Climate's role in the distribution of the Cretaceous terrestrial Crocodyliformes throughout Gondwana

A high diversity of terrestrial crocodyliform species has been found in the continental Cretaceous deposits of Gondwana. They are widespread in the sedimentary basins of Brazil, Uruguay, Argentina, Bolivia, Morocco, Cameroon, Niger, Malawi, Madagascar and Pakistan, from alluvial, fluvial and lacustrine deposits. A peculiar aspect of these terrestrial crocodyliforms is that only some of them are cosmopolitan. They comprise distinct groups as the basal Notosuchia, baurusuchids, sphagesaurids and Sebecian peirosaurids. There is a distribution pattern of the terrestrial Crocodyliformes faunas throughout the Cretaceous. The oldest are composed of the small and probably omnivorous Notosuchia and Araripesuchus, found in Early Cretaceous deposits. In the Late Cretaceous this fauna was enriched by the medium-sized to large-size baurusuchids, sphagesaurids, peirosaurids and larger Araripesuchus which present specializations as active terrestrial predators. The distribution analysis of the terrestrial Crocodyliformes from Early and Late Cretaceous palaeogeographic and palaeoclimatic maps indicates that temperature was the principal influence on their Gondwanan distribution. Although expressed seasonality, aridity is a limiting factor for the distribution of extant crocodilians. The Cretaceous basal Notosuchia, baurusuchids, sphagesaurids, Araripesuchus, Sebecian peirosaurids, are found in arid climatic belts during Early and Late Cretaceous. To live in a hot and arid climate they have presumably developed ecological strategies that allowed such habits. The aridity or seasonal warm and cyclic dry and wet climate periods plays a role, that have not yet been analyzed, that may explain the domain of bizarre Crocodyliformes in Gondwana during the Cretaceous.     

The Significance of the Found Fossil Species of Wodehouseia in China

Fossil pollen of Wodehouseia were first found by E. A. Stanley in 1961.There are several species of Wodehouseia found from the drill cores of the Lower member of Taizhou Formation in Rutung, North Jiangsu. It provides an evidence forjudging the Lower member of Taizhou Formation to be Maestrichtian age. In the assemblage of the Lower member of Taizhou Formation, there are not onlyAquiapollenites and Wodehouseia but also Morinoipollenites and Jianghanpollis. TheAquilapollenites and Wodehouseia are typical genera occurred in the humid, subtropicalclimatic phytogeoprovince in the “Aquilapolles District”, circumpacific and Siberiaduring Late Cretaceous age. The Morinoipollenites and Jianghanpollis are typical ge-nera occurred in the arid(or semiarid), subtropical climatic phytogeoprovince in widesouthern areas of our country at that time. This mixed assemblage provides a key tocompare the assemblage in wide southern areas of our country with those in other countries. The author discusses the geographical distributions and distributions of geologicalperiod of Wodehouseia in detail. The structure of Wodehouseia and the paleoecology of the parent plant of Wodehouseia pollen are explored. 7 species including 3 new spe-cies are described in this paper. The new species are Wodehouseia jiangsuensis sp. nov., W. circinata sp. nov., W. zoros sp. nov.     

On the Fossil Proteaceous Pollen in China

In this paper, some results of the study on the Proteaceous pollen collected recently in China are reported. On the basis of their different evolutionary features in geological history, their evolution is subdivided four stages in the Late Cretaceous—Early Tertiary, namely Cenomanian-Turonian, Senonian-Danian, Paleocene-Eocene and Oligocene. Each of the stages is typified by its own representative elements of Pro- teaceous and a certain domain of their predominance. According to the geographical distribution of Proteaceous pollen in China, four predominant districts are recognized, i.e. Northeast district of Late Cretaceous, Northwest-North-Inner Mongolia district of Early Tertiary, Centrial district of Late Cretaceous and Early Tertiary and South district of Early Tertiary. In addition, the problems concerning the stratigraphical, palaeoecological and pa- laeoelimatic significances of the Proteaceous pollen, and its origin are discussed in this paper too. Three new species are discribed here and they are: Proteacidites partitus sp. nov., P. tenellus sp. nov., P. bullatus sp. nov.     

