Early Paleogene decapod crustaceans from the Sulaiman and Kirthar Ranges,Pakistan

A set of Paleocene and Eocene decapod crustaceans is described from the Sulaiman and Kirthar Ranges of Pakistan. The fossil crabs Proxicarpilius planifrons Collins and Morris, 1978 and Pakicarcinus orientalis (Collins and Morris, 1978), already known in the Eocene of northern Pakistan, are reported for the first time in the Paleocene of southern Pakistan, enlarging the stratigraphic and the palaeobiogeographical ranges of these species. The callianassid genus Calliax de Saint Laurent, 1973 is reported for the first time in the Paleocene of southern Pakistan; this is the oldest record for the genus.     

Diverse fossil Onagraceae pollen from a Miocene palynoflora of north-east China: early steps in resolving the phytogeographic history of the family

The origin and evolution of angiosperms can be unravelled by using fossil records to determine first occurrences and phytogeographic histories of plant families and genera. Many angiosperm families, for example the Onagraceae, have a poor macrofossil record, but are more common in palynological records. Modern Onagraceae produce pollen clearly distinct from that of other angiosperms. Combined morphological features obtained by use of light and scanning electron microscopy have enabled assignment of fossil Onagraceae pollen to extant genera, and therefore tracing of the origin and past distributions of extant Onagraceae lineages. We studied a Miocene palynoflora from the Daotaiqiao Formation of north-east China. Using the single-grain technique, we examined individual Onagraceae pollen/tetrads using both light and scanning electron microscopy. Fossil Onagraceae pollen is more frequent than macrofossil remains, but is still rare, and usually represented by a single taxon in palynological samples. Remarkably, samples from the Miocene of north-east China contain five different species: two of Circaea, one of Epilobium, and two of Ludwigia. Such a large number of Onagraceae taxa from a single palynoflora is unknown elsewhere. Whereas Ludwigia pollen is known from Cenozoic sediments of the northern hemisphere, the Circaea pollen is the first fossil pollen assignable to this extant genus. This is also the first fossil record of Epilobium from China. Although the young geological age of this sample does not enable consideration of time of origin for the genera encountered, the co-occurrence of Circaea, Epilobium, and Ludwigia in the mid to late-Miocene of East Asia sheds some light on their phytogeographic histories.     

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.     

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.     

MUSOPSIS N. GEN.: A BANANA-LIKE LEAF GENUS FROM THE EARLY TERTIARY OF EASTERN NORTH GREENLAND

A fossil flora from the Late Paleocene-Early Eocene Thyra Ø Formation of eastern North Greenland (paleolatitude 77° N) has yielded monocotyledon leaf impressions with characters seen only in the closely related modem species in the families of Heliconiaceae, Musaceae, and Strelitziaceae. The combination of large costae widths and parallel, nonanastomosing, lateral veins that depart at right angles from the costae in the fossil material are features present only in leaves of extant species from these families. Three basic venation patterns also are recognized in the modem species of these families, but except for the genera Strelitzia and Phenakospermum, none of these patterns are present exclusively in any one family. Musopsis n. gen. is created for the fossil material from Greenland, but it is considered a form genus due to the lack of gross morphological features that can be used for separating leaves of the modem genera in Heliconiaceae, Musaceae, and Strelitiziaceae. It is the first known Arctic occurrence of fossil leaf material resembling this modem group of taxa.     

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.     

Middle Eocene Pelagornithidae and Gaviiformes (Aves) from the Ukrainian Paratethys

Abstract: We describe avian remains from Novopskov, a new middle Eocene marine locality in Ukraine. The fossils constitute the most substantial collection of Palaeogene bird bones from Eastern Europe and contribute to a better knowledge of the Paratethyan seabird fauna. Most of the specimens belong to Pelagornithidae (bony‐toothed birds), and two species of very different size can be distinguished. The larger of these is tentatively referred to Dasornis sp., the smaller to Odontopteryx toliapica. The specimens include skeletal elements that were not described for Palaeogene bony‐toothed birds and document previously unknown morphological differences between Palaeogene and Neogene Pelagornithidae. It is argued that the purported crane Eobalearica tugarinovi, from the middle Eocene of Kyrgyzstan, is probably also a bony‐toothed bird. A new genus and species of small Gaviiformes, Colymbiculus udovinchenkoi, is described, which is the earliest fossil record of a loon from Europe, preceding the next oldest specimens by more than 10 myr. The Ukrainian fossils document profound differences between middle Eocene and extant marine avifaunas of Europe, and whereas the middle Eocene Paratethyan avifauna appears to have been similar to that of the North Sea with regard to pelagornithid diversity, the absence of prophaethontids and relative abundance of Gaviiformes may indicate faunistic differences concerning the remaining seabirds.     

