中国中生代的马通蕨科植物:化石记录、多样性与时空分布特征*

现生真蕨目马通蕨科(Matoniaceae)植物仅存Matonia和Phanerosorus两属, 集中分布于马来西亚、印度尼西亚等热带地区。马通蕨科植物在中生代时期全球广布, 且主要分布于热带、亚热带地区, 有近9个属, 被作为热带、亚热带气候的标志性植物化石之一。本文梳理总结了中国中生代的马通蕨科化石记录并分析其多样性特征, 共计有2属16种, 包括异脉蕨属(Phlebopteris) 15种和准马通蕨属(Matonidium) 1种。对其化石记录和地质地理分布分析表明, 晚三叠世时期, 异脉蕨属植物广泛分布于热带—亚热带湿热气候区, 包括华南一带的四川、湖北、 云南、西藏、福建等地; 早侏罗世时期, 其分布逐渐向北方扩展, 在南、北方植物区系界线附近均有发现; 中侏罗世局限于湖北、青海等地; 早白垩世时仅在黑龙江和西藏少量发现。准马通蕨属仅在黑龙江地区的早白垩世地层中发现。整体上, 马通蕨科在中国中生代的分布范围变迁与气候带范围变化相吻合。     

Seed ferns survived the end-Cretaceous mass extinction in Tasmania

Seed ferns, dominant elements of the vegetation in many parts of the world from the Triassic to Cretaceous, were considered to have disappeared at the end of the Cretaceous together with several other groups that had occupied key positions in terrestrial and marine ecosystems such as dinosaurs, plesiosaurs, and ammonoids. Seed-fern demise is generally correlated with competition from diversifying flowering plants through the Cretaceous and the global environmental crisis related to the Chicxulub impact event in the paleotropics at the end of the period. New fossils from Tasmania show that one seed-fern lineage survived into the Cenozoic by at least 13 million years. These fossils are described here as a new species, Komlopteris cenozoicus. Komlopteris is a genus of seed ferns attributed to Corystospermaceae and until now was not known from sediments younger than the Early Cretaceous. Discovery of this "Lazarus taxon," together with the presence of a range of other relictual fossil and extant organisms in Tasmania, other southern Gondwanan provinces, and some regions of northern North America and Asia, underscores high-latitude regions as biodiversity refugia during global environmental crises and highlights their importance as sources of postextinction radiations.     

Early Cretaceous angiosperm invasion of Western Europe and major environmental changes

BACKGROUND AND AIMS: At the beginning of the Late Cretaceous, angiosperms already inhabited all the environments and overtopped previously gymnosperm-dominated floras, especially in disturbed freshwater-related environments. The aim of this paper is to define what fossil plant ecology occurred during the early Cretaceous in order to follow the early spread of angiosperm taxa. METHODS: Floristic lists and localities from the Barremian to the Albian of Europe are analysed with the Wagner's Parsimony Method. KEY RESULTS: The Wagner's Parsimony Method indicates that (a) during the Barremian, matoniaceous ferns formed a savannah-like vegetation, while angiosperms composed freshwater aquatic vegetation; (b) during the Late Aptian humid phase, conifers increased, while matoniaceous ferns decreased, reflecting the closure of the vegetation; and (c) from the Albian, warmer and drier conditions induced the recovery of the matoniaceous ferns, while core angiosperms first developed in floodplains. CONCLUSIONS: During the late Early Cretaceous (Barremian-Albian), angiosperms showed a stepwise widening of their ecological range, being recorded first during the Barremian as aquatic plant mega-remains and at the Cenomanian onwards occurred in all the environments.     

The lower cretaceous ferns in the genus Anemia (Schizaeceae), Potomac group of Virginia, and relationships within the genus

Based on characters of the fertile fronds, two species of fossil ferns from the Lower Cretaceous Potomac Group of the U.S.A., originally described as species of Osmunda by Fontaine in 1889, are transferred to the genus Anemia. The two species display primitive characters for that genus. Analysis of these and other characters of Anemia and related species, using parsimony and compatibility programs, suggests possible phylogenetic arrangements and facilitates delimination of three taxonomic units within Anemia that were already distinct during Cretaceous time. The subgenera Coptophyllum and Anemia are shown to be paraphyletic in the suggested phylogenies and the schemes agree with the known stratigraphy of the fossil ferns.     

