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The evolution of tooth enamel microstructure in both extinct and extant mammalian groups has been extensively documented, but is poorly known in reptiles, including dinosaurs. Previous intensive sampling of dinosaur tooth enamel microstructure revealed that: (1) the three‐dimensional arrangement of enamel types and features within a tooth—the schmelzmuster—is most useful in diagnosing dinosaur clades at or around the family level; (2) enamel microstructure complexity is correlated with tooth morphology complexity and not necessarily with phylogenetic position; and (3) there is a large amount of homoplasy within Theropoda but much less within Ornithischia. In this study, the examination of the enamel microstructure of 28 additional dinosaur taxa fills in taxonomic gaps of previous studies and reinforces the aforementioned conclusions. Additionally, these new specimens reveal that within clades such as Sauropodomorpha, Neotheropoda, and Euornithopoda, the more basal taxa have simpler enamel that is a precursor to the more complex enamel of more derived taxa and that schmelzmusters evolve in a stepwise fashion. In the particularly well‐sampled clade of Euornithopoda, correlations between the evolution of dental and enamel characters could be drawn. The ancestral schmelzmuster for Genasauria remains ambiguous due to the dearth of basal ornithischian teeth available for study. These new specimens provide new insights into the evolution of tooth enamel microstructure in dinosaurs, emphasizing the importance of thorough sampling within broadly inclusive clades, especially among their more basal members.  相似文献   

3.
Nucleotide sequence data from the mitochondrial 12S rRNA and cytochrome b genes were used to analyze phylogenetic relationships among sciurognath rodents. Our sample taxa included representatives of 11 sciurognath and 3 hystricognath families with two marsupial species, Didelphis virginiana and Macropus robustus, as outgroups. The dataset was analyzed using both maximum-parsimony (weighted and unweighted) and likelihood methods. Three suprafamilial groupings are strongly supported: Geomyidae + Heteromyidae (Geomyoidea), Sciuridae + Aplodontidae (Sciuroidea), and Pedetidae + Anomaluridae (Anomaluroidea). Although moderately supported, two sister group relationships were identified between Gliridae and Sciuroidea and between Castor and Geomyoidea. In contrast to previous nuclear DNA evidence, the evolutionary affinities between Ctenodactylidae and Hystricognathi (Ctenohystrica) and between Muridae and Dipodidae (Myodonta) are not supported by the mitochondrial data. Molecular divergence dates based on the combined data were estimated for suprafamilial groupings and are discussed in the light of current morphological and paleontological interpretations of rodent phylogeny.  相似文献   

4.
Many genera of arvicolid rodents (voles) contain species that spend a considerable amount of time underground. Incisors provide a considerable effort in the digging process and a difference in the enamel microstructure of incisors is expected between species that spend most of their life above ground and those who have greater specializations for a subterranean habitat. The ratio between the thicknesses of the two layers composing the enamel of the incisors has proved to be an effective proxy to infer fossorial activity of extinct forms of Arvicola but this ratio exhibited no phylogenetic signal (although lifestyle was in a greater extent related to phylogeny). We were able to infer ancestral lifestyles (of extinct populations and of nodes of the phylogeny) from enamel morphologies. Inclusion of fossils did not solve the uncertainties associated with the lifestyle for the last common ancestor of extant Arvicola species and it did not change the inferred lifestyle of several other, less inclusive, clades.  相似文献   

