragali, including a specimen from Silveirinha that constitutes the earliest occurrence of an astragalus of the genus Diacodexis in the European fossil record. Investigation of the enamel microstructure reveals that early European species had a simple enamel pattern with one-layered Schmelzmuster composed of ‘basic’ radial enamel only, instead of the two-layered Schmelzmuster (thin radial enamel + thick layer of Hunter-Schreger bands) observed on North American species and so far considered to represent the primitive condition within Artiodactyla. In accordance with previous studies, our observations highlight that Diacodexis gigasei from Belgium is morphologically closer to the North American species D. ilicis than to D. antunesi from Portugal. The latter species, together with D. aff. antunesi from Fordones, appears to be morphologically closer to the Asiatic taxa D. indicus and D. pakistanensis. Finally, we found numerous similarities between D. cf. gigasei from Palette and D. gigasei, a result that challenges the intra-European provincialism that characterizes the earliest Ypresian. Diacodexis gigasei could be one of the rare species shared by the northwestern and southwestern European bioprovinces.     

Sinopaninae and Arfianinae (Hyaenodontida,Mammalia) from the Early Eocene of Europe and Asia; evidence for dispersal in Laurasia around the Paleocene/Eocene boundary and for an unnoticed faunal turnover in Europe

Hyaenodontida are represented in Europe by three subfamilies: Proviverrinae, Arfianinae and Sinopaninae. Here, we review all the specimens of Arfianinae and Sinopaninae known to date in Europe and Asia. A new Galecyon species is erected: Galecyon gallus nov. sp. We discuss the taxonomic position of the two Asian hyaenodontidans Anthracoxyaena palustris and Arfia langebadreae; the genus Anthracoxyaena is synonymized with Arfia. The analysis of the European and Asian arfianines and sinopanines provides new data concerning the dispersals and faunal events that occurred during the Early Eocene in Laurasia. The Arfianinae and Sinopaninae appeared in Europe around the Paleocene/Eocene boundary (reference-level MP7). The sinopanines are widespread in Europe; they are known in Dormaal (Belgium, reference-locality of the level MP7), Rians, Soissons, Pourcy, Try, Le Quesnoy (France), and Abbey Wood (England). The analysis of the paleogeographic distribution of all Oxyaenodonta and Hyaenodontida at and after the MP7 supports the existence of two European provinces: the North Province and Mesogean Province. We show that the Arfianinae and Sinopaninae rapidly disappeared from Europe; they are unknown in Avenay (reference-locality of the level MP8+9) and younger localities. Their disappearance from Europe is synchronous with that of the Oxyaenodonta. These observations support the existence of a faunal turnover, which occurred between the reference-levels MP7 (Dormaal) and MP8+9 (Avenay). The hypothesis of a dispersal from Europe to North America during the Paleocene-Eocene transition for the Arfianinae and Sinopaninae is supported. Moreover, the study of Arfianinae supports a dispersal from Europe to Asia around the P/E boundary, followed by a short period of endemic evolution. However, our study does not support a close relationship between Arfia and the “Arfia-like South Asian Proviverrinae” (Kyawdawia, Indohyaenodon, Paratritemnodon and Yarshea).     

New remains of the Eocene Prophaethon and the early evolution of tropicbirds (Phaethontiformes)

A partial skeleton of the early Eocene tropicbird Prophaethon shrubsolei (Prophaethontidae) from the London Clay of Walton‐on‐the‐Naze (UK) is described. Most of the bones preserved in this fossil have not been described previously for Prophaethon or prophaethontids in general. The pelvis and leg bones of Prophaethon, including the previously unknown tarsometatarsus, are very different from modern tropicbirds and show a strong similarity to procellariiform birds, especially albatrosses. Rather than employing plunge‐diving like their extant relatives, prophaethontids therefore probably used foraging strategies similar to those of modern albatrosses, which seize food on the sea surface. Prophaethontids also appear to have been less pelagic than extant tropicbirds, and these different life modes, as well as the disappearance of Phaethontiformes from northern latitudes, attest to major changes in Northern Hemispheric marine avifaunas during the Cenozoic, which also affected other pelagic birds. The reasons for the profound changes in the historical biogeography and way of living of tropicbirds are unknown, but ecological competition and predation at breeding sites are likely to have played a role.     