Evolution and biogeography of the austral genus Phyllocladus (Podocarpaceae)

Aim To infer evolutionary relationships within the genus Phyllocladus and among its close relatives by phylogenetic analysis of DNA sequences. Interpret the inferred relationships in association with the fossil record to examine the origin and diversification of the genus. Location Australasia. Methods Phylogenetic analyses of rbcL, matK and internal transcribed spacer (ITS) sequences representing all of the extant species of Phyllocladus and a selection of outgroups from Podocarpaceae and Araucariaceae. Results The rbcL and matK sequences exhibit little variation within Phyllocladus, but ally its members to Podocarpaceae although its immediate sister remains unclear. The ITS sequences resolve all five species of Phyllocladus and two intraspecific ecotypes of P. alpinus. Main conclusions Phyllocladus forms a distinct lineage that diverged early in the evolutionary history of Podocarpaceae. The fossil record indicates that the genus was more widely distributed and morphologically diverse during the early Tertiary than at present. Although of Mesozoic origin, the level of sequence variation within Phyllocladus suggests that the extant species radiated during the late Tertiary c. 6.3 ± 0.9 Ma. New Zealand is the present centre of species diversity.     

The fossil record of extant elasmobranchs

Sharks and their relatives (Elasmobranchii) are highly threatened with extinction due to various anthropogenic pressures. The abundant fossil record of fossil taxa has allowed the tracing of the evolutionary history of modern elasmobranchs to at least 250 MYA; nonetheless, exactly how far back the fossil record of living taxa goes has never been collectively surveyed. In this study, the authors assess the representation and extent of the fossil record of elasmobranchs currently living in our oceans by collecting their oldest records and quantifying first appearance dates at different taxonomic levels (i.e., orders, families, genera and species), ecological traits (e.g., body size, habitat and feeding mechanism) and extinction risks (i.e., threatened, not threatened and data deficient). The results of this study confirm the robust representation of higher taxonomic ranks, with all orders, most of the families and over half of the extant genera having a fossil record. Further, they reveal that 10% of the current global species diversity is represented in the geological past. Sharks are better represented and extend deeper in time than rays and skates. While the fossil record of extant genera (e.g., the six gill sharks, Hexanchus) goes as far back as c. 190 MYA, the fossil record of extant species (e.g., the sand shark, Carcharias taurus Rafinesque 1810) extends c. 66 MYA. Although no significant differences were found in the extent of the fossil record between ecological traits, it was found that the currently threatened species have a significantly older fossil record than the not threatened species. This study demonstrate that the fossil record of extant elasmobranchs extends deep into the geologic time, especially in the case of threatened sharks. As such, the elasmobranch geological history has great potential to advance the understanding of how species currently facing extinction have responded to different stressors in the past, thereby providing a deep-time perspective to conservation.     

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.     

Hepatics from Rovno amber (Ukraine), 6. Frullania rovnoi,sp. nov.

A fossil species of the extant liverwort genus Frullania Raddi is described and illustrated, based on a single inclusion in a piece of Rovno amber (Ukraine) that shares its age with Late Eocene Baltic amber, its northern contemporary. Frullania rovnoi is characterised by leaves with a rounded dorsal lobe and the absence of ocelli. The ventral lobe is inflated and forms a saclike lobule, which is bell-shaped and somewhat constricted above the mouth. The bifid underleaves have several blunt teeth or angulations along the shoulder. The Rovno fossil differs sufficiently from morphologically similar species preserved in Baltic and Bitterfeld amber as to be described as new to science. The shape of the lobules and underleaves, as well as the absence of ocelli, indicate an affiliation to F. sect. Australes, hitherto represented in Eocene amber inclusions solely by F. schumannii (Casp.) Grolle. The Rovno fossil is distinguished from extant species of F. subg. Australes and from F. schumannii by having roughly and irregularly dentate-angulate underleaf margins.     

