Phylogeny and Systematics of Multituberculate Mammals

We present a synopsis of high-rank multituberculate systematicsand a manually generated cladogram illustrating multituberculate interrelationships. We divide the Multituberculata into the paraphyletic suborder 'Plagiaulacida', an apparently monophyletic suborder Cimolodonta, and one family incertae sedis. Within 'Plagiaulacida' we recognise three informal lines: paulchoffatiid (three families), plagiaulacid (three families) and allodontid (two families and the genus Glirodon). The Cimolodonta are divided into an informal Paracimexomys group; three superfamilies: Ptilodontoidea, Djadochtatherioidea (new), and Taeniolabidoidea (restricted to Taeniolabididae); and five families (superfamily incertae sedis): Eucosmodontidae, Microcosmodontidae, Cimolodontidae, Boffiidae, and Kogaionidae; and some genera incertae sedis. New characters used in our analysis are (1) a tendency of molar cusps to coalesce; and (2) ornamentation of grooves, pits, and ridges on the molars. We argue that the Ptilodontoidea, and less certainly also the Cimolodontidae and Boffiidae, might have originated from amongthe plagiaulacid line, a possible intermediate link being the Paracimexomys group. The remaining Cimolodonta might have originated from unknown members of the Paracimexomys group with separated molar cusps and smooth enamel. The origin of two types of prismatic enamel and a relationship between them are stumbling blocks in understanding the origin of the Cimolodonta; we conclude that microprismatic enamel made its appearance only once. Revised diagnoses of high-rank multituberculate taxa, including lists of all known genera, are given.     

Absence of ptilodontoidean multituberculates from Asia and its palaeogeographic implications

Gobibaatar parvus Kielan-Jaworowska 1970, formerly the only representative of the Ptilodontoidea in Asia, is recognized to be a junior synonym of Kryptobaatar dashzevegi Kielan-Jaworowska 1970 and allocated to the Taeniolabidoidea. Members of the Ptilodontoidea are thus currently unknown in Asian faunas. Chulsanbaataridae Kielan-Jaworowska 1974 is a junior synonym of the Eucosmodontidae Jepsen 1940. The absence of Ptilodontoidea in Asia gives further support to the theory of one-way Late Cretaceous dispersal of mammals from Asia to North America. The Marsupialia and Ptilodontoidea originated in the New World and never reached Asia, whereas the Taeniolabidoidea and Eutheria originated in Asia and dispersed to North America during the Late Cretaceous.     

早期哺乳动物三尖齿兽类牙齿的超微结构

本文用扫描电镜研究了中国云南禄丰盆地下禄丰组发现的以及英国威尔士晚三迭世的三尖齿兽类牙齿的超微结构.其釉质具有准釉柱结构,它们呈并行排列从釉牙质界延伸到釉质外表面.这种准釉柱结构由很多羽支状排列的磷灰石微晶组成.微晶的C轴与准釉柱的长轴形成交叉的角度大约为10-20度.早期哺乳动物牙齿釉质的准釉柱结构,很可能是从爬行动物无釉柱结构向哺乳动物真釉柱结构进化过程中的标志.     

Evolutionary analysis of the middle and inner ear of Late Jurassic multituberculates

We describe three previously unreported specimens of petrosal bones of paulchoffatiid multituberculate mammals, collected from strata of Late Jurassic age in the Guimarota lignite mine of Leiria, west-central Portugal. The new fossils allow correction, supplementation, and confirmation of anatomical details, thus refining knowledge of general adaptation in the ear region among Jurassic multituberculates. Virtually all observed characters in the paulchoffatiid otic region are primitive relative to homologous features seen among Late Cretaceous and younger representatives of the Multituberculata; we recognize few unique otic specializations in paulchoffatiids that would preclude ancestry to later multituberculates. The plesiomorphic nature of paulchoffatiid ear regions provides no evidence in support of the hypothesis of a special, sister-group relationship between multituberculates and Late Cretaceous/Cenozoic marsupials plus placentals. Used in isolation, objective evidence derived from paulchoffatiid ear regions is consistent with interpretation of multituberculate divergence from other mammals predating the stem to living monotremes and postdating the stem to extinct morganucodontids. More broadly based comparative studies among Mesozoic mammals, however, suggest that independent acquisition of similarly advanced mammalian features was a pervasive theme among evolutionary histories of early mammals, probably including multituberculates. Although the phylogenetic position of multituberculates relative to other mammalian groups has yet to be unequivocally resolved, we suggest that a very early divergence of the group remains a distinct possibility.     