Recurrent genome duplication events likely contributed to both the ancient and recent rise of ferns

Ferns, the second largest group of vascular plants, originated ～400 mil ion years ago(Mya). They became dominant in the ancient Earth landscape before the angiosperms and are stil important in current ecosystems.Many ferns have exceptional y high chromosome numbers,possibly resulting from whole-genome duplications(WGDs).However, WGDs have not been investigated molecularly across fern diversity. Here we detected and dated fern WGDs using a phylogenomic approach and by calculating synonymous substitution rates(Ks). We also investigated a possible correlation between proposed WGDs and shifts in species diversification rates. We identified 19 WGDs: three ancient events along the fern phylogenetic backbone that are shared by 66%–97% of extant ferns, with additional lineage-specific WGDs for eight orders, providing strongevidence for recurring genome duplications across fern evolutionary history. We also observed similar Ks peak values for more than half of these WGDs, with multiple WGDs occurring close to the Cretaceous(～145–66 Mya). Despite the repeated WGD events, the biodiversity of ferns declined during the Cretaceous, implying that other factors probably contributed to the floristic turnover from ferns to angiosperms. This study provides molecular evidence for recurring WGDs in ferns and offers important clues to the genomic evolutionary history of ferns.     

黑龙江省东部鸡西盆地早白垩世穆棱组植物化石新材料

系统描述了最近采集的穆棱组植物化石22属23种,其中的两种被子植物叶化石均为具原始被子植物叶特征的小型叶,在该植物群中尚属首次发现。运用扫描电镜(SEM)等方法对其中的Tyrmia sp．,Nilssoniopteris sp．,Ginkgoites cf．．sibirica,．Sphenobaiern sp．,Elatocladus manchurica等化石的叶角质层进行了研究。据已有的资料统计,穆棱组迄今已知有化石植物31属38种。该植物群以真蕨类占据绝对优势,裸子植物也相当丰富,其中的大多数种类主要见于早白垩世,穆棱组的地质时代应属早白垩世巴列姆期或巴列姆-阿普梯早期。植物组合中以丰富的热带及亚热带型植物为主要特点,未发现针叶类型的植物,说明早白垩世穆棱组沉积时,该地区的气候温暖湿润;另一方面,落叶的银杏类和松柏类,以及裸子植物木化石中具有清楚的生长轮,又说明该地区的古气候存在着季节变化。该植物群属于早白垩世西伯利亚植物区或加拿大-西伯利亚植物区。     

Distinguishing angiophytes from the earliest angiosperms: A Lower Cretaceous (Valanginian-Hauterivian) fruit-like reproductive structure

A remarkably diverse Lower Cretaceous (Valanginian-Hauterivian) flora at Apple Bay, Vancouver Island, preserves seed plants at an important time of floristic evolutionary transition, about the same time as the earliest flowering plant megafossils. The fossils are permineralized in carbonate concretions and include tetrahedral seeds within cupule- or carpel-like structures. These enclosing structures, composed of elongate sclerenchyma cells with spiral thickenings that grade externally to a few layers of parenchyma, are vascularized by one collateral vascular bundle and lack trichomes. They apparently broke open to release the tightly enclosed seeds by valves. Seeds are similar to those of the Triassic seed fern Petriellaea, but are about 100 million years younger and differ in size, vascularization, integumentary anatomy, seed attachment, and number of seeds/cupule. These new seeds are described as Doylea tetrahedrasperma gen. et sp. nov., tentatively assigned to Corystospermales. Inverted cupules are reminiscent of an outer angiosperm integument rather than a carpel. Like fruits, cupules opened to release seeds at maturity, thereby foretelling several aspects of angiospermy. They show that nearly total ovule enclosure, a level of organization approaching angiospermy, was achieved by advanced seed ferns during the Mesozoic.     