5.
There are two types of cheek pouches in extant rodents. Internal cheek pouches are evaginations of the oral cavity deep to M. platysma and M. sphincter colli profundus, and have evolved independently in some species of the superfamilies Sciuroidea and Muroidea. External, furlined cheek pouches open lateral to and separate from the oral cavity, (also deep to M. platysma and M. sphincter colli profundus), and occur in all species of the families Geomyidae and Heteromyidae. The presence of external, furlined cheek pouches is a synapomorphy for the superfamily Geomyoidea. The posterior retractor muscle of the pouch is derived from facial musculature in sciurids, from trapezius musculature in cricetids, and from both facial and trapezius muscle groups in the Geomyoidea. Differences also exist in the musculature associated with the pouch opening. In the Sciuridae and Cricetidae, the M. buccinatorius muscle group acts as a sphincter to control the size of the pouch opening. In the Geomyoidea, the size of the opening is controlled by the M. orbicularis sacculi in concert with a slip of the M. platysma myoides. Thin sections and scanning electron micrographs of the pouch tissue reveal the presence of dermal papillae in Phodopus sungorus but not in a close relative, Mesocricetus auratus. All members of the subfamily Cricetinae have a peninsula of highly folded tissue projecting anteriorly from the posteromedial pouch wall. This folding allows for expansion of the pouch walls when food is stored in the pouch.  相似文献   

6.
To investigate the diversity of the enamel structures in rodent molars, the schmelzmuster of more than 270 genera from the various fossil and extant groups was investigated. Only three basic types of schmelzmuster were recognized. The most prominent one, the C-type, is characterized by a basal ring of lamellar enamel (BRLE) surrounding the molars at the base of the crown. It occurs in most murids (cricetids) and other Myodonta. It is shared by the Dipodidea, Eomyidae, Geomyidae, and Gliridae. The C-type schmelzmuster was not found in any of the other rodents groups (Sciuromorpha, Anomaluromorpha, Sciuravida, Hystricognathi, Bathyergomorphi, and Caviida). Despite the rare occurrence of the P-type schmelzmuster with radial enamel only, most of these rodents have thick Hunter–Schreger bands (HSB) in their molars (S-type schmelzmuster). Although the C-type schmelzmuster is strictly limited to the Myomorpha (sensu McKenna and Bell), the fossil record shows that this structure does not form a synapomorphy but developed in parallel mostly from the P-type, but in Gliridae from the S-type schmelzmuster. On the basis of molar and incisor enamel the systematic position of Protoptychus can be evaluated.  相似文献   

7.
In the enamel microstructure of the incisors, porcupines (Hystricidae) are characterized by multiserial Hunter-Schreger bands (HSB) while beavers (Castoridae) basically have uniserial HSB. In addition, two groups can be differentiated among beavers based on enamel microstructure.Palaeocastor, Steneofiber andCastor show rather typical uniserial HSB. OnlyCastor shows a slight tendency of fusion of the bands. In contrast,Trogontherium andCastoroides are characterized by the regular occurrence of fused bands.Anchitheriomys, having uniserial HSB and a high frequency of fused bands fits well with theTrogontherium group, proving the castorid nature ofAnchitheriomys. Furthermore, the enamel differentiation observed among the Castoridae thus provides an important character for further systematic subdivision of the beavers. In the enamel of the cheek teeth, porcupines can be told apart from beavers as well.Anchitheriomys shows a two-layered schmelzmuster with inner radial enamel like other beavers. Porcupines have thick HSB throughout the enamel layer. This character in the cheek teeth facilitates the separation of isolated beaver teeth from porcupine teeth.  相似文献   