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).     

<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.     

虔州豫俊兽(Yubaatarqianzhouensissp.nov.):江西赣州盆地首例晚白垩世哺乳动物化石*

本文报道了产自江西省赣州市上白垩统赣县河口组的一件多瘤齿兽类标本, 这是江西省报道的首例中生代哺乳动物化石。此标本头骨后部横向扩展, 额骨较小, 后端尖并构成眼眶的内侧边缘, M1具三列齿尖, 系统发育分析支持其归入纹齿兽超科。与河南晚白垩世的中原豫俊兽形态较为相似, 但是两者间也存在一些明显的区别, 因此建立一个豫俊兽属新种——虔州豫俊兽(Yubaatar qianzhouensis sp. nov.), 鉴定特征为: m1齿尖式为7︰6; M2齿尖式为1︰3︰3; m1颊侧后部存在一道小脊; 虔州豫俊兽m2和m1的长度比例小于中原豫俊兽; 冠状突呈楔状, 末端尖。虔州豫俊兽的发现不仅扩展了晚白垩世多瘤齿兽类在东亚地区的地理分布和物种多样性, 并且也扩展了中生代哺乳动物的地理分布。     

Evolution of metatherian and eutherian (mammalian) characters: a review based on cladistic methodology

Cladistic methodology is used to test the hypothesis that three major monophyletic groups exist among living mammals–the oviparous monotremes (Prototheria), and the viviparous marsupials (Metatheria) and placentals (Eutheria). Evaluation is made of the polarity (i.e. the direction of change in a primitive-to-derived sequence) of numerous characters which distinguish some or all of these groups, and of the usefulness of these characters in phylogenetic inference. An attempt is made to establish the state of these characters in the common Late Jurassic-Early Cretaceous therian ancestor of marsupials and placentals.
It is concluded that the most basic division of the Mammalia is the dichotomy into the subclasses Prototheria (including Monotremata, Multituberculata, Triconodonta, Docodonta) and Theria (including Metatheria, Eutheria, Pantotheria and Symmetrodonta). Two major groups exist among living viviparous mammals, the Metatheria and Eutheria; in a cladistic framework these are sister-groups. It is demonstrated that there is no special (sister-group) relationship between monotremes and marsupials, and there is no justification for placing them in a group Marsupionta.     

The origin and early evolution of metatherian mammals: the Cretaceous record

Metatherians, which comprise marsupials and their closest fossil relatives, were one of the most dominant clades of mammals during the Cretaceous and are the most diverse clade of living mammals after Placentalia. Our understanding of this group has increased greatly over the past 20 years, with the discovery of new specimens and the application of new analytical tools. Here we provide a review of the phylogenetic relationships of metatherians with respect to other mammals, discuss the taxonomic definition and diagnosis of Metatheria, outline the Cretaceous history of major metatherian clades, describe the paleobiology, biogeography, and macroevolution of Cretaceous metatherians, and provide a physical and climatic background of Cretaceous metatherian faunas. Metatherians are a clade of boreosphendian mammals that must have originated by the Late Jurassic, but the first unequivocal metatherian fossil is from the Early Cretaceous of Asia. Metatherians have the distinctive tightly interlocking occlusal molar pattern of tribosphenic mammals, but differ from Eutheria in their dental formula and tooth replacement pattern, which may be related to the metatherian reproductive process which includes an extended period of lactation followed by birth of extremely altricial young. Metatherians were widespread over Laurasia during the Cretaceous, with members present in Asia, Europe, and North America by the early Late Cretaceous. In particular, they were taxonomically and morphologically diverse and relatively abundant in the Late Cretaceous of western North America, where they have been used to examine patterns of biogeography, macroevolution, diversification, and extinction through the Late Cretaceous and across the Cretaceous-Paleogene (K-Pg) boundary. Metatherian diversification patterns suggest that they were not strongly affected by a Cretaceous Terrestrial Revolution, but they clearly underwent a severe extinction across the K-Pg boundary.     