New notharctine primate fossils from the early Eocene of New Mexico and southern Wyoming and the phylogeny of Notharctinae

Previously undescribed notharctine primate fossils are reported from the early Eocene San Jose Formation, San Juan Basin, New Mexico, and the early Eocene Wasatch Formation, southern Wyoming. These collections include the most complete specimens yet discovered of the poorly known species Copelemur tutus and Copelemur praetutus; the first upper dentitions of Cantius angulatus and Cantius frugivorus from the type area of these taxa; and fossils attributable to two new notharctine species, Copelemur australotutus and Smilodectes gingerichi. These new fossils reveal that current ideas concerning notharctine phylogeny are incorrect. Two major, monophyletic clades are apparent within the subfamily: the tribe Copelemurini, consisting of the genera Copelemur and Smilodectes, and the tribe Notharctini, comprising the genera Cantius, Pelycodus, and Notharctus. Analysis of the paleobiogeographic distribution of the Copelemurini indicates that this clade was limited to more southerly regions of western North America during early Eocene time. Northward migration of more tropical habitats during the late Wasatchian and early Bridgerian in western North America, associated with an overall climatic warming trend through the early and middle Eocene, appears to have allowed several mammalian taxa, including Smilodectes, to extend their ranges northward during this time interval. Such taxa thus possess diachronous distributions and have been partly responsible for the long-standing confusion regarding the biostratigraphic correlation of early Eocene faunas from New Mexico with those from Wyoming. Based on several taxa which are also known from the Wasatchian of Wyoming, the age of the San Jose Formation appears to be middle Wasatchian.     

New early Eocene anaptomorphine primate (Omomyidae) from the Washakie Basin,Wyoming, with comments on the phylogeny and paleobiology of anaptomorphines

Recent paleontological collecting in the Washakie Basin, southcentral Wyoming, has resulted in the recovery of over 100 specimens of omomyid primates from the lower Eocene Wasatch Formation. Much of what is known about anaptomorphine omomyids is based upon work in the Bighorn and Wind River Basins of Wyoming. This new sample documents greater taxonomic diversity of omomyids during the early Eocene and contributes to our understanding of the phylogeny and adaptations of some of these earliest North American primates. A new middle Wasatchian (Lysitean) anaptomorphine, Anemorhysis savagei, n. sp., is structurally intermediate between Teilhardina americana and other species of Anemorhysis and may be a sister group of other Anemorhysis and Trogolemur. Body size estimates for Anemorhysis, Tetonoides, Trogolemur, and Teilhardina americana indicate that these animals were extremely small, probably less than 50 grams. Analysis of relative shearing potential of lower molars of these taxa indicates that some were primarily insectivorous, some primarily frugivorous, and some may have been more mixed feeders. Anaptomorphines did not develop the extremes of molar specialization for frugivory or insectivory seen in extant prosimians. Incisor enlargement does not appear to be associated with specialization in either fruits or insects but may have been an adaptation for specialized grooming or food manipulation. © 1994 Wiley-Liss, Inc.     

Curtisia (Cornales) from the Eocene of Europe and its phytogeographical significance

Fossil remains of Curtisia Aiton (Cornales) are recognized for the first time from the Tertiary of Europe, based on early Eocene anatomically preserved fruits from the London Clay and Poole Formations of southern England. The modern distribution of this monotypic genus is limited to the cape of South Africa. Curtisia quadrilocularis (Reid & Chandler) comb. nov. fruits have globose tetralocular endocarps composed of isodiametric sclereids with a single seed per locule, a prominent axial vascular canal, apical placentation, and four germination valves. All of these characters, as well as size, correspond to extant Curtisia . Although many fossil taxa from the Eocene of Europe have been shown to have their closest extant relatives in Asia, this occurrence of Curtisia highlights Tertiary floristic exchange between Europe and Africa. The newly recognized fossil occurrences suggest a Laurasian origin for Curtisia , in conformity with the fossil record for several other genera of the Cornales. In addition, our rejection of the former assignment of this species to Leucopogon causes us to question whether Epacridaceae were present in the Tertiary of Europe.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 127–134.     

