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1.
Hupehsuchia is a group of enigmatic Triassic marine reptiles that is known exclusively from two counties in Hubei Province, China. One of the common features of the group was a modestly long neck with nine to ten cervical vertebrae. We report a new species of Hupehsuchia, Eohupehsuchus brevicollis gen. et sp. nov., which for the first time shows a short neck in this group, with six cervicals. The configuration of the skull roof in Eohupehsuchus is also unique among Hupehsuchia, with narrow frontals and posteriorly shifted parietals, warranting recognition of a new species. The taxon superficially resembles Nanchangosaurus in retaining hupehsuchian plesiomorphies, such as low neural spines and small body size. However, its limbs are well-developed, unlike in Nanchangosaurus, although the latter genus is marginally larger in body length. Thus, the individual is unlikely to be immature. Also, Eohupehsuchus shares a suite of synapomorphies with Hupehsuchus, including the second and third layers of dermal ossicles above the dorsal neural spines. A phylogenetic analysis suggests that the new species is not the most basal hupehsuchian despite its short neck, and instead forms the sister taxon of Hupehsuchidae. Until recently, Hupehsuchia contained only two monotypic genera. Now there are at least four genera among Hupehsuchia, and the undescribed diversity is even higher. The left forelimb of the only specimen is incomplete, ending with broken phalanges distally. The breakage could only have occurred pre-burial. The individual may have been attacked by a predator and escaped, given that scavenging is unlikely.  相似文献   

2.
The viviparous sea snakes (Hydrophiinae) comprise ~90% of living marine reptiles and display many physical and behavioral adaptations for breathing, diving, and achieving osmotic balance in marine habitats. Among the most important innovations found in marine snakes are their paddle-shaped (dorsoventrally expanded) tails, which provide propulsive thrust in the dense aquatic medium. Here, we reconstruct the evolution of caudal paddles in viviparous sea snakes using a dated molecular phylogeny for all major lineages and computed tomography of internal osteological structures. Bayesian ancestral state reconstructions show that extremely large caudal paddles supported by elongated vertebral processes are unlikely to have been present in the most recent common ancestor of extant sea snakes. Instead, these characters appear to have been acquired independently in two highly marine lineages of relatively recent origin. Both the Aipysurus and Hydrophis lineages have elongated neural spines that support the dorsal edge of their large paddles. However, whereas in the Aipysurus lineage the ventral edge of the paddle is supported by elongated haemapophyses, this support is provided by elongated and ventrally directed pleurapophyses in the Hydrophis lineage. Three semi-marine lineages (Hydrelaps, Ephalophis, and Parahydrophis) form the sister group to the Hydrophis clade and have small paddles with poorly developed dorsal and ventral supports, consistent with their amphibious lifestyle. Overall, our results suggest that not only are the viviparous hydrophiines the only lineage of marine snakes to have acquired extremely large, skeletally supported caudal paddles but also that this innovation has occurred twice in the group in the past ~2-6 million years.  相似文献   

3.

Background

Secondary adaptation to aquatic life occurred independently in several amniote lineages, including reptiles during the Mesozoic and mammals during the Cenozoic. These evolutionary shifts to aquatic environments imply major morphological modifications, especially of the feeding apparatus. Mesozoic (250–65 Myr) marine reptiles, such as ichthyosaurs, plesiosaurs, mosasaurid squamates, crocodiles, and turtles, exhibit a wide range of adaptations to aquatic feeding and a broad overlap of their tooth morphospaces with those of Cenozoic marine mammals. However, despite these multiple feeding behavior convergences, suction feeding, though being a common feeding strategy in aquatic vertebrates and in marine mammals in particular, has been extremely rarely reported for Mesozoic marine reptiles.

Principal Findings

A relative of fossil protostegid and dermochelyoid sea turtles, Ocepechelon bouyai gen. et sp. nov. is a new giant chelonioid from the Late Maastrichtian (67 Myr) of Morocco exhibiting remarkable adaptations to marine life (among others, very dorsally and posteriorly located nostrils). The 70-cm-long skull of Ocepechelon not only makes it one of the largest marine turtles ever described, but also deviates significantly from typical turtle cranial morphology. It shares unique convergences with both syngnathid fishes (unique long tubular bony snout ending in a rounded and anteriorly directed mouth) and beaked whales (large size and elongated edentulous jaws). This striking anatomy suggests extreme adaptation for suction feeding unmatched among known turtles.