The oldest higher true crabs (Crustacea: Decapoda: Brachyura): insights from the Early Cretaceous of the Americas

Despite the extensive fossil record of higher crabs (Eubrachyura) from Late Cretaceous and Cenozoic rocks worldwide, their Early Cretaceous occurrences are scarce and fragmentary, obscuring our understanding of their early evolution. Until now, representatives of only two families of eubrachyuran‐like crabs were known from the Early Cretaceous: Componocancridae and Tepexicarcinidae fam. nov., both monospecific lineages from the Albian (~110–100 Ma) of North and Central America, respectively. The discovery of Telamonocarcinus antiquus sp. nov. (Telamonocarcinidae) from the early Albian of Colombia, South America (~110 Ma), increases to three the number of known Early Cretaceous eubrachyuran‐like families. The ages and geographical distributions of the oldest eubrachyuran‐like taxa (i.e. Componocancridae, Telamonocarcinidae and Tepexicarcinidae fam. nov.) suggest that the oldest higher true crabs might have originated in the Americas; that they were already morphologically diverse by the late Early Cretaceous; and that their most recent common ancestor must be rooted in the Early Cretaceous, or even the Late Jurassic.     

The Phylogenetic Affinities of the Enigmatic Mammalian Clade Gondwanatheria

Gondwanatheria is a group of extinct mammals known from the Cretaceous and Paleogene of Gondwana. Resolution of the phylogenetic affinities of gondwanatherians has proven problematical, with the group currently considered Mammalia incertae sedis. We briefly review the morphology of known gondwanatherians, and argue that isolated upper premolars and a partial dentary preserving a blade-like p4 originally referred to the ferugliotheriid gondwanatherian Ferugliotherium windhauseni but subsequently identified as Multituberculata incertae sedis do indeed belong to F. windhauseni. We also suggest that the recently described ?cimolodontan multituberculate Argentodites coloniensis, based on an isolated lower premolar, may in fact be an unworn p4 of Ferugliotherium or a closely related taxon. We present the first phylogenetic analyses to include gondwanatherians, using maximum parsimony and Bayesian methods. Both methods place Ferugliotherium and sudamericid gondwanatherians in a clade with cimolodontan and “plagiaulacidan” multituberculates, although relationships within this clade are largely unresolved. The Gondwanatheria + Multituberculata clade supported here may reflect the convergent evolution of similar dental features, but it is the best supported hypothesis based on currently available data. However, denser sampling of multituberculate taxa and the discovery of more complete gondwanatherian fossils will be required to clarify the precise relationship between gondwanatherians and multituberculates, specifically to determine whether or not gondwanatherians are members of Multituberculata. We hypothesize that the anterior molariforms of sudamericid gondwanatherians evolved from blade-like precursors similar to the p4 of Ferugliotherium, possibly in response to the appearance of grasses in Gondwana during the Cretaceous.     