Morphofunctional analysis of Svobodaina species (Brachiopoda,Heterorthidae) from south‐western Europe

Geometric morphometric methods applied to the ventral muscle field outline of the various species of Svobodaina from south‐western Europe identify criteria for discriminating among this key cluster of brachiopod species. These data indicate the close relationship between the patterns of the ventral muscle field in the Svobodaina species and the environmental conditions where each inhabited; the fields are better developed in species related to high‐energy environments. For example, S. armoricana, with the smallest diductor scars, would have inhabited the quiet marine environments of the lower offshore or within protected lagoonal settings. S. feisti would have inhabited the upper offshore, a more energetic environment than S. armoricana. Finally, S. havliceki, with the largest diductor scars, would have thrived in the most energetic environments among all the south‐western European Svobodaina species, living just above the fair‐weather wave base in the lower shoreface. The palaeoecological results suggest a distribution of Svobodaina species during the Late Ordovician along an onshore–offshore transect across the shallow marine platforms of the Mediterranean margin of Gondwana. On the other hand, the occurrence in some localities of several species with overlapping ranges or within the same assemblage indicates that the biostratigraphical efficacy of the genus is restricted. Thus, the previously defined taxon‐range biozones characterized by Svobodaina species of the north Gondwanan margin are in need of reassessment. The morphology of Svobodaina may be a considerable aid to environmental analyses rather than to precise biostratigraphical correlations.     

Silurian bolbozoids and cypridinids (Myodocopa) from Europe: pioneer pelagic ostracods

Abstract: Bolbozoid and cypridinid myodocope ostracods from France, the Czech Republic and Sardinia consist of four genera (one new: Silurocypridina) and at least ten species (eight new: Bolbozoe acuta, B. rugosa, B. parvafraga, Parabolbozoe armoricana, Silurocypridina retroreticulata, S. variostriata, S. calva and Calocaria robusta). They have a functional design, palaeogeographical distribution, depositional setting and faunal associates that suggest that they were swimmers in the water column, living in dim light, relatively deep environments, above near bottom dysoxic/anoxic conditions. These myodocope faunas include some possible pelagic species sensu stricto, with widespread palaeogeographical (transoceanic) distribution, and some species that occupied hyperbenthic niches and were capable of making short incursions to the sediment to scavenge on carrion. These data support the model that Silurian myodocopes were pioneer pelagic ostracods.     

New ceratomorph perissodactyls (Mammalia) from the Middle and Late Eocene of Mongolia: their implications for phylogeny and dating

Eight species rank taxa arc described from Eocene sites in the Gobi Desert, Mongolia. Irdinolophus gen. now is erected for Desmalotherium mongoliense Osborn and for Irdinolophus? tuiensis sp. nov. The genus is placed at the stem of the family Depcrctellidae. Intraspecific variation in lekolophus magnus is documented for the first time; Pachylophus is synonymized with leleoloplms; and Deperetella cf.birmanica is recorded far north of its previously known range in Burma. The first known lower teeth of Colodon inceptus support referral ofDesmalotherium Jissum to Colodon. The stratigraphic range of the hyracodontid Ardynia is extended back from the late Eocene to the early middle Eocene. A range of primitive tapiroids and rhinocerotoids arc analysed cladistically to resolve the problematic affinities of the extinct families Dc-peretellidae and Rhodopagidae. The Depcrctellidae are shown to belong to a monophyletic superfamily Tapiroidca, whilst the Rhodopagidae belongs to the Rhinocerotoidea, but is distinct from the Hyracodontidae. Veragromovia, previously synonymized with the tapiroid Helaleles, is resurrected as a valid genus within the Rhodopagidae. Tsagan Khutcl and other sites in the Kholbolchi formation may be older than Irdinmanhan faunas in Asia, equaling with those from the Arshanto Formation of China of probable early Lutetian or latest Ypresian age. The Irdinmanhan Mergen fauna may be slightly older than that from Irdin Manha, because of the primitive aspect of its Lophialetes.     

A new basal caniform (Mammalia: Carnivora) from the middle Eocene of North America and remarks on the phylogeny of early carnivorans

Background

Despite a long history of research, the phylogenetic origin and initial diversification of the mammalian crown-group Carnivora remain elusive. Well-preserved fossil materials of basal carnivorans are essential for resolving these issues, and for constraining the timing of the carnivoran origin, which constitutes an important time-calibration point in mammalian phylogenetics.