8.
Afropithecus turkanensis, a 17-17.5 million year old large-bodied hominoid from Kenya, has previously been reported to be the oldest known thick-enamelled Miocene ape. Most investigations of enamel thickness in Miocene apes have been limited to opportunistic or destructive studies of small samples. Recently, more comprehensive studies of enamel thickness and microstructure in Proconsul, Lufengpithecus, and Dryopithecus, as well as extant apes and fossil humans, have provided information on rates and patterns of dental development, including crown formation time, and have begun to provide a comparative context for interpretation of the evolution of these characters throughout the past 20 million years of hominoid evolution. In this study, enamel thickness and aspects of the enamel microstructure in two A. turkanensis second molars were quantified and provide insight into rates of enamel apposition, numbers of cells actively secreting enamel, and the time required to form regions of the crown. The average value for relative enamel thickness in the two molars is 21.4, which is a lower value than a previous analysis of this species, but which is still relatively thick compared to extant apes. This value is similar to those of several Miocene hominoids, a fossil hominid, and modern humans. Certain aspects of the enamel microstructure are similar to Proconsul nyanzae, Dryopithecus laietanus, Lufengpithecus lufengensis, Graecopithecus freybergi and Pongo pygmaeus, while other features differ from extant and fossil hominoids. Crown formation times for the two teeth are 2.4-2.6 years and 2.9-3.1 years respectively. These times are similar to a number of extant and fossil hominoids, some of which appear to show additional developmental similarities, including thick enamel. Although thick enamel may be formed through several developmental pathways, most Miocene hominoids and fossil hominids with relatively thick enamel are characterized by a relatively long period of cuspal enamel formation and a rapid rate of enamel secretion throughout the whole cusp, but a shorter total crown formation time than thinner-enamelled extant apes.  相似文献   

9.
Fossil pollen believed to be related to extant Hagenia abyssinica were discovered in the early Miocene (21.73 Ma) Mush Valley paleoflora, Ethiopia, Africa. Both the fossil and extant pollen grains of H. abyssinica were examined with combined light microscopy, scanning electron microscopy, and transmission electron microscopy to compare the pollen and establish their relationships. Based on this, the fossil pollen grains were attributed to Hagenia. The presence of Hagenia in the fossil assemblage raises the questions if its habitat has changed over time, and if the plants are/were wind pollinated. To shed light on these questions, the morphology of extant anthers was also studied, revealing specialized hairs inside the anthers, believed to aid in insect pollination. Pollen and anther morphology are discussed in relation to the age and origin of the genus within a molecular dated phylogenetic framework, the establishment of complex topography in East Africa, other evidence regarding pollination modes, and the palynological record. The evidence presented herein, and compiled from the literature, suggests that Hagenia was an insect‐pollinated lowland rainforest element during the early Miocene of the Mush Valley. The current Afromontane habitat and ambophilous (insect and wind) pollination must have evolved in post‐mid‐Miocene times.  相似文献   

10.
Traditional contributions of the insect fossil record are listed. Fossil material indicates the earliest occurrence of a group, which in turn is useful for inferring clade divergence dates and net diversification rates. Fossil material provides complementary information on the dynamics of taxonomic diversity. Geographical occurrences outside the extant range of a taxon can be used to infer climatic macro‐fluctuations. In short, the fossil record of insects is essential for pointing out the major factors responsible for the mega‐diversity of the group, and of some of its internal lineages. Reliable taxonomic assignments and phylogenetic hypotheses underpin broader generalizations. In that respect, a problem is the inadequate integration of data from fossil and extant insect taxa in phylogenetic investigations. Stumbling blocks lie at various systematic levels. Unreliability of specimen‐based data, of species delimitation, and of homology assumptions, might have been responsible for a disdain by some entomologists for palaeoentomological literature. Idiosyncratic (and in cases flawed) methods aimed at investigating phylogenetic relationships used by a fraction of the palaeoentomological community might also have contributed to this situation. Concurrently, the traditional nomenclatural procedure might prevent effective communication between neo‐ and palaeoentomologists. Augmenting the available information on the wing venation of extant taxa would significantly advance palaeoentomology, and provide a relevant broad‐scale character system. Furthermore, the entomological community should contribute to experimentations of various nomenclatural procedures, with the aim of developing an optimal approach in terms of communication and information retrieval.  相似文献   