Paleocene reefs on the Maiella Platform Margin, Italy: An example of the effects of the cretaceous/tertiary boundary events on reefs and carbonate platforms

Summary Reef facies, reef types and their biotic associations in the Maiella platform margin (central Italy) provide qualitative evidence for a significant reef decline across the Cretaceous/Tertiary (K/T) boundary, and indicate two phases of reef recovery during the Paleocene. Rudists dominated the reef community until the latest Cretaceous. A significant sea-level fall around the time of the K/T boundary is documented by a truncation surface associated with emersion. During sea-level highstands in the Danian to Early Thanetian and, more extensively, during the Late Thanetian, coral-algal patch-reefs grew along the platform margin and top. Already in the Danian to Early Thanetian, the reef communities were more diverse and the constructional types more evolved than previously known from this time. Differences between the Danian to Early Thanetian coral association, the Late Thanetian association, and Late Cretaceous coral faunas may have ecological or evolutionary causes. Repeated emergence produced a complex diagenetic history in the Danian to Lower Thanetian limestones. All Paleocene reefs were displaced by gravitative redeposition. Coral-algal reefs are less important in the Early to mid Eocene, when alveolinid foraminifera dominated on the Maiella shelf. Reefs on the Maiella platform diversified and attained large sizes in the Late Eocene to Early Oligocene, as known from other Mediterranean platforms. The external controls on the Late Cretaceous to Oligocene evolution and demise of reef communities that are most easily demonstrated with our data are sealevel fluctuations and climate change. We propose that the change in reef biota and reef types across the K/T boundary and during the Early Tertiary were important causes of the parallel changes in platform growth style.     

Vegetational patterns and distribution of relict taxa in humid temperate forests and wetlands of Georgia (Transcaucasia)

Small-scale vegetational patterns of Georgian lowland and montane forests and wetlands are documented by species per vegetation unit, function of species in particular vegetation units and vegetational sketches. The humid warm-temperate climate of western Georgia contributes to a different spatial pattern than known for most of Europe. One characteristic feature of western Georgian relict vegetation is the co-occurrence of 'Mediterranean', 'Sub-mediterranean', temperate, and Tertiary relict species, as well as of species normally confined to either zonal or azonal vegetation, in the same or closely related vegetation units. The extant distribution of Tertiary relict taxa strongly depends on how they are niching into changing environments. Niching strategies are assumed crucial for extinction or survival in times of rapid climatic/environmental changes. For Tertiary relict plant taxa in western Georgia we found four different niching strategies which also hold true of other northern hemispheric Tertiary relict plant taxa. Some amphibians and reptiles display similar distribution patterns and niching strategies as do relict plant taxa. A number of relict taxa in the warm humid regions of western Georgia occupy niches in swamp forests which might represent 'primeval' environments of species which are at present also elements of meso-Mediterranean vegetation and of thermophilous forest edges and hedges in Central Europe.     

Interplate dispersal paths for megathermal angiosperms

The dispersal of megathermal angiosperms between tectonic plates is reviewed on the basis of fossil evidence for the Cretaceous and Tertiary periods, since the radiation of the angiosperms, and the period of break-up of Gondwana. The combination of tectonic plate disassembly and redistribution, coupled with phases of global warming followed by pronounced cooling, has resulted in the formation of intermittent dispersal opportunities for frost-intolerant plants, and has been a major factor in determining the direction of angiosperm diversification. The Early Cretaceous radiation of angiosperms seems to show little relationship to the formation of Tethys. However, for the Late Cretaceous and Tertiary nine relevant dispersal routes can be differentiated that can be divided into two distinct categories: routes which formed following the break-up of Gondwana during the Late Cretaceous and Earlier Tertiary, when warm climates encouraged dispersal of megathermal elements globally, and routes which formed since the Middle Eocene, following phases of plate collision, as global climates were cooling down, inhibiting such dispersal. Most inter-plate dispersal of megathermal angiosperms took place in the Late Cretaceous and Early Tertiary at a time when global climates were markedly different from those of today, and the global area of megathermal vegetation several times greater than at present. Under such a scenario, it is likely than opportunities for speciation were much higher than for present-day megathermal plants.     