Conclusion/Significance

The feeding apparatus of Ocepechelon, a bony pipette-like snout, is unique among tetrapods. This new taxon exemplifies the successful systematic and ecological diversification of chelonioid turtles during the Late Cretaceous. This new evidence for a unique trophic specialization in turtles, along with the abundant marine vertebrate faunas associated to Ocepechelon in the Late Maastrichtian phosphatic beds of Morocco, further supports the hypothesis that marine life was, at least locally, very diversified just prior to the Cretaceous/Palaeogene (K/Pg) biotic crisis.  相似文献   

4.
The Ichthyosauria is the group of Mesozoic marine reptiles that was most highly adapted to the aquatic environment. The first ichthyosaurs from the upper Lower Triassic (Spathian) already show a suite of unique characters (very large eyes, elongate snout, deeply amphicoelous vertebrae, limb modified to fins) correlated with a fully aquatic existence and probably were unable to leave the water. The key evolutionary innovation was vivipary, giving birth to live young, which is documented by the fossil record since the end of the Anisian. Major evolutionary trends in the locomotor apparatus are the increasing modification of the fin skeleton to a mosaic of bones and the change from anguiliform swimming in the earliest forms to thunniform swimming in the Jurassic and later forms, as evidenced by the shortening of the body and the evolution of a semilunate tail fin. Almost from the beginning, ichthyosaurs had a cosmopolitan distribution which was retained until their extinction in the Cenomanian. Ichthyosaurian diversity is greatest in the Middle Triassic with piscivorous, heterodont, and durophagous forms. Jurassic diversity is greatest in the Liassic, declining to one genus (Platypterygius) in the Cretaceous. Although skull characters indicate that ichthyosaurs were diapsids, their exact position within Diapsida is unclear. A cladistic analysis of the well known genera clarifies relationships within the Ichthyosauria. Most basal areGrippia andUtatsusaurus, followed by the Mixosauridae (Mixosaurus andPhalarodon). The Shastasauridae (Cymbospondylus, Shonisaurus, Besanosaurus) are the most advanced Triassic forms and represent the sistergroup of all post-Triassic ichthyosaurs. These are clearly monophyletic and are termed here the Neoichthyosauria.  相似文献   

5.
Marine reptiles and mammals are phylogenetically so distant from each other that their marine adaptations are rarely compared directly. We reviewed ecophysiological features in extant non-avian marine tetrapods representing 31 marine colonizations to test whether there is a common pattern across higher taxonomic groups, such as mammals and reptiles. Marine adaptations in tetrapods can be roughly divided into aquatic and haline adaptations, each of which seems to follow a sequence of three steps. In combination, these six categories exhibit five steps of marine adaptation that apply across all clades except snakes: Step M1, incipient use of marine resources; Step M2, direct feeding in the saline sea; Step M3, water balance maintenance without terrestrial fresh water; Step M4, minimized terrestrial travel and loss of terrestrial feeding; and Step M5, loss of terrestrial thermoregulation and fur/plumage. Acquisition of viviparity is not included because there is no known case where viviparity evolved after a tetrapod lineage colonized the sea. A similar sequence is found in snakes but with the haline adaptation step (Step M3) lagging behind aquatic adaptation (haline adaptation is Step S5 in snakes), most likely because their unique method of water balance maintenance requires a supply of fresh water. The same constraint may limit the maximum body size of fully marine snakes. Steps M4 and M5 in all taxa except snakes are associated with skeletal adaptations that are mechanistically linked to relevant ecophysiological features, allowing assessment of marine adaptation steps in some fossil marine tetrapods. We identified four fossil clades containing members that reached Step M5 outside of stem whales, pinnipeds, sea cows and sea turtles, namely Eosauropterygia, Ichthyosauromorpha, Mosasauroidea, and Thalattosuchia, while five other clades reached Step M4: Saurosphargidae, Placodontia, Dinocephalosaurus, Desmostylia, and Odontochelys. Clades reaching Steps M4 and M5, both extant and extinct, appear to have higher species diversity than those only reaching Steps M1 to M3, while the total number of clades is higher for the earlier steps. This suggests that marine colonizers only diversified greatly after they minimized their use of terrestrial resources, with many lineages not reaching these advanced steps. Historical patterns suggest that a clade does not advance to Steps M4 and M5 unless these steps are reached early in the evolution of the clade. Intermediate forms before a clade reached Steps M4 and M5 tend to become extinct without leaving extant descendants or fossil evidence. This makes it difficult to reconstruct the evolutionary history of marine adaptation in many clades. Clades that reached Steps M4 and M5 tend to last longer than other marine tetrapod clades, sometimes for more than 100 million years.  相似文献   