Review of the monotreme fossil record and comparison of palaeontological and molecular data

Monotremes have traditionally been considered a remnant group of mammals descended from archaic Mesozoic stock, surviving to the present day on the relatively isolated Australian continent. Challenges to this orthodoxy have been spurred by discoveries of 'advanced' Cretaceous monotremes (Steropodon galmani, Archer, M., et al., 1985. First Mesozoic mammal from Australia-an Early Cretaceous monotreme, Nature. 318, 363-366) as well as by results from molecular data linking monotremes to therian mammals (specifically to marsupials in some studies). This paper reviews the monotreme fossil record and briefly discusses significant new information from additional Cretaceous Australian material. Mesozoic monotremes (including S. galmani) were a diverse group as evidenced by new material from the Early Cretaceous of New South Wales and Victoria currently under study. Although most of these new finds are edentulous jaws (limiting dental comparisons and determination of dietary niches), a range of sizes and forms has been determined. Some of these Cretaceous jaws exhibit archaic features-in particular evidence for the presence of a splenial bone in S. galmani-not seen in therian mammals or in post-Mesozoic (Tertiary and Quaternary) monotreme taxa. Tertiary monotremes were either archaic ornithorhynchids (toothed platypuses in the genera Monotrematum and Obdurodon) or tachyglossids (large echidnas in the genera Megalibgwilia and Zaglossus). Quaternary ornithorhynchid material is referable to the sole living platypus species Ornithorhynchus anatinus. Quaternary echidnas, however, were moderately diverse and several forms are known (Megalibgwilia species; 'Zaglossus' hacketti; Zaglossus species and Tachyglossus aculeatus).     

Early Cretaceous rhynchonellids of Dagestan: System, morphology, stratigraphic and paleobiogeographic significance

A comprehensive report on Early Cretaceous Dagestan rhynchonellids is provided for the first time. It includes the rhynchonellid system based on detail morphofunctional studies, recognition of the boundary in the evolutionary development of Late Mesozoic rhynchonelloids at the Barremian-Aptian boundary, analysis of changes in shell matter structure during successive time periods of the Early Cretaceous, elucidation of stratigraphic importance of species assemblages allocated stratigraphically by layers to six sections of the Lower Cretaceous deposits described by the author, and revelation of possibilities of using this group in paleobiogeographic reconstructions. The study describes 37 species, 7 genera, and 2 families; 16 species and 2 genera are new. This paper is useful for a wide range of paleontologists, experts on the Mesozoic fauna, biostratigraphers, and geologists dealing with Mesozoic deposits. The study provides 8 plates, 58 figures, 71 references.     

Crocodyliform biogeography during the Cretaceous: evidence of Gondwanan vicariance from biogeographical analysis

Explanations of the distributions of terrestrial vertebrates during the Mesozoic are currently vigorously contested and debated in palaeobiogeography. Recent studies focusing on dinosaurs yield conflicting hypotheses. Dispersal, coupled with regional extinction or vicariance driven by continental break-up, have been cited as the main causal factors behind dinosaur distributions in the Mesozoic. To expand the scope of the debate and test for vicariance within another terrestrial group, I herein apply a cladistic biogeographical method to a large sample of Cretaceous crocodyliform taxa. A time-slicing methodology is employed and a refinement made to account for the divergence times of the analysed clades. The results provide statistically significant evidence that Gondwana fragmentation affected crocodyliform diversification during the Mid-Late Cretaceous. Detection of a vicariant pattern within crocodyliforms is important as it helps corroborate vicariance hypotheses in other fossil and extant groups as well as furthers the move towards more taxonomically diverse approaches to palaeobiogeographical research.     