Structure and affinities of the petrified plants from the Cretaceous of Japan and Saghalien,V. Tree fern stems from Hokkaido,Paracyathocaulis ogurae gen. et comb. nov. andCyathocaulis yezopteroides sp. nov.

Paracyathocaulis ogurae gen. et comb. nov. andCyathocaulis yezopteroides sp. nov. are Late Cretaceous tree-fern stems with cyatheaceous affinity. They are comparable with the Recent scaly cyatheoids (Cyathea s.l.), as well as with the Cretaceous and Tertiary fossil stems that have similar anatomy. All of these tree ferns, including the Late Jurassic to Early CretaceousOguracaulis Tidwellet al., comprise a well-defined morphological complex within Cyatheaceae, and are considered to be monophyletic. The taxonomy of this complex is analyzed cladistically, and the cladogram is phylogenetically evaluated by reference to the chronological data of fossils.     

Early Cretaceous angiosperms of the Soviet Union based on leaf and fruit remains

Five newly collected Early Cretaceous angiosperms from the Kolyma River in Siberia are described. There are two new species of leaf, Cinnamomoides ievlevii and Celastrophyllum serrulatus and a new genus of fruit, Kenella harrisiana together with two leaves, Nelumbites aff. minimus Vachrameev and Celastrophyllum aff. hunteri Ward. Early Cretaceous angiosperm records for the Soviet Union are given in a table and their significance is discussed. It is argued that by the end of the Early Cretaceous whon the flora underwent a substantial change angiosperms were already distinguished by their high state of morphological development judging by leaf form and systematic composition. They were widely spread occurring in regions of both temperate and warm humid climates. They appear to have been plants of mountains, including trees and herbs, terrestrial and aquatic. With the advent of physical and climatic changes these angiosperms were ready for rapid expansion. Their domination of the vegetation of the Late Cretaceous was not as inexplicable as previously thought.     

The Tertiary Vegetation and Flora and Floristic Regions in China

The development or adaptation radiation of angiosperms has been closely related to the paleoclimate and paleoenvironment. Several environmental factors, influencing each other, have affected the evolution or development of angiosperms, which falls into different stages. The lines between these stages are not consistent with the traditional geological ages. Floristic regionalization might be better based on the stages of the development. Angiosperms have become important or dominant elements in the flora since the middle and late period of the Late Cretaceous. The development of angiosperms may be divided into four stages: (1) Initial stage: The angiosperms were not yet well established during the Early Cretaceous. The number of species and individuals was limited, the leaf small, mostly with entire margine, irregular venation and poor differentiation of venation; (2) Flourished stage: The angiosperms increased dramatically and became abundant. The leaf was larger with more regular venation and the differentiation of venation was remarkable. The percentage of angiosperms was from 40%-60% in the flora, and eventually arose, becoming predominant, with important families well presented. This was probably from the Late Cretaceous to the Early Tertiary; (3) Herbs-Flourished stage: Some of woody plants were extinct, while herbs greatly increased, because of the changes in climate, sea level and mountain uplift during the Neogene period; (4)Quaternary stage: The mountain glacier in China, influenced by global glacial-interglacial alternation due to the climatic fluctuation, advanced and regressed during the Quaternary. The climatic fluctuation apparently affected the distribution of plants. But the components of the flora were similar to or slightly different from the features of the present one. The aim of the present paper is to discuss the floristic regions at the second and third stages. The Late Cretaceous-Early Tertiary stage: A. The Late Cretaceous-Paleogene flora of northern China 1. Warm-temperate to subtropical deciduous broad-leaved and conifer forests in North and Northeastern China 2. Dry subtropical flora of Northwestern and Central China B. Southern China Late Cretaceous-Paleogene flora 3. Deciduous and evergreen forests with coniferous elements in the subtropical coastal region of east China 4. Evergreen forests of South China 5. Evergreen forests of lower mountains in the Tethys The Neogene stage-Herbs-flourished stage: 1. Temperate forests and grasslands to semidesert-desert floras of northwestern China; 2. Warm temperate deciduous forests of north and northeastern China; 3. Warm temperate to subtropical deciduous and evergreen forests of central and east China; 4. Evergreen forests of the subtropical and tropics mangrove vegetation of south China; 5. Subtropical evergreen and deciduous forests on the Yunnan and Xizang plateau.     