11.
We describe the first endocast reconstruction of a hyaenodont mammal based on X‐ray microtomography. The endocast belongs to the type material of the European hyaenodont Proviverra typica. We performed phylogenetic analysis to contextualize the evolution of endocranial size and complexity in Hyaenodonta. We added several European hyaenodonts and modified several codings of the most recent character–taxon matrix established to question the relationships within Hyaenodonta. Including these new species in a phylogenetic analysis reveals a new clade: Hyaenodontoidea. Comparisons with several previously described endocasts show that there was an increase in complexity in the convolutions of the encephalon within Hyaenodontidae history. Moreover, the analysis of the encephalization quotient reveals that the endocranium of the Hyaenodonta is not smaller than those of fossil Carnivora or some extant Carnivora. Therefore, the extinction of Hyaenodonta may not be linked to the relative size of hyaenodont brains.  相似文献   

12.
It is widely acknowledged that integrating fossils into data sets of extant taxa is imperative for proper placement of fossils, resolution of relationships, and a better understanding of character evolution. The importance of this process has been further magnified because of the crucial role of fossils in dating divergence times. Outstanding issues remain, including appropriate methods to place fossils in phylogenetic trees, the importance of molecules versus morphology in these analyses, as well as the impact of potentially large amounts of missing data for fossil taxa. In this study we used the angiosperm clade Juglandaceae as a model for investigating methods of integrating fossils into a phylogenetic framework of extant taxa. The clade has a rich fossil record relative to low extant diversity, as well as a robust molecular phylogeny and morphological database for extant taxa. After combining fossil organ genera into composite and terminal taxa, our objectives were to (1) compare multiple methods for the integration of the fossils and extant taxa (including total evidence, molecular scaffolds, and molecular matrix representation with parsimony [MRP]); (2) explore the impact of missing data (incomplete taxa and characters) and the evidence for placing fossils on the topology; (3) simulate the phylogenetic effect of missing data by creating "artificial fossils"; and (4) place fossils and compare the impact of single and multiple fossil constraints in estimating the age of clades. Despite large and variable amounts of missing data, each of the methods provided reasonable placement of both fossils and simulated "artificial fossils" in the phylogeny previously inferred only from extant taxa. Our results clearly show that the amount of missing data in any given taxon is not by itself an operational guideline for excluding fossils from analysis. Three fossil taxa (Cruciptera simsonii, Paleoplatycarya wingii, and Platycarya americana) were placed within crown clades containing living taxa for which relationships previously had been suggested based on morphology, whereas Polyptera manningii, a mosaic taxon with equivocal affinities, was placed firmly as sister to two modern crown clades. The position of Paleooreomunnea stoneana was ambiguous with total evidence but conclusive with DNA scaffolds and MRP. There was less disturbance of relationships among extant taxa using a total evidence approach, and the DNA scaffold approach did not provide improved resolution or internal support for clades compared to total evidence, whereas weighted MRP retained comparable levels of support but lost crown clade resolution. Multiple internal minimum age constraints generally provided reasonable age estimates, but the use of single constraints provided by extinct genera tended to underestimate clade ages.  相似文献   

13.
Relationships among extant and fossil echimyids (Rodentia: Hystricognathi)   总被引:1,自引:0,他引:1  
The echimyid rodents are the most diverse group of Neotropical hystricognaths, with approximately 40 extant and fossil genera. Craniodental characters are proposed in order to formulate hypotheses of phylogenetic relationships within the Echimyidae. A data matrix of 54 taxa and 50 characters is constructed and submitted to parsimony analyses using PAUP and WinClada programs. Analysis of the complete data set results in 47 448 most parsimonious trees 107 steps long. These trees are summarized in a strict consensus tree, which is taken as the main phylogenetic hypothesis resulting from this study. The monophyly of several currently recognized supraspecific taxa is not corroborated. These are: the subfamilies Eumysopinae, Echimyinae, Myocastorinae and Adelphomyinae; and the genera Proechimys , Echimys and Makalata . Conversely, the monophyly of Dactylomyinae and Trinomys is supported. New associations are proposed: (1) a clade comprising the extant Carterodon , Clyomys and Euryzygomatomys and the fossil Pampamys and Theridomysops placed at the base of the crown-group Echimyidae; (2) a clade uniting Proechimys , Hoplomys and Trinomys , which is the sister-taxon of (3) a clade including Mesomys , Lonchothrix , Myocastor and a clade with extant dactylomyines and echimyines and associated fossil taxa. Based on this phylogenetic hypothesis, patterns of tooth evolution in Echimyidae are discussed, and minimum ages for the divergence events within the family are estimated.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 445–477.  相似文献   