New Studies of Enamel Microstructure in Mesozoic Mammals: A Review of Enamel Prisms as a Mammalian Synapomorphy

Characters from enamel microstructure have not been used in recent phylogenetic analyses of Mesozoic Mammalia. Reasons are that enamel characters have been perceived as (A) variable without regard to systematic position of taxa, (B) inconsistently reported within the literature, and (C) simply scored as either prismatic or not prismatic in earlier mammals. Our work on Mesozoic mammals such as Sinoconodon, Gobiconodon, Triconodontidae, Docodon, Laolestes, and others suggests that synapsid columnar enamel (SCE) structure was easily transformed into plesiomorphic prismatic enamel (PPE) and that PPE may be described with at least five independent character states. Two PPE characters—a flat, open prism sheath and a planar prism seam—were present in the cynodont Pachygenelus and in several Jurassic and Cretaceous mammals. We propose that appearance of a prism sheath transforms SCE into PPE and that reduction and loss of a prism sheath reverse PPE into SCE, in both phylogeny and ontogeny. We further propose that no amniote vertebrates other than the trithelodontid cynodont, Pachygenelus, plus Mammalia have ever evolved an ameloblastic Tomes process capable of secreting PPE and that the genetic potential to secrete PPE is a synapomorphy of Pachygenelus plus Mammalia, whether or not all lineages of the clade have expressed that potential.     

Mammalian tooth marks on the bones of dinosaurs and other Late Cretaceous vertebrates

Abstract: We describe bones from the Late Cretaceous of Alberta – including bones of large dinosaurs, a femur from the aquatic reptile Champsosaurus, and a dentary from the marsupial Eodelphis– that bear tooth marks made by animals with opposing pairs of teeth. Of the animals known from the Late Cretaceous of North America, only mammals are capable of making such tooth marks. In particular, multituberculates, which have paired upper and lower incisors, are the most likely candidates for the makers of these traces. The traces described here represent the oldest known mammalian tooth marks. Although it is possible that some of these tooth marks represent feeding traces, the tooth marks often penetrate deep into the dense cortices of the bone. This raises the possibility that, much as extant mammals gnaw bone and antler, some Cretaceous mammals may have consumed the bones of dinosaurs and other vertebrates as a source of minerals. However, none of the tooth marks described here resemble the extensive gnaw traces produced by Cenozoic multituberculates or rodents. This suggests that specialized gnawing forms may have been rare or absent in the Late Cretaceous of North America.     

Revision of <Emphasis Type="Italic">Actinastrea</Emphasis>, the most common Cretaceous coral genus

The very common and species-rich Scleractinian genus Actinastrea (family Actinastraeidae, suborder Archeocaeniina) is revised on the basis of the type material of its type species and additional material from the type locality. A lectotype is designated for the type species. It was discovered that Jurassic to Early Cretaceous corals currently assigned to Actinastrea do not fit into the concept of this genus. These species belong to the genus Stelidioseris, which is also revised on the basis of the type of the type species, including designating a lectotype. These two genera are distinguished by various characteristics: septal external parts are swollen in Actinastrea but not in Stelidioseris, the costae are confluent in Stelidioseris but not in Actinastrea, the coenosteum is granulated in Actinastrea but narrow than in Actinastrea and only with costae in Stelidioseris. Actinastrea is restricted to the Late Cretaceous (Late Turonian—Maastrichtian), whereas Stelidioseris originates in the Jurassic and reaches into the Late Cretaceous, but is less common from the Turonian on.     

INTRODUCTION TO MESOZOIC BIRDS FROM LIAONING,CHINA

l.IntroductionThestudyofMesozoicbirdsinChinadatedbacktotheearlyeightiesofthiscentury,whenGansuswasdiscoveredanddescribed(HouetLiu,l984).Sincethelateeighties,anumberofEarlyCretaceousbirdshavebeenfound,firstlyinWesternLiaoning(Zhou,l995),andshortlylaterinInnerMongolia(Dong,l993;Hou,l994)andHebeiProvince.Atthesametime,featherimpressionswerealsorecoveredinShandongProvince(Zhang,l992)andNingxiaAutonomousRegion.Particularlyimpor-tantisthatsincel994someLateJurassicbirdshavebeenfoundfromtheY…     

Paramesopsocidae,a new Mesozoic psocid family (Insecta: Psocodea “Psocoptera”: Psocomorpha)

Paramesopsocus lu n. gen., n. sp. and Paramesopsocus adibi n. sp. are respectively described from the Early Cretaceous amber of Lebanon and from the Late Jurassic limestone of Karatau (Kazakhstan). They are placed within the suborder Psocomorpha, and in the Mesozoic extinct family Paramesopsocidae n. fam. A cladistic phylogeny for Psocomorpha is given including our fossil taxa. The discovery of these new taxa demonstrates the necessity of a deep cladistic redefinition of the currently admitted major subdivisions of this suborder.     