Hemiasterid echinoids (Echinodermata: Spatangoida) from the Cretaceous Tethys to the present-day Mediterranean

Hemiasterid echinoids form one of the oldest and most varied spatangoid groups. Current data on stratigraphical and geographical distribution pattern show that their biogeographical history is closely related to Mediterranean-Tethyan structural and climatic changes. The group first appeared in the Aptian in the central Tethys (southwestern Europe) and developed and diversified remarkably in the Late Cretaceous through the subgenera Hemiaster (Hemiaster), H. (Bolbaster) and H. (Leymeriaster). Marked morphological changes arose in the early Paleogene when new Tertiary forms H. (Trachyaster), Opissaster and Ditremaster, originating in the Caribbean and Madagascan regions, respectively, replaced H. (Bolbaster) which partly survived and H. (Hemiaster) and H. (Leymeriaster) which disappeared entirely. These new forms achieved their greatest diversity during the Eocenee and thrived in the Mediterranean basin until it was sealed off from the Indo-Pacific in the Middle Miocene. Then Late Neogene climatic and structural events reduced hemiasterids to only a few Mediterranean species. The survivors of modern groups persisted in refuge areas of the western Mediterranean until middle Messinian times before being reduced to only one species, the last fossil one, during the Pliocene. Then, the Cenozoic groups H. (Trachyaster) and Opissaster definitively disappeared. Outside the Mediterranean basin primitive H. (Bolbaster) and modern Ditremaster, which colonized deep-water Atlantic environments from the Late Cretaceous and Middle Miocene respectively, survived the Messinian and Plio-Pleistocene climatic crises. They flourished in the northern Atlantic where their present-day descendants Ditremaster (Sarsiaster) griegii and H. (Bolbaster) expergitus still survive. H. (Bolbaster) expergitus spread worldwide and colonized the western Mediterranean in the Plio-Pleistocene via the Straits of Gibraltar.     

Seed Size, Fruit Size, and Dispersal Systems in Angiosperms from the Early Cretaceous to the Late Tertiary

Fossil data from 25 angiosperm floras from the Early Cretaceous ( approximately 124 million years ago) to the Pliocene ( approximately 2 million years ago) were compiled to estimate sizes of seeds and fruits and the relative proportion of two different seed-dispersal systems by animals and by wind. The results suggest that, first, seed and fruit sizes were generally small during most of the Cretaceous, in agreement with previous suggestions, but the trend of increasing sizes started before the Cretaceous-Tertiary boundary; second, there was a decrease in both seed and fruit sizes during late Eocene and Oligocene, reaching a level that has continued to the Late Tertiary; third, the fraction of animal dispersal was, in contrast to previous suggestions, rather high also during the Cretaceous but increased drastically in the Early Tertiary and declined congruently with the declining seed and fruit sizes from the late Eocene; and fourth, the fraction of wind dispersal showed a bimodal pattern, being high in the Late Cretaceous and in the Oligocene-Miocene but with a drop in between. We find that the observed trends are only weakly related to the availability of animal fruit dispersers. Instead, the trends are congruent with a climate-driven change in environmental conditions for recruitment, where larger seeds are favored by closed forest vegetation. The prevalence of semiopen, dry, and probably herbivore-disturbed vegetation during the Cretaceous, the development of closed multistratal forests in the Eocene, and the later development of a more open vegetation and grasslands starting in the Oligocene-Miocene, are reflected in the distribution of angiosperm seed and fruit sizes and in the dispersal systems.     