6.
We describe an Oligocene newt specimen from western Germany that has gone practically unnoticed in the literature despite having been housed in the Museum für Naturkunde (Berlin) for a century. It is referable to the coeval Chelotriton, but is unusually peramorphic; for many characters it is more peramorphic than all other caudates or even all other lissamphibians. Most noticeable are the position of the jaw joints far caudal to the occiput, the honeycombed sculpture on the maxilla, and the possible presence of a septomaxilla (which would be unique among salamandrids). Referral to a species would require a revision of the genus, but the specimen likely does not belong to the type species. A phylogenetic analysis of nonmolecular characters of Salamandridae, far larger than all predecessors, confirms the referral to Chelotriton. It further loosely associates the Oligocene Archaeotriton and the Miocene Carpathotriton with the extant Lissotriton, though the former may alternatively lie outside Pleurodelinae altogether. The Miocene? I. randeckensis may not belong to the extant Ichthyosaura. The Miocene “Triturus” roehrsi is found neither with the extant Ommatotriton nor with Lissotriton, but inside an Asian/aquatic clade or, when geographic distribution is included as a character, as the sister-group to all other European molgins. The main cause for discrepancies between the results and the molecular consensus is not heterochrony, but adaptations to a life in mountain streams; this is the most likely reason why the Paleocene Koalliella from western Europe forms the sister-group to some or all of the most aquatic extant newts in different analyses. We would like to urge neontologists working on salamandrids to pay renewed attention to the skeleton, not limited to the skull, as a source of diagnostic and phylogenetically informative characters.  相似文献   

7.
The origin of turtles is one of the most contentious issues in systematics with three currently viable hypotheses: turtles as the extant sister to (i) the crocodile–bird clade, (ii) the lizard–tuatara clade, or (iii) Diapsida (a clade composed of (i) and (ii)). We reanalysed a recent dataset that allied turtles with the lizard–tuatara clade and found that the inclusion of the stem turtle Proganochelys quenstedti and the ‘parareptile’ Eunotosaurus africanus results in a single overriding morphological signal, with turtles outside Diapsida. This result reflects the importance of transitional fossils when long branches separate crown clades, and highlights unexplored issues such as the role of topological congruence when using fossils to calibrate molecular clocks.  相似文献   

8.
Megachirella wachtleri Renesto et Posenato, 2003, a well preserved partial reptile skeleton from the Middle Triassic of the Dolomites (N. Italy), was originally considered a lepidosauromorph, but no phylogenetic analysis was carried out. Consequently, the taxon was overlooked in later phylogenetic analyses of the Diapsida. Here, the holotype and only known specimen of M. wachtleri is redescribed, allowing an investigation of its phylogenetic relationships. Phylogenetic analyses confirm that Megachirella is a lepidosauromorph close to the crown group lepidosaurs (Squamata + Rhynchocephalia). Megachirella enhances our knowledge of the series of morphological modifications that led to the origin of the Lepidosauria, the most diverse clade of extant reptiles.  相似文献   

9.
The structure of caudal neural spines of protoceratopoids displays adaptation for aquatic and terrestrial mode of life. The increasing height of caudal neural spines in the series Leptoceratops, Udanoceratops, Protoceratops, Bagaceratops is connected with the extent of adaptation for swimming and changes in inclination of neural spines are connected with the mechanical balance of the lever. Thus, the anterior caudal vertebrae (1cd–15cd) of Protoceratops and Bagaceratops show an anticliny, which promotes extension (rise) of a heavy tail in terrestrial conditions. In the middle part of the tail (16cd–23cd), with the greatest height of neural spines, a decrease in width and increase in thickness counteract transverse loads accompanying movements on land. At the same time, the supraspinal ligament prevents divergence of neural spines caused by curvature of the tail as it is raised above the ground.  相似文献   

10.
Sander PM  Chen X  Cheng L  Wang X 《PloS one》2011,6(5):e19480

Background

Ichthyosaurs were an important group of Mesozoic marine reptiles and existed from the Early Triassic to the early Late Cretaceous. Despite a great diversity in body shapes and feeding adaptations, all share greatly enlarged eyes, an elongated rostrum with numerous conical teeth, and a streamlined body.