Methodology/Principal Findings

A new carnivoramorphan from the middle Eocene of southern California, Lycophocyon hutchisoni, is described. The new taxon exhibits stages of dental and basicranial evolution that are intermediate between earlier carnivoramorphans and the earliest representatives of canoid carnivorans. The evolutionary affinity of the new taxon was determined by a cladistic analysis of previously-published and newly-acquired morphological data for 30 Paleogene carnivoramorphans. The most-parsimonious trees identified L. hutchisoni as a basal caniform carnivoran, and placed (1) Tapocyon robustus, Quercygale angustidens, “Miacis” sylvestris, “M.” uintensis, and “M.” gracilis inside or outside the Carnivora, (2) nimravids within the Feliformia, and (3) the amphicyonid Daphoenus outside the crown-group Canoidea. Parsimony reconstructions of ancestral character states suggest that loss of the upper third molars and development of well-ossified entotympanics that are firmly fused to the basicranium (neither condition is observed in L. hutchisoni) are not associated with the origin of the Carnivora as traditionally thought, but instead occurred independently in the Caniformia and the Feliformia. A discriminant analysis of the estimated body weight and dental ecomorphology predicted a mesocarnivorous diet for L. hutchisoni, and the postcranial morphology suggests a scansorial habit.

Conclusions/Significance

Lycophocyon hutchisoni illuminates the morphological evolution of early caniforms leading to the origin of crown-group canoids. Considerable uncertainty remains with respect to the phylogenetic origin of the Carnivora. The minimum date of caniform-feliform divergence is provisionally suggested to be either 47 million years ago or 38 million years ago, depending on the position of “Miacis” sylvestris within or outside the Carnivora, respectively.     

A view of early vertebrate evolution inferred from the phylogeny of polystome parasites (Monogenea: Polystomatidae)

The Polystomatidae is the only family within the Monogenea to parasitize sarcopterygians such as the Australian lungfish Neoceratodus poisteri and freshwater tetrapods (lissamphibians and chelonians). We present a phylogeny based on partial 18S rDNA sequences of 26 species of Polystomatidae and three taxon from the infrasubclass Oligonchoinea (= Polyopisthocotylea) obtained from the gills of teleost fishes. The basal position of the polystome from lungfish within the Polystomatidae suggests that the family arose during the evolutionary transition between actinopterygians and sarcopterygians, ca. 425 million years (Myr) ago. The monophyly of the polystomatid lineages from chelonian and lissamphibian hosts, in addition to estimates of the divergence times, indicate that polystomatids from turtles radiated ca. 191 Myr ago, following a switch from an aquatic amniote presumed to be extinct to turtles, which diversified in the Upper Triassic. Within polystomatids from lissamphibians, we observe a polytomy of four lineages, namely caudatan, neobatrachian, pelobatid and pipid polystomatid lineages, which occurred ca. 246 Myr ago according to molecular divergence-time estimates. This suggests that the first polystomatids of amphibians originated during the evolution and diversification of lissamphibian orders and suborders ca. 250 Myr ago. Finally, we report a vicariance event between two major groups of neobatrachian polystomes, which is probably linked to the separation of South America from Africa ca. 100 Myr ago.     

New hyaenodonta (Mammalia) from the middle Eocene of Myanmar

The middle Eocene Pondaung Formation in Myanmar has yielded a rich mammalian fauna including several Primate taxa. Hyaenodonta are known by the genera Kyawdawia, Yarshea, Orienspterodon, and two other indeterminate taxa. We describe here new material of Kyawdawia, including some morphological details, a new species of the hypercarnivorous genus Propterodon and an indeterminate species, different from those described earlier in Myanmar, and characterized by a reduction of m3 and would belong to a third lineage with the same evolutionary trend as Galecyon and the Limnocyoninae. The hyainailourines (Orienspterodon) and hyaenodontines (Propterodon) are recorded for the first time in Southeast Asia and these subfamilies appeared in quasi the same time in Europe illustrating probably a profound change in the carnivorous fauna among Laurasia.     