<Emphasis Type="Italic">Protungulatum</Emphasis>, Confirmed Cretaceous Occurrence of an Otherwise Paleocene Eutherian (Placental?) Mammal

Neither pre-Cenozoic crown eutherian mammals (placentals) nor archaic ungulates (“condylarths”) are known with certainty based on the fossil record. Herein we report a new species of the Paleocene archaic ungulate (“condylarth”) Protungulatum from undisputed Late Cretaceous aged rocks in Montana USA based on an isolated last upper premolar, indicating rare representatives of these common early Tertiary mammals appeared in North America a minimum of 300 k  years before the extinction of non-avian dinosaurs. The other 1200 mammal specimens from the locality are characteristic Late Cretaceous taxa. This discovery overturns the current hypothesis that archaic ungulates did not appear in North America until after the Cretaceous/Tertiary (K/T) boundary and also suggests that other reports of North American Late Cretaceous archaic ungulates may be correct. Recent studies, including ours, cannot determine whether Protungulatum does or does not belong to the crown clade Placentalia.     

壳斗科的地质历史及其系统学和植物地理学意义

在收集整理现有壳斗科化石资料的基础上,讨论了壳斗科及其各属的起源时间、地史分布和地史 演替过程以及这些化石资料在系统学和植物地理学上的意义。白垩纪尚无壳斗科可靠的大化石记录, 微化石需要进一步研究才能确定亲缘关系以及古新世壳斗科已经分化出两个类群。从以上这些事实推 论壳斗科起源于白垩纪晚期,而壳斗科现代各属出现的时间应不晚于古新世。最早发现的壳斗科化石和现代栗亚科和水青冈亚科在形态结构上非常相似,这一事实表明,壳斗科分为两个亚科的观点更接近客观事实。在水青冈亚科中,三棱栎类的化石最早出现;在栎属中,青冈亚属更接近祖先类群;在地史中全缘栎类较具齿栎类出现早,粗齿的落叶栎类出现最晚。三棱栎属、栲属和石栎属的化石在老第三纪出现于北美和欧洲的事实说明,北美、欧洲和东亚在老第三纪时有一个相通的壳斗科植物区系。南美的三棱栎是通过北美进入南美的。中国横断山、欧洲地中海沿岸和北美西北部有一类形态特征相似、亲缘关系相近的硬叶栎类,它们之间有相同的地质演替历史,它们现代分布边界可能就是古地中海的边界。美洲的栎类有两个来源,常绿硬叶栎类是通过古地中海沿岸而经北美-欧洲陆桥到达的,落叶栎类则是在中新世以后通过白令海峡到达的。     

Fossils of the Fagaceae and their implications in systematics and biogeography

The fossil history of the Fagaceae from China and its systematic and biogeographic implications are discussed based on revisionary studies of the fossil records. No creditable macrofossil record of the Fagaceae exists in the Cretaceous deposits and all the Cretaceous microfossil reports remain equivocal and require further study. The Paleocene fossils show the appearance and diversification of the two groups corresponding to the subfamilies Fagoideae and Castaneoideae sensu Nixon. By the Eocene, all modern genera had been present. The oldest fagaceous fossils represent subfamily Fagoideae with affinities to the extant genus Trigonobalanus. The leaf fossil genus Berryophyllum, with affinities to Quercus subg. Cyclobalanopsis, has been documented by the early Eocene and might have occurred earlier than other fossils assignable to Quercus. The appearance of evergreen sclerophyllous Ouercus with entire leaves might have occurred earlier than those with toothed leaves. Deciduous, urticoid-leaved oak fossils (Quercus subg. Quercus sect. Quercus) had not appeared until the Miocene. Fossil equivalents of Trigonobalanus, Castanopsis and Lithocarpus had occurred in Europe and North America by the early Tertiary, suggesting that continuous distributions were achieved via the northern hemisphere land bridges. Three groups of evergreen sclerophyllous oaks of apparent close phylogenetic relationships occurred in the Hengduan mountains, the Mediterranean area and northwestern North America. Their fossil forms have become dominant elements of those vegetation zones since the Miocene. A shared fossil history indicates a possible biogeographic boundary formed by the ancient Mediterranean. The evidence suggests that the oaks might arrive in North America during two distinct geologic periods: evergreen sclerophyllous entire-leaved oaks appeared by the Early Tertiary, whereas thedeciduous oaks with urticoid leaves appeared in the Late Tertiary.     