Methodology/Principal Findings

Based on new material from China and the restudy of Shastasaurus pacificus, we here reinterpret the classical large-bodied Late Triassic ichthyosaur genus Shastasaurus to differ greatly from the standard ichthyosaurian body plan, indicating much greater morphological diversity and range of feeding adaptations in ichthyosaurs than previously recognized. Phylogenetic analysis indicates a monophyletic clade consisting of the giant Shonisaurus sikanniensis, Guanlingsaurus liangae, and Shastasaurus pacificus to which the genus name Shastasaurus is applied. Shastasaurus liangae comb. nov. is from the Late Triassic (Carnian) Xiaowa Formation of Guizhou Province, southwestern China. The species combines a diminutive head with an entirely toothless and greatly reduced snout. The species also has by far the highest vertebral count among ichthyosaurs (86 presacral vertebrae and >110 caudal vertebrae), a count that is also very high for tetrapods in general. A reduced toothless snout and a diminutive head is also apparently present in the giant S. sikanniensis and presumably in S. pacificus.

Conclusions/Significance

In analogy to many modern odontocetes, Shastasaurus is interpreted as a specialized suction feeder on unshelled cephalopods and fish, suggesting a unique but widespread Late Triassic diversification of toothless, suction-feeding ichthyosaurs. Suction feeding has not been hypothesized for any of the other diverse marine reptiles of the Mesozoic before, but in Shastasaurus may be linked to the Late Triassic minimum in atmospheric oxygen.  相似文献   

11.
Clitellata (earthworms, leeches, and allies) is a clade of segmented annelid worms that comprise more than 5000 species found worldwide in many aquatic and terrestrial habitats. According to current views, the first clitellates were either aquatic (marine or freshwater) or terrestrial. To address this question further, we assessed the phylogenetic relationships among clitellates using parsimony, maximum likelihood and Bayesian analyses of 175 annelid 18S ribosomal DNA sequences. We then defined two ecological characters (Habitat and Aquatic‐environment preferences) and mapped those characters on the trees from the three analyses, using parsimony character‐state reconstruction (i.e. Fitch optimization). We accommodated phylogenetic uncertainty in the character mapping by reconstructing character evolution on all the trees resulting from parsimony and maximum likelihood bootstrap analyses and, in the Bayesian inference, on the trees sampled using the Markov chain Monte Carlo algorithm. Our analyses revealed that an ‘aquatic’ ancestral state for clitellates is a robust result. By using alterations of coding characters and constrained analyses, we also demonstrated that the hypothesis for a terrestrial origin of clitellates is not supported. Our analyses also suggest that the most recent ancestor of clitellates originated from a freshwater environment. However, we stress the importance of adding sequences of some rare marine taxa to more rigorously assess the freshwater origin of Clitellata. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 447–464.  相似文献   

12.
The physical nature of water and the environment it presents to an organism have long been recognized as important constraints on aquatic adaptation and evolution. Little is known about the dermal cover of mosasauroids (a group of secondarily aquatic reptiles that occupied a wide array of predatory niches in the Cretaceous marine ecosystems 92–65 Myr ago), a lack of information that has hindered inferences about the nature and level of their aquatic adaptations. A newly discovered Plotosaurus skeleton with integument preserved in three dimensions represents not only the first documented squamation in a mosasaurine mosasaur but also the first record of skin in an advanced member of the Mosasauroidea. The dermal cover comprises keeled and possibly osteoderm-reinforced scales that presumably contributed to an anterior–posterior channelling of the water flow and a reduction of microturbulent burst activities along the surface of the skin. Thus, hydrodynamic requirements of life in the water might have influenced the evolution of multiple-keeled body scales in advanced mosasauroids.  相似文献   