Fossil history of Mesozoic weevils (Coleoptera: Curculionoidea)

Abstract The first synopsis of Mesozoic weevils (Curculionoidea: Coleoptera) is presented. Changes of family, genera and species abundance during the Mesozoic revealed three distributional patterns. The Jurassic (Karatau) fauna was dominated by the Nemonychidae. During the Early Cretaceous (beginning at the Jurassic/Cretaceous border), the Ithyceridae was the prevalent group with a significant role played by the Nemonychidae. In the Late Cretaceous (Cenomanian and Turonian), the major groups were the Curculionidae and Brentidae. Obviously, the change of weevil fauna during this period was due to the expansion of the angiosperms, which provided multiple niches in their vegetative and reproductive organs for weevil development.     

A revision of the genusCyathorachis in Japan

Cyathorachis, a genus of Cretaceous tree fern rachises is revised. In addition to the type specimens of the sole species,C. fujiiana Ogura, two specimens collected near the type locality were examined histologically. The histological data being used for the taxonomical comparison were obtained from anatomical studies on the rachises of extant tree ferns. As a result of the study three species are recognized, two of which are new. As a result of this study, it appears that species differentiation of the tree ferns in the late Cretaceous age of Hokkaido was rather high, as it is now in tropical and subtropical regions. Contributions from the Laboratory of Phylogenetic Botany, Chiba University, No. 74.     

On the genus Tomaxellia (Coniferae) from the Lower Cretaceous of Patagonia (Argentina) and its male and female cones

This paper reports the organic attachment of male and female cones to twigs previously described as Tomaxellia biforme Archangelsky. The male cones produce Classopollis pollen, while the scales of the female structures are comparable to the Rhaeto-Liassic northern genus Cheirolepidium. A new interpretation of the possible female cone of Cheirolepidium is presented, based on the new evidence now available with the knowledge of Tomaxellia cones. Other female cones found in Mesozoic formations may be comparable to some extent with Tomaxellia , such as Indostrobus (Cretaceous of India) or Pararaucaria (Jurassic of Argentina), and they may be grouped in the family Cheirolepidiaceae. Comparisons with older conifer genera with known female cones are also included (Voltziaceae). Some morphological changes of the female cones, which probably took place during the Mesozoic (in these particular groups) are also inferred.     

Contrasting Predictors of Fern versus Angiosperm Decomposition in a Common Garden

We evaluated differences in the rates and correlates of decomposition among 32 fern and angiosperm litter types collected in Hawai'i. Leptosporangiate ferns were separated into groups based on phylogeny: 'polypod' ferns, a monophyletic clade of ferns that diversified in the Cretaceous, and all other ('non-polypod') ferns that diversified earlier. We measured initial litter chemistry (nutrients and carbon chemistry), and mass loss and nitrogen (N), phosphorus (P), and calcium (Ca) of litter tissue during a 1-yr incubation in a common garden. Nutrient concentrations and carbon (C) chemistry differed significantly among litter types, and litter turnover ( k -values) ranged from 0.29 to 8.31. Decomposition rates were more closely correlated with nutrient concentration than is typically observed. Lignin:N was the best predictor of decomposition across all litter types combined; however, among plant groups different predictors of decomposition were important. Nitrogen and P concentrations best predicted fern decomposition, whereas C chemistry, particularly lignin concentration, was more important for angiosperm (monocot and dicot) decomposition. Among native plants, non-polypod ferns decomposed significantly more slowly than both polypod ferns and angiosperms. Contrary to our hypothesis, fern litter did not decompose more slowly than angiosperm litter overall. Nutrient dynamics in litter were affected by initial litter concentration more than phylogeny; low-nutrient litter immobilized more nutrients than high-nutrient litter. Systematic differences in rates of decomposition, and the importance of nutrients in predicting fern decomposition, imply that changes in species composition within ferns and between ferns and angiosperms could influence the functioning of ecosystems where ferns are important forest components.     