Phylogeny and Phytogeography of the Betulaceae (to be continued)

In the present thesis, a detailed review of the literature was given on the Betulaceae. Some new data was provided based on the author’s studies in order to analyze various characters, such as inflorescence, flower, pollen and leaf epidermis, among the genera treated. The polarity of the characters was determined according to the criteria of outgroup comparison and compatibility analysis. A cladistic analysis was made using the methods of Maximal Same Steps and Minimal Parallel Evolution developed by Xu (1993). The generic distributions in both modern and fossil times were outlined, and their distribution center, place and time of origin and the way of dispersal were discussed. An attempt was made to discuss early differentiation, geographical radiation and morphological evolution of the family on the background of paleogeography and paleoclimatology, tracing back as far as the Late Cretaceous and Early Tertiary. Finally, supraspecific taxa, from section to tribe, were systematically arranged.     

The historical biogeography of sturgeons (Osteichthyes: Acipenseridae): a synthesis of phylogenetics, palaeontology and palaeogeography

Abstract. The historical biogeography of sturgeons is explored using information from palaeogeography, palaeontology and phylogenetic interrelationships. The integration of information from these diverse sources indicates that sturgeons reached a wide Laurasian distribution in the Cretaceous and Tertiary by freshwater and coastal dispersal routes across land connections and along newly forming continental margins. The fossil record also suggests a considerable degree of morphological stasis and also supports an estuarine habit, and perhaps diadromy, as an old and conserved life history trait. While a ‘centre of origin’ for sturgeons remains elusive, phylogenetic relationships indicate that diversification appears to have been associated with fragmentation of biota, and of landmasses and basins, by late Tertiary geological and climatic phenomena, such as orogeny and unequal glaciation over North America, the desiccation of central Asia and alteration of its drainages, and the formation of discrete Ponto-Caspian basins by the fragmentation of the Paratethys. Amphi-oceanic distributions of certain species (Acipenser medirostris Ayres) and sister taxa (e.g. A. oxyrhynchus Mitchill and A. sturio L.) are explained by coastal dispersal and subsequent vicariance by geological (sea-floor spreading and development of new continental margins) and climatic (Pliocene cooling) changes during the Tertiary. An hypothesis is developed for the relationships of the North American sturgeons and their potential relationships with the Siberian sturgeon A. baeri. Late Tertiary climatic and geological phenomena are hypothesized as mediators of vicariance and subsequent diversification of these acipenserids. It appears that although acipenserids are a geologically old group, the historical biogeography of surviving lineages is best explained by more recent geological and climatic changes.     

The Paleocene-Eocene Palynoflora from the Tantou Basin in West Henan

On the basis of palynological data the Paleoeene-Eocene palynoflora from the Tantou Basin in west Itenan are divided into three sporo-pollen assemblages. They are given ill ascending order as follows: 1. The first assemblage (Dazhang Formation) characterized mainly by the predominance of palynoforms of Ulmaeeae anal various Proteaeeous pollen, being assigned to Late Paleoeene and indicating a dry-hot climate. 2. The second assemblage (Lower part of the Tantou Formation) marked by the presence of various Triporopollenites and Tricolporopollenites, belonging to the Early Eocene. A moist-hot climate is reflected. 3. The third assemblage (the Upper part of the Tantou Formation) represented by aboundant pollen grains of saeeated elements (consisting of Pinaeeae and Podocarpaeeae) and the occurrence of Aquilapollenites pollen which plays an important part in this assemblage. It is interpreted as belonging to the middle to early Eocene. The climate was moist and warm. As indicated by the sporo-polleu assemblage the present Paleoeene to Eocene flora in the whole phytoprovienee may belong to a flora transitional from western to eastern China. In this paper the distribution of Aquilapollenites pollen and its stratigraphicai significance are also discussed and it is considered that this kind of pollen distributed in the Paleoeene of China is eertainly not redeposited. At the same time, sonm palynoforms of certain significance for delimitation of Paleoeene and Eocene are provided.     