A basal thunnosaurian from Iraq reveals disparate phylogenetic origins for Cretaceous ichthyosaurs

Cretaceous ichthyosaurs have typically been considered a small, homogeneous assemblage sharing a common Late Jurassic ancestor. Their low diversity and disparity have been interpreted as indicative of a decline leading to their Cenomanian extinction. We describe the first post-Triassic ichthyosaur from the Middle East, Malawania anachronus gen. et sp. nov. from the Early Cretaceous of Iraq, and re-evaluate the evolutionary history of parvipelvian ichthyosaurs via phylogenetic and cladogenesis rate analyses. Malawania represents a basal grade in thunnosaurian evolution that arose during a major Late Triassic radiation event and was previously thought to have gone extinct during the Early Jurassic. Its pectoral morphology appears surprisingly archaic, retaining a forefin architecture similar to that of its Early Jurassic relatives. After the initial latest Triassic radiation of early thunnosaurians, two subsequent large radiations produced lineages with Cretaceous representatives, but the radiation events themselves are pre-Cretaceous. Cretaceous ichthyosaurs therefore include distantly related lineages, with contrasting evolutionary histories, and appear more diverse and disparate than previously supposed.     

Eutherians experienced elevated evolutionary rates in the immediate aftermath of the Cretaceous–Palaeogene mass extinction

The effect of the Cretaceous–Palaeogene (K–Pg) mass extinction on the evolution of many groups, including placental mammals, has been hotly debated. The fossil record suggests a sudden adaptive radiation of placentals immediately after the event, but several recent quantitative analyses have reconstructed no significant increase in either clade origination rates or rates of character evolution in the Palaeocene. Here we use stochastic methods to date a recent phylogenetic analysis of Cretaceous and Palaeocene mammals and show that Placentalia likely originated in the Late Cretaceous, but that most intraordinal diversification occurred during the earliest Palaeocene. This analysis reconstructs fewer than 10 placental mammal lineages crossing the K–Pg boundary. Moreover, we show that rates of morphological evolution in the 5 Myr interval immediately after the K–Pg mass extinction are three times higher than background rates during the Cretaceous. These results suggest that the K–Pg mass extinction had a marked impact on placental mammal diversification, supporting the view that an evolutionary radiation occurred as placental lineages invaded new ecological niches during the Early Palaeocene.     

Early Cenozoic increases in mammal diversity cannot be explained solely by expansion into larger body sizes

A prominent hypothesis in the diversification of placental mammals after the Cretaceous–Palaeogene (K/Pg) boundary suggests that the extinction of non-avian dinosaurs resulted in the ecological release of mammals, which were previously constrained to small body sizes and limited species richness. This ‘dinosaur incumbency hypothesis’ may therefore explain increases in mammalian diversity via expansion into larger body size niches, that were previously occupied by dinosaurs, but does not directly predict increases in other body size classes. To evaluate this, we estimate sampling-standardized diversity patterns of terrestrial North American fossil mammals within body size classes, during the Cretaceous and Palaeogene. We find strong evidence for post-extinction diversity increases in all size classes. Increases in the diversity of small-bodied species (less than 100 g, the common body size class of Cretaceous mammals, and much smaller than the smallest non-avialan dinosaurs (c. 400 g)) were similar to those of larger species. We propose that small-bodied mammals had access to greater energetic resources or were able to partition resources more finely after the K/Pg mass extinction. This is likely to be the result of a combination of widespread niche clearing due to the K/Pg mass extinctions, alongside a suite of biotic and abiotic changes that occurred during the Late Cretaceous and across the K/Pg boundary, such as shifting floral composition, and novel key innovations among eutherian mammals.