Systematics and evolution of the Pan‐Alcidae (Aves,Charadriiformes)

Puffins, auks and their allies in the wing‐propelled diving seabird clade Pan‐Alcidae (Charadriiformes) have been proposed to be key pelagic indicators of faunal shifts in Northern Hemisphere oceans. However, most previous phylogenetic analyses of the clade have focused only on the 23 extant alcid species. Here we undertake a combined phylogenetic analysis of all previously published molecular sequence data (～ 12 kb) and morphological data (n = 353 characters) with dense species level sampling that also includes 28 extinct taxa. We present a new estimate of the patterns of diversification in the clade based on divergence time estimates that include a previously vetted set of twelve fossil calibrations. The resultant time trees are also used in the evaluation of previously hypothesized paleoclimatic drivers of pan‐alcid evolution. Our divergence dating results estimate the split of Alcidae from its sister taxon Stercorariidae during the late Eocene (～ 35 Ma), an evolutionary hypothesis for clade origination that agrees with the fossil record and that does not require the inference of extensive ghost lineages. The extant dovekie Alle alle is identified as the sole extant member of a clade including four extinct Miocene species. Furthermore, whereas an Uria + Alle clade has been previously recovered from molecular analyses, the extinct diversity of closely related Miocepphus species yields morphological support for this clade. Our results suggest that extant alcid diversity is a function of Miocene diversification and differential extinction at the Pliocene–Pleistocene boundary. The relative timing of the Middle Miocene climatic optimum and the Pliocene–Pleistocene climatic transition and major diversification and extinction events in Pan‐Alcidae, respectively, are consistent with a potential link between major paleoclimatic events and pan‐alcid cladogenesis.     

First fossil record of the genus Nageia (Podocarpaceae) in south China and its phytogeographic implications

The first fossil record of the genus Nageia Gaertner from the Tertiary of southern China is reported. Nageia hainanensis sp. nov. is described from the Eocene Changchang Formation of Hainan Island. The extant species of Nageia are widely distributed in southeastern Asia, from northeastern India to southeastern China and southern Japan, and southward through Indochina to the Malay Archipelago and New Guinea. The fossil evidence of Nageia has been discovered from the Lower Cretaceous of the Far East of Russia and Japan. The distribution of both modern and fossil species of Nageia indicates that this genus could have originated in the northeastern part of Asia in the Early Cretaceous and spread to south China at least in the Eocene. Discovery of Nageia fossil record in south China provides important evidence for the geological and phytogeographic history of the genus.     

Marsileaceaephyllum, a new genus for marsileaceous macrofossils: leaf remains from the Early Cretaceous (Albian) of southern Gondwana

Three new taxa from Albian, Early Cretaceous assemblages in Gondwana (Australia and Antarctica) and two previously described fossils from the Late Cretaceous and Eocene of North America are attributable to the heterosporous semi-aquatic fern family Marsileaceae. They are assigned to Marsileaceaephyllum, a morphotaxon erected here for sterile remains (whole plants, and isolated leaves and leaflets) of Marsileaceae. The Gondwanan taxa, Marsileaceaephyllum lobatum and Marsileaceaephyllum spp. B-C, have either a cruciform leaflet arrangement or dichotomous and anastomosing venation characteristic of modern Marsileaceae. Two previously established taxa, Marsilea johnhallii and Marsilea sp., which represent sterile Marsileaceae, are also transferred to the new genus (now Marsileaceaephyllum johnhallii and Marsileaceaephyllum sp. A, respectively). Examination of all fossil venation patterns reveals four new venation types not present in extant taxa, suggesting that most fossil Marsileaceae (leaves) are distinct from extant genera, and are likely members of extinct lineages. This is further supported by the absence of modern megaspore types in the Early Cretaceous.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: A CATKIN WITH JUGLANDACEOUS AFFINITIES

Three specimens of one type of fossil catkin from the Middle Eocene of Tennessee are excellently preserved and have been investigated morphologically. The flowers on these catkins are subtended by elongate, three-lobed bracts, are exclusively staminate, and have three conspicuous, obovate, perianth parts that bear large peltate scales. The stamens are well preserved and contain triporate pollen grains that are equivalent to the dispersed pollen genus Momipites. Floral morphology, cuticular features, and pollen indicate close affinities with the extant genera Engelhardia, Oreomunnea, and Alfaroa of the Juglandaceae; but because the fossil catkins are distinct and are a dispersed plant organ, they are placed in a new form genus: Eokachyra. These fossil flowers represent a rare opportunity to correlate the micro- and macrofossil record and to compare the relative rates of evolution of these features. The fossil catkins also demonstrate that much structural information may be gained from the study of fossil angiosperm flowers. The similarities between the staminate flowers of the fossil catkins and the staminate flowers of Engelhardia, Oreomunnea, and Alfaroa confirm the idea that this complex has had a long evolutionary history and suggest that the pollination system of certain extant genera was well developed during Middle Eocene times.     