A new global palaeobiogeographical model for the late Mesozoic and early Tertiary

Late Mesozoic palaeobiogeography has been characterized by a distinction between the northern territories of Laurasia and the southern landmasses of Gondwana. The repeated discovery of Gondwanan lineages in Laurasia has led to the proposal of alternative scenarios to explain these anomalous occurrences. A new biogeographical model for late Mesozoic terrestrial ecosystems is here proposed in which Europe and "Gondwanan" territories possessed a common Eurogondwanan fauna during the earliest Cretaceous. Subsequently, following the Hauterivian, the European territories severed from Africa and then connected to Asiamerica resulting in a faunal interchange. This model explains the presence of Gondwanan taxa in Laurasia and the absence of Laurasian forms in the southern territories during the Cretaceous. In order to test this new palaeobiogeographical model, tree reconciliation analyses (TRAs) were performed based on biogeographical signals provided by a supertree of late Mesozoic archosaurs. The TRAs found significant evidence for the presence of an earliest Cretaceous Eurogondwanan fauna followed by a relatively short-term Gondwana-Laurasia dichotomy. The analysis recovered evidence for a biogeographical reconnection of the European territories with Africa and South America-Antarctica during the Campanian to Maastrichtian time-slice. This biogeographical scenario appears to continue through the early Tertiary and sheds light on the trans-Atlantic disjunct distributions of several extant plant and animal groups.     

Intercontinental disjunctions between eastern Asia and western North America in vascular plants highlight the biogeographic importance of the Bering land bridge from late Cretaceous to Neogene

This review shows a close biogeographic connection between eastern Asia and western North America from the late Cretaceous to the late Neogene in major lineages of vascular plants (flowering plants, gymnosperms, ferns and lycophytes). Of the eastern Asian–North American disjuncts, conifers exhibit a high proportion of disjuncts between eastern Asia and western North America. Several lineages of ferns also show a recent disjunct pattern in the two areas. In flowering plants, the pattern is commonly shown in temperate elements between northeastern Asia and northwestern North America, as well as elements of the relict boreotropical and Neogene mesophytic and coniferous floras. The many cases of intercontinental biogeographic disjunctions between eastern Asia and western North America in plants supported by recent phylogenetic analyses highlight the importance of the Bering land bridge and/or the plant migrations across the Beringian region from the late Cretaceous to the late Neogene, especially during the Miocene. The Beringian region has permitted the filtering and migration of certain plant taxa since the Pliocene after the opening of the Bering Strait, as many conspecific taxa or closely related species occur on both sides of Beringia.     

The Gondwanan and South American Episodes: Two Major and Unrelated Moments in the History of the South American Mammals