13.
Exceptional occurrences have fundamental interest in evolution relevant to understanding adaptations and origins. Monogeneans primarily infect aquatic lower vertebrates, i.e. fish, amphibians and chelonian reptiles, but there is a single instance of colonisation of a mammal: Oculotrema hippopotami Stunkard, 1924 infecting the eye of Hippopotamus amphibius Linnaeus. Its combination of systematic characters is amongst the most diverse in the Polystomatidae Gamble, 1896 and relationships are obscure. This study emphasises the primary significance of two features: the reinforcement of haptoral suckers with an internal skeleton and the pattern of ciliated cells on the oncomiracidium, especially the presence of conjoined cells. Closest relationships are with polystomatids infecting chelonians, specifically species of Polystomoides Ward, 1917 from the oral cavity/pharynx, or more likely (but with currently incomplete evidence) species of Neopolystoma Price, 1939 from the eye. Morphological characters of polystomoidines, all of which infect chelonians, appear to have remained relatively stable since at least the Jurassic (from zoogeographical evidence), but the highly derived characters of species of Oculotrema may have evolved during the comparatively short period (16 million years) since the Miocene origin of Hippopotaminae Gray. However, the initial host switch may plausibly have been to hippo ancestors, the anthracotheres, with similar semi-aquatic ecology and an Eocene origin (41 million years ago). Over the same time-scale, the oncomiracidial cell pattern remained closely comparable with that of presumed ancestors, emphasising its value in phylogenetic analyses.  相似文献   

14.
The ‘tree lobsters’ are an enigmatic group of robust, ground-dwelling stick insects (order Phasmatodea) from the subfamily Eurycanthinae, distributed in New Guinea, New Caledonia and associated islands. Its most famous member is the Lord Howe Island stick insect Dryococelus australis (Montrouzier), which was believed to have become extinct but was rediscovered in 2001 and is considered to be one of the rarest insects in the world. To resolve the evolutionary position of Dryococelus, we constructed a phylogeny from approximately 2.4 kb of mitochondrial and nuclear sequence data from representatives of all major phasmatodean lineages. Our data placed Dryococelus and the New Caledonian tree lobsters outside the New Guinean Eurycanthinae as members of an unrelated Australasian stick insect clade, the Lanceocercata. These results suggest a convergent origin of the ‘tree lobster’ body form. Our reanalysis of tree lobster characters provides additional support for our hypothesis of convergent evolution. We conclude that the phenotypic traits leading to the traditional classification are convergent adaptations to ground-living behaviour. Our molecular dating analyses indicate an ancient divergence (more than 22 Myr ago) between Dryococelus and its Australian relatives. Hence, Dryococelus represents a long-standing separate evolutionary lineage within the stick insects and must be regarded as a key taxon to protect with respect to phasmatodean diversity.  相似文献   

15.
We performed molecular phylogenetic analyses of four morphotypes of the benthic foraminiferal genus Bulimina (B. aculeata, B. marginata f. marginata, B. marginata f. denudata, and B. elongata) based on sequences of internal transcribed spacer region of ribosomal DNA (ITS rDNA). Six genetically distinct phylotypes were revealed by our phylogenetic analyses. The six phylotypes basically correspond to the fundamental morphotypes: clades A + B (B. aculeata); clade C (B. elongata); clade D (B. marginata f. denudata); clade E (B. marginata f. marginata genotype 1); and clade F (B. marginata f. marginata genotype 2). All six phylotypes are well distinguished, except phylotype B, which shows only little sequence divergence compared to clade A, possibly indicating that genetic differentiation is in progress. Morphological characters including the direction, placement, and shape of spines, the angle of undercutting of chamber periphery, and the roundness of the chambers were stable among specimens of each clade. In contrast, the length and density of spines, and chamber size, were variable within each clade. These intermediate morphological characters may reflect ecophenotypic variation. Our study clearly shows that the examined B. acuelata, elongata, and denudata morphospecies are genetically separated and that B. marginata is a species complex comprising several genotypes. A novel phylotype represent different morphotype compare to B. marginata f. marginata that can be distinguished based on differences of chamber angularity, the direction, placement, and shape of spines, and test dimensions.  相似文献   