Out of the Neotropics: newly discovered relictual species sheds light on the biogeographical history of spider ants (Leptomyrmex,Dolichoderinae, Formicidae)

Spider ants of the genus Leptomyrmex Mayr (Hymenoptera: Formicidae: Dolichoderinae) are conspicuous species of Australasian rainforests, with putative fossil relatives in the Neotropics and Europe. There is longstanding debate over the biogeographical history of the genus, with the Palaearctic and Neotropical regions proposed as alternate centres of origin. We propose a resolution of this debate with the recent discovery and analysis of an extant species from central Brazil, L. relictus sp.n. , which we describe from workers, males and brood. We sequence ten nuclear genes in the new species and in several Australian Leptomyrmex species, and append these data to a 54‐taxon, 10‐gene data matrix previously generated for the subfamily Dolichoderinae. We conduct phylogenetic and divergence dating analyses, and re‐evaluate the fossil record of the group. We recover Leptomyrmex relictus sp.n. as a member of the Leptomyrmex clade with high support. It is sister to the Australasian species, and the genus Leptomyrmex is, in turn, sister to a pair of Neotropical genera, Forelius and Dorymyrmex. We infer a Neotropical origin for the genus and estimate a mid‐Eocene (46 Ma, 95% CI 56 to 36 Ma) origin for the crown genus and an Oligocene origin for the Australasian clade (29 Ma, 95% CI 40 to 19 Ma). We confirm placement of the Dominican amber species ?L. neotropicus Baroni Urbani in the genus but reject a close relationship with the Palaearctic fossil taxa ?Leptomyrmula Emery and ?Usomyrma Dlussky, Radchenko & Dubovikoff, considering them incertae sedis in the subfamily (Dolichoderinae). In contrast to the mesophilic preferences of the Australasian species of Leptomyrmex, the new Brazilian species inhabits cerrado (dry savannah). Our results support a Neotropical origin for spider ants with dispersal to Australia. Rafting on west‐bound currents and/or a historical diversity imbalance between Australia and South America are proposed as alternate hypotheses to explain a pattern of biased E–W mid‐Tertiary dispersal for ants with austral distributions. This pattern is suggested by our results in conjunction with observations of other ant clades. Overall, our findings highlight the value of integrated taxonomy, critical interpretation of morphology, and a comparative phylogenetic framework when conducting palaeontological and biogeographical studies of insect species. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:6E9E6617‐6E53‐40B8‐82C7‐67F89A83C553 .     

Evolution of Philopotinae,with a revision and phylogeny of the New World spider fly genus Philopota Wiedemann (Diptera,Acroceridae)

Philopotinae are hunchbacked spider flies represented by 63 fossil and extant species in 15 genera worldwide. Philopota Wiedemann, 1830, is the most species‐rich genus within the subfamily. Here, the evolution of Philopotinae is discussed, and a revision and phylogeny of Philopota based on adult morphology are presented. Nine of the 12 extant Philopotinae genera were included in our analysis, and 22 species were recognized in Philopota, of which 13 are described as new. Seven new synonymies are proposed. The phylogenetic analysis included 33 terminal taxa (22 ingroup and 11 outgroup species) and used 53 morphological characters, resulting in a single most parsimonious tree under equal weights. The monophyly of Philopota is recovered, and the Palaearctic genus Oligoneura is hypothesized as its sister‐group.     

First fossil record of an East Asian endemic genus Sladenia (Sladeniaceae) from its modern range: Implications for floristic evolution and conservation biology

Fossil records of endemic plants play an important role in recognizing the floristic history of East Asia and thereby facilitate the conservation of plant diversity in the region. However, the fossil record of many extant East Asian endemic genera remains poorly documented thus far. Here, we report an infructescence fossil of an East Asian endemic genus, Sladenia (Sladeniaceae), from the early Miocene of southeastern Yunnan, China. The fossil is characterized by: (i) dichasial cymes; and (ii) flask‐shaped ovary with dense subparallel ribs on the surface extending from the base to the distal end of the united style. It represents the first fossil record of Sladenia in Asia, showing that the genus was established in the region at least by the early Miocene. Given that a much older fossil record of Sladeniaceae has been reported from Africa and the sister group of Sladenia is distributed only in Africa, Sladenia is not likely of East Asian origin. The present endemic status of Sladenia was possibly achieved by regional extirpation in Africa and taking refuge in East Asia. This case thus supports the “Museum” rather than “Cradle” hypothesis for the genesis of high plant species in the flora of East Asia. A comparison of the present fossil with extant Sladenia infructescence shows morphological stasis from the early Miocene to present. Such evolutionary tardiness might have resulted in the reduced fitness of the genus, which further caused its current endangered situation.