14.
The estimation of phylogenetic relationships and divergence times among a group of organisms is a fundamental first step toward understanding its biological diversification. The time of the most recent or last common ancestor (LCA) of extant platyrrhines is one of the most controversial among scholars of primate evolution. Here we use two molecular based approaches to date the initial divergence of the platyrrhine clade, Bayesian estimations under a relaxed-clock model and substitution rate plus generation time and body size, employing the fossil record and genome datasets. We also explore the robustness of our estimations with respect to changes in topology, fossil constraints and substitution rate, and discuss the implications of our findings for understanding the platyrrhine radiation. Our results suggest that fossil constraints, topology and substitution rate have an important influence on our divergence time estimates. Bayesian estimates using conservative but realistic fossil constraints suggest that the LCA of extant platyrrhines existed at ca. 29 Ma, with the 95% confidence limit for the node ranging from 27–31 Ma. The LCA of extant platyrrhine monkeys based on substitution rate corrected by generation time and body size was established between 21–29 Ma. The estimates based on the two approaches used in this study recalibrate the ages of the major platyrrhine clades and corroborate the hypothesis that they constitute very old lineages. These results can help reconcile several controversial points concerning the affinities of key early Miocene fossils that have arisen among paleontologists and molecular systematists. However, they cannot resolve the controversy of whether these fossil species truly belong to the extant lineages or to a stem platyrrhine clade. That question can only be resolved by morphology. Finally, we show that the use of different approaches and well supported fossil information gives a more robust divergence time estimate of a clade.  相似文献   

15.
The phylogenetic position of the Pedetidae, represented by a single species Pedetes capensis, is controversial, reflecting in part the retention of both Hystricomorphous and Sciurognathous characteristics in this rodent. In an attempt to clarify the species evolutionary relationships, mtDNA gene sequences from 10 rodent species (representing seven families) were analyzed using phenetic, parsimony, and maximum-likelihood methods of phylogenetic inference; the rabbit, Oryctolagus cuniculus (Order Lagomorpha), and cow, Bos taurus (Order Artiodactyla), were used as outgroups. Investigation of 714 base pairs of the protein-coding cytochrome b gene indicate strong base bias at the third codon position with significant rate heterogeneity evident between the three structural domains of this gene. Similar analyses conducted on 816 base pairs of the 12S rRNA gene revealed a transversion bias in the loop sections of all taxa. The cytochrome b gene sequences proved useful in resolving associations between closely related species but failed to produce consistent tree topologies at the family level. In contrast, phylogenetic analysis of the 12S rRNA gene resulted in strong support for the clustering of Pedetidae/Heteromyidae/Geomyidae and Muridae in one clade to the exclusion of the Hystricidae/Thryonomyidae and Sciuridae, a finding which is concordant with studies of rodent fetal membranes as well as reproductive and other anatomical features.   相似文献   