The first steps in the history of South American mammals took place ca. 130 Ma., when the South American plate, still connected to the Antarctic Peninsula, began to drift away from the African-Indian plate. Most of the Mesozoic history of South American mammals is still unknown, and we only have a few enigmatic taxa (i.e., a Jurassic Australosphenida and an Early Cretaceous Prototribosphenida) that pose more evolutionary and biogeographic questions than answers. The best-known Mesozoic, South American land-mammal fossils are from Late Cretaceous Patagonian beds. These fossils represent the last survivors of non- and pre-tribosphenic Pangaean lineages, all of them with varying endemic features: some with few advanced features (e.g., ?Eutriconodonta and “Symmetrodonta”), some very diversified as endemic groups (e.g., ?Docodonta Reigitheriidae), and others representing vicariant types of well known Laurasian Mesozoic lineages (e.g., Gondwanatheria as vicariant of Multituberculata). These endemic mammals lived as relicts (although advanced) of pangeic lineages when a primordial South American continent was still connected to the Antarctic Peninsula and, at the northern extreme, near the North American Plate. By the beginning of the Late Cretaceous, the volcanic and diastrophic processes that finally led to the differentiation of the Caribbean region and Central America built up transient geographic connections that permitted the initiation of an overland inter-American exchange that included, for example, dinosaurian titanosaurs from South America and hadrosaurs from North America. The immigration of other vertebrates followed the same route, for example, polydolopimorphian marsupials. These marsupials were assumed to have differentiated in South America prior to new discoveries from the North American Late Cretaceous. The complete extinction of endemic South American Mesozoic mammals by the Late Cretaceous-Early Paleocene, and the subsequent and in part coetaneous immigration of North American therians, respectively, represent two major moments in the history of South American mammals: a Gondwanan Episode and a South American Episode. The Gondwanan Episode was characterized by non- and pre-tribosphenic mammal lineages that descended from the Pangeic South American stage (but already with a pronounced Gondwanan accent, and wholly extinguished during the Late Cretaceous-Early Paleocene span). The South American Episode, in turn, was characterized only by therian mammals, mostly emigrated from the North American continent and already with a South American accent obtained through isolation. The southernmost extreme of South America (Patagonia) remained connected to the present Antarctic Peninsula at least up until about 30 Ma., and both provided the substratum where the primordial cladogenesis of “South American” mammals occurred. The resulting cladogenesis of South American therian mammals followed Gould's motto: early experimentation, later standardization. That is to say, early cladogenesis engendered a great variety of taxa with scarce morphological differentiation. After this early cladogenesis (Late Eocene-Early Oligocene), the variety of taxa became reduced, but each lineage became clearly recognizable distinctive by a constant morphologic pattern. At the same time, those mammals that underwent the “early experimentation” were part of communities dominated by archaic lineages (e.g., brachydont types among the native “ungulates”), whereas the subsequent communities were dominated by mammals of markedly “modern” stamp (e.g., protohypsodont types among the native “ungulates”). The Gondwanan and South American Episodes were separated by a critical latest Cretaceous-earliest Paleocene hiatus, it is as unknown as it is important in which South American land-mammal communities must have experienced extinction of the Gondwanan mammals and the arrival and radiation of the North American marsupials and placentals (with the probable exception of the xenarthrans, whose biogeographic origin is still unclear).     

A New European Marsupial Indicates a Late Cretaceous High-Latitude Transatlantic Dispersal Route

The first record of an undoubted opossum-like marsupial from the Mesozoic of Europe indicates an invasion from North America at the end of Late Cretaceous (Maastrichtian). The new 66.1 million-year-old marsupial, Maastrichtidelphys meurismeti n. gen., n. sp., represented by a right upper molar, comes from the type Maastrichtian of The Netherlands. The Maastricht marsupial exhibits affinities with earlier (early Maastrichtian) North American herpetotheriids providing definitive evidence of a high-latitude North Atlantic dispersal route between North America and Europe during the latest Cretaceous. Previously, the first major interchange for marsupials was thought to have occurred nearly 10 million years later in the Eocene. The occurrence of this new marsupial in Europe implies that at some time during the latest Cretaceous, sea level and climatic conditions must have been sufficiently favorable to allow for such a high-latitude dispersal. The fragmentary remains of hadrosaurid and theropod dinosaurs, as well as boid snakes from northwestern Europe which have affinities with North American taxa help substantiate assumptions made by the occurrence of the herpetotheriid marsupial in Maastricht.     

Diversity variation and tempo-spatial distributions of the Dipteridaceae ferns in the Mesozoic of China

The extant family Dipteridaceae is a remarkable leptosporangiate fern because it includes only one genus with a restricted distribution to tropical regions. The fossil record of this family has been widely reported from the Mesozoic strata in Eurasia, America, Australia, and Greenland. In China, numerous fossils of the Dipteridaceae have been documented, in total, about 74 species of 6 genera. Geographically, they are distributed both in the Southern and Northern Floristic Provinces, and were particularly well developed in the Southern Floristic Province during the Late Triassic and the Early Jurassic intervals. Fossil diversity of Dipteridaceae varies in the different episodes of the Mesozoic in China. It is shown that Dipteridaceae has undergone a diversity development process and a distinct turnover during the Mesozoic. They appear to have diversified in the warm and humid Late Triassic–Early Jurassic, but declined sharply as aridity developed in the Middle Jurassic, and became extinct at the end of the Early Cretaceous. The diversity variation and tempo-spatial distribution pattern is suggested to be linked with paleoclimatic variations during the Mesozoic.