16.
17.
Metriorhynchoid crocodylians represent the pinnacle of marine specialization within Archosauria. Not only were they a major component of the Middle Jurassic–Early Cretaceous marine ecosystems, but they provide further examples that extinct crocodilians did not all resemble their modern extant relatives. Here, we use a varied toolkit of techniques, including phylogenetic reconstruction, geometric morphometrics, diversity counts, discrete character disparity analysis, and biomechanical finite‐element analysis (FEA), to examine the macroevolutionary history of this clade. All analyses demonstrate that this clade became more divergent, in terms of biodiversity, form, and function, up until the Jurassic–Cretaceous boundary, after which there is no evidence for recovery or further radiations. A clear evolutionary trend towards hypercarnivory in Dakosaurus is supported by phylogenetic character optimization, morphometrics, and FEA, which also support specialized piscivory within Rhacheosaurus and Cricosaurus. Within Metriorhynchoidea, there is a consistent trend towards increasing marine specialization, with the hypermarine Cricosaurus exhibiting numerous convergences with other Mesozoic marine reptiles (e.g. loss of the deltopectoral crest and retracted external nares). In addition, biomechanics, morphometrics, and character‐disparity analyses consistently distinguish the two newly erected metriorhynchid subfamilies. This study illustrates that together with phylogeny, quantitative assessment of diversity, form, and function help elucidate the macroevolutionary pattern of fossil clades. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 801–859.  相似文献   

18.
During a parasitological survey, Myxidium-like spores were identified in the gall bladders of marine fishes from Australian waters. This paper describes four novel species of Ellipsomyxa Køie, 2003, three novel species of Myxidium Bütschli, 1882 and six novel species of Zschokkella Auerbach, 1910 from teleosts from Australian waters using a combination of morphological, biological and molecular characters. Phylogenetic analyses showed a monophyletic relationship of all Ellipsomyxa spp. sequences with Sigmomyxa sphaerica (Thélohan, 1895) and Myxidium queenslandicus Gunter & Adlard, 2008 as sister species to the clade. The validity of genus Sigmomyxa Karlsbakk & Køie, 2012 is discussed. In phylogenetic analyses, the novel species of Myxidium fell within the ‘marine’ clade of Fiala (2006). However, the novel species of Zschokkella fell within the ‘freshwater’ clade of Fiala (2006) and formed a distinct clade with all other sequences of Zschokkella spp. from the gall bladder of marine fish and a sequence of a species of Myxobolus Bütschli 1882, also from the gall bladder of a marine fish. This is the second distinct marine lineage to emerge within the freshwater clade.  相似文献   

19.
20.
Kånneby, T., Todaro, M. A., Jondelius, U. (2012). Phylogeny of Chaetonotidae and other Paucitubulatina (Gastrotricha: Chaetonotida) and the colonization of aquatic ecosystems. —Zoologica Scripta, 42, 88–105. Chaetonotidae is the largest family within Gastrotricha with almost 400 nominal species represented in both freshwater and marine habitats. The group is probably non‐monophyletic and suffers from a troubled taxonomy. Current classification is to a great extent based on shape and distribution of cuticular structures, characters that are highly variable. We present the most densely sampled molecular study so far where 17 of the 31 genera belonging to Chaetonotida are represented. Bayesian and maximum likelihood approaches based on 18S rDNA, 28S rDNA and COI mtDNA are used to reconstruct relationships within Chaetonotidae. The use of cuticular structures for supra‐specific classification within the group is evaluated and the question of dispersal between marine and freshwater habitats is addressed. Moreover, the subgeneric classification of Chaetonotus is tested in a phylogenetic context. Our results show high support for a clade containing Dasydytidae nested within Chaetonotidae. Within this clade, only three genera are monophyletic following current classification. Genera containing both marine and freshwater species never form monophyletic clades and group with other species according to habitat. Marine members of Aspidiophorus appear to be the sister group of all other Chaetonotidae and Dasydytidae, indicating a marine origin of the clade. Halichaetonotus and marine Heterolepidoderma form a monophyletic group in a sister group relationship to freshwater species, pointing towards a secondary invasion of marine environments of these taxa. Our study highlights the problems of current classification based on cuticular structures, characters that show homoplasy for deeper relationships.  相似文献   

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