16.
Lampridiformes is a peculiar clade of pelagic marine acanthomorph (spiny‐rayed) teleosts. Its phylogenetic position remains ambiguous, and varies depending on the type of data (morphological or molecular) used to infer interrelationships. Because the extreme morphological specializations of lampridiforms may have overwritten the ancestral features of the group with a bearing on its relationships, the inclusion of fossils that exhibit primitive character state combinations for the group as a whole is vital in establishing its phylogenetic position. Therefore, we present an osteological data set of extant (ten taxa) and fossil (14 taxa) acanthomorphs, including early Late Cretaceous taxa for which a close relationship with extant Lampridiformes has been suggested: ?Aipichthyoidea, ?Pharmacichthyidae, and ?Pycnosteroididae. We find that all three taxa plus Lampridiformes form a clade that we call Lampridomorpha. Under this hypothesis, ?Aipichthyoidea is paraphyletic. The inclusion of fossils in the analysis changes the topology, highlighting their critical importance in phylogenetic studies of morphological characters. When fossils are included, Lampridomorpha is sister to Euacanthomorpha (all other extant acanthomorphs), concurring with most previous anatomical studies, but conflicting with most molecular results. Lampridomorpha as a whole was a major component of the earliest acanthomorph faunas, notably in the Cenomanian. © 2014 The Linnean Society of London  相似文献   

17.
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.  相似文献   

18.
The enamel microstructure in molars of Arsinoitherium is reinvestigated and a new modification of radial enamel (RE), ‘arsinoitheriid radial enamel (ARE)’, is defined. It is characterised by alternating stripes with different organisation of the interprismatic matrix but no prism decussation. Recognition of this new subtype leads to a reinterpretation of structure previously identified as modified radial enamel and of Hunter–Schreger bands in Arsinoitherium. The newly differentiated ARE of Arsinoitherium is more derived in relation to corresponding microstructures of Palaeoamasia and Crivadiatherium. A careful reinvestigation of RE in other Paenungulata will be required to provide additional data bearing on phylogenetic reconstruction. The enamel of Phenacolophus argues against inclusion of this genus in the Embrithopoda.  相似文献   

19.
Living fossils are lineages that have retained plesiomorphic traits through long time periods. It is expected that such lineages have both originated and diversified long ago. Such expectations have recently been challenged in some textbook examples of living fossils, notably in extant cycads and coelacanths. Using a phylogenetic approach, we tested the patterns of the origin and diversification of liphistiid spiders, a clade of spiders considered to be living fossils due to their retention of arachnid plesiomorphies and their exclusive grouping in Mesothelae, an ancient clade sister to all modern spiders. Facilitated by original sampling throughout their Asian range, we here provide the phylogenetic framework necessary for reconstructing liphistiid biogeographic history. All phylogenetic analyses support the monophyly of Liphistiidae and of eight genera. As the fossil evidence supports a Carboniferous Euramerican origin of Mesothelae, our dating analyses postulate a long eastward over-land dispersal towards the Asian origin of Liphistiidae during the Palaeogene (39–58 Ma). Contrary to expectations, diversification within extant liphistiid genera is relatively recent, in the Neogene and Late Palaeogene (4–24 Ma). While no over-water dispersal events are needed to explain their evolutionary history, the history of liphistiid spiders has the potential to play prominently in vicariant biogeographic studies.  相似文献   

20.
The shape of the enamel-dentine junction (EDJ) in primate molars is regarded as a potential indicator of phylogenetic relatedness because it may be morphologically more conservative than the outer enamel surface (OES), and it may preserve vestigial features (e.g., cuspules, accessory ridges, and remnants of cingula) that are not manifest at the OES. Qualitative accounts of dentine-horn morphology occasionally appear in character analyses, but little has been done to quantify EDJ shape in a broad taxonomic sample. In this study, we examine homologous planar sections of maxillary molars to investigate whether measurements describing EDJ morphology reliably group extant anthropoid taxa, and we extend this technique to a small sample of fossil catarrhine molars to assess the utility of these measurements in the classification of fossil teeth. Although certain aspects of the EDJ are variable within a taxon, a taxon-specific cross-sectional EDJ configuration predominates. A discriminant function analysis classified extant taxa successfully, suggesting that EDJ shape may a reliable indicator of phyletic affinity. When considered in conjunction with aspects of molar morphology, such as developmental features and enamel thickness, EDJ shape may be a useful tool for the taxonomic assessment of fossil molars.  相似文献   

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