Dietary adaptations and palaeoecology of Lophialetidae (Mammalia,Tapiroidea) from the Eocene of the Erlian Basin,China: combined evidence from mesowear and stable isotope analyses

Lophialetidae is an extinct group of endemic Asiatic tapiroids that are widely distributed in the Eocene sediments of Asia. Schlosseria magister and Lophialetes expeditus are the most abundant species in this family. However, their dietary and ecological characteristics are largely unknown. For the first time, we reconstruct the palaeodiet and habitat of these two lophialetids using a combination of mesowear and stable carbon isotope analysis of fossil teeth excavated from the Erlian Basin, China. Mesowear analysis (n = 141) suggests that the dietary structure of S. magister and L. expeditus shifted from less to more abrasive diets from ~52 to ~42 Ma. Stable carbon isotope analysis (n = 137) suggests that the habitats of S. magister and L. expeditus became drier and/or more open through time. The dietary shifts of the two lophialetids are consistent with evident changes in habitat. The changes in the diet and habitat were probably related to global climate change during that time period. The gradual drop in global temperatures during the early–middle Eocene led to a drier and more open terrestrial ecosystem in the Erlian Basin, probably resulting in changes in floral composition of the environment inhabited by S. magister and L. expeditus. Hence, herbivores highly susceptible to vegetation modification had to develop new resource exploitation strategies to adapt to these changes. Schlosseria magister, considered to be the sister-group of L. expeditus and with a low level of ecological flexibility, was unable to adapt to the habitat changes finally becoming extinct at ~45 Ma.     

Giant ants from the Paleogene of Denmark with a discussion of the fossil history and early evolution of ants (Hymenoptera: Formicidae)

Pachycondyla rebekkae sp. nov. (Formicidae, Ponerinae) is described from the Fur and Ø1st Formations (transitional between Paleocene and Eocene, Denmark). About 95 complete or fragmentary gyne specimens and one almost complete male specimen were examined, revealing characters which permit classification and placement of the fossil species in the phylogenetic system of extant Formicidae. The palaeoecology and taphonomy of the insect fauna of the Fur and Ø1st Formations are discussed. Finally, the significance of the new species is discussed in light of the fossil history and early evolution of ants.     

A new species of Limnofregata (Pelecaniformes: Fregatidae) from the Early Eocene Wasatch Formation of Wyoming: implications for palaeoecology and palaeobiology

A humerus and a coracoid from the Early Eocene Wasatch Formation in the Washakie Basin of south‐western Wyoming are the oldest materials (by ~2 million years) of the pelecaniform Limnofregata (Aves) and represent a new large species, Limnofregata hutchisoni sp. nov. This fossil is the oldest known member of the frigatebird lineage. Other than its large size relative to Limnofregata azygosternon and L. hasegawai, the new material is very similar morphologically to other known Limnofregata specimens. The size of this new species is comparable to the largest living species (e.g. Fregata minor and Fregata magnifiscens) and much larger than the two described species of Limnofregata. This fossil indicates that the hard minimum date previously advocated for molecular calibration of the split between Fregatidae and Suloidea is an underestimate by approximately two million years. The presence of early pelecaniform bird lineages (represented by Limnofregata and Masillastega) in limnic ecosystems prior to their known occurrences in marine deposits/habitats appears to indicate that some clades of pelecaniform birds may have undergone an evolutionary transition from freshwater to marine habitats in a pattern reminiscent of what has been suggested during the evolution of pinnipeds or that their palaeoecology included broader niches ranging across a variety of aquatic habitats. That transition in habitat occupation and the origin of many of the characteristic biological aspects present in the crown frigatebird clade likely occurred during a significant temporal gap (> 45 million years) in the fossil record of the frigatebird lineage after these earliest occurrences in the Early Eocene and before the oldest records of the extant Fregata species in the Pleistocene.     

Dispersed cuticles from the Eocene of North America

KOVACH, W. L. & DILCHER, D. L., 1984. Dispersed cuticles from the Eocene of North America. Macerations of organic-rich clay from the Claiborne Formation (Middle Eocene) of Tennessee have yielded a wide variety of well-preserved dispersed cuticles. Details of the epidermal cell patterns, arrangement of the stomata1 complex, trichomes and trichome attachment have made possible the association of some dispersed cuticles with leaf types known from this formation which have similar cuticle, and with modern families. All dispersed cuticles are classified in a morphologic system. Through both our own work and a review of previous investigations we have found that distinct dispersed cuticle types can be recognized and may be used biostratigraphically to characterize geologic strata, palaeo-ecologically to provide insights into environmental reconstruction, and systematically to follow the history of certain taxa.