Quantitative Analysis of the Timing of the Origin and Diversification of Extant Placental Orders

Fossil evidence is consistent with origination and diversification of extant placental orders in the early Tertiary (Explosive Model), and with the possibility of some orders having stem taxa extending into the Cretaceous (Long Fuse Model). Fossil evidence that 15 of 18 extant placental orders appeared and began diversification in the first 16 m.y. of the Cenozoic is, however, at odds with molecular studies arguing some orders diversified up to 40 m.y. earlier in the Early Cretaceous (Short Fuse Model). The quality of the fossil record was assessed by tabulating localities of all mammals in the last 105 m.y. Global locality data (except Africa) for 105 m.y. of eutherian evolution indicate discernible biogeographic patterns by the last 15 m.y. of the Cretaceous. Eutherian genera increase from 11 in latest Cretaceous to 139 in earliest Tertiary, although both are represented by about 50 localities. Yet even in the Late Cretaceous of North America and Asia where eutherians are abundant, none of the 18 extant orders are definitely known. A series of Monte Carlo simulations test whether the rapid appearance of most mammalian orders is statistically significant, and if so, whether it is a radiation event or an artifact of a limited fossil record. Monte Carlo tests affirm that the clustering of appearances in the early Cenozoic is statistically significant. Quantitative analysis of the locality data suggests that the number of genera described is a function of the number of localities sampled. In contrast, the number of orders is not a simple function of localities and thus does not appear to be limited by localities. A second set of Monte Carlo simulations confirms that the increase in orders cannot be explained by the limited number of localities sampled. Even for best-fit simulations, the observed pattern of ordinal appearances is steeper than expected under a variety of null models. These quantitative analyses of the fossil record demonstrate that the rapid ordinal appearances cannot be ascribed to limited Late Cretaceous sample sizes; thus, early Tertiary ordinal diversification is real. Although the fossil record is incomplete, it appears adequate to reject the hypothesis that orders of placentals began to diversify before the K/T boundary.     

Evolution of Lycopodiaceae (Lycopsida): estimating divergence times from rbcL gene sequences by use of nonparametric rate smoothing

By use of nonparametric rate smoothing and nucleotide sequences of the rbcL gene, divergence times in Lycopodiaceae are estimated. The results show that much extant species diversity in Lycopodiaceae stems from relatively recent cladogenic events. These results corroborate previous ideas based on paleobotanical and biogeographical data. Previous molecular phylogenetic analyses recognized a split into neotropical and paleotropical clades in Huperzia, which contains 85-90% of all living species. Connecting this biogeographical pattern with continent movements, the diversification of this epiphytic group was suggested to coincide with that of angiosperms in the mid to Late Cretaceous. Results presented here are consistent with this idea, and the diversification of the two clades is resolved as Late Cretacous (78 and 95 Myr). In the related genera Lycopodium and Lycopodiella, the patterns are somewhat different. Here species diversity is scattered among different subgeneric groups. Most of the high-diversity subgeneric groups seem to have diversified very recently (Late Tertiary), whereas the cladogenic events leading to these groups are much older (Early to Late Cretaceous). Our analysis shows that, although much living species diversity stems from relatively recent cladogenesis, the origins of the family (Early Carboniferous) and generic crown groups (Early Permian to Early Jurassic) are much more ancient events.     

Osteopygis (Testudines: Cheloniidae) from the Lower Tertiary of the Ouled Abdoun phosphate basin, Morocco

Osteopygis emarginatus Cope 1868 is described from the Lower Tertiary of the Ouled Abdoun phosphate basin, Morocco, on the basis of skulls and lower jaws. Osteopygis is a cosmopolitan turtle that had a wide geographical distribution during the Late Cretaceous and Paleocene. Osteopygis emarginatus is a very conservative species which crossed the Cretaceous/Tertiary boundary without major changes.