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1.
Mass accumulations of belemnite rostra ('belemnite battlefields') are common in Mesozoic sediments, and accumulations of belemnoids are also known from older rocks. Many Recent teuthid species suffer mortality immediately after spawning, and some authors have suggested that belemnite accumulations record a similar phenomenon. Conversely, it is clear that many belemnite battlefields actually formed in an environment of net sediment loss, with current alignment and winnowing. A proposed model for the production of belemnite battlefields involves five possible pathways: post-spawning mortality, catastrophic mass mortality, predation concentration, stratigraphical condensation, and resedimentation. Although accumulations that have not been reworked are rare, it is possible to recognise the generating agents of belemnite accumulations through their intrinsic features. A genetic classification of belemnite battlefields, based on the identified pathways, is tested through field examples in Britain and Antarctica, as well as through published examples. □ Belemnites, fossil accumulations, palaeoecology, Mesozoic, Antarctica, Yorkshire.  相似文献   

2.
Trace element contents and stable isotopic composition of Middle Campanian marl-limestone rhythmites and belemnite rostra of Belemnitella mucronata were investigated. High strontium and low iron as well as manganese and magnesium contents of belemnite calcite and bulk rock samples suggest no diagenetic overprint. However, the orange-coloured cathodoluminescence of coccolith-rich sediments indicates diagenetic cementation and/or recrystallization. The non-luminescent belemnite rostra reveal an extraordinary preservation of the microstructures that is interpreted to have been favoured by a silification of the outer rim of the belemnite rostra. Carbon isotope ratios of the coccolith limestones and belemnite rostra are comparable, with higher δ13C variations observed for belemnite calcite. The 1.5-2‰ depletion in δ18O of the marl-limestone rhythmites relative to belemnite calcite is explained by diagenetic alteration of the sediments. Palaeotemperatures, calculated from the δ18O values of the well-preserved belemnite rostra, are around 12.5 ± 2 °C and suggest rather low sea-surface temperatures for the Middle Campanian epicontinental sea of north Germany assuming a water depth of less than 100 m.  相似文献   

3.
The late Kimmeridgian Nusplingen Plattenkalk (palaeolatitude ~30°N) is one of the well‐known European Plattenkalk konservatlagerstätten. The laminated limestones of this lagerstätte have been deposited in a shallow lagoon, surrounded by sponge‐microbial mounds, some of which formed small islands. Plattenkalk sediments are often thought to have been deposited below a halocline, which was induced by high evaporation rates. By measuring the stable isotope composition of belemnite rostra (n = 27) of the species Hibolithes semisulcatus, the depositional environment of the Nusplingen Plattenkalk has been investigated. Cathodoluminescence‐microscopy and trace element analysis have been applied to check for diagenetic alteration of the studied rostra. The mean δ13C of the well‐preserved rostra is +0.03 ± 0.27‰, the mean δ18O ?0.68 ± 0.3‰. A narrow range of stable isotope data acquired from an accumulation of rostra, regurgitated by a fish or reptile, supports the notion that belemnite calcite reflects environmental conditions. The palaeontological and sedimentological findings suggest that H. semisulcatus was autochthonous to the Nusplingen Plattenkalk. Anoxic conditions prevailed in the sediments and temporarily in the water‐column above the sediment/bottom water interface. A nektobenthic life style can be excluded for the Nusplingen belemnites; a demersal life style seems unlikely. Comparison with δ18Obel data from other latitudes indicates that a latitudinal gradient played a role in the δ18O composition of seawater. A pelagic, inner shelf setting is therefore suggested for the Plattenkalk, an interpretation that is supported by palaeontological evidence. It is here proposed that the formation of the Nusplingen Plattenkalk was not caused by salinity stratification. It is further suggested that the belemnites were not restricted to deep‐water settings, but also occurred in shallow waters and higher in the water‐column.  相似文献   

4.
Oxygen and carbon isotope data of well-preserved belemnite rostra and ammonite shells are presented from the Callovian–Oxfordian boundary (uppermost Lamberti to lowermost Cordatum zones) of the Dubki section near Saratov in the Russian Platform. Palaeotemperatures calculated for nektobenthic belemnites (averages of 5 °C and 8 °C for cylindroteuthids and belemnopseids, respectively) show the presence of cold bottom waters in the central part of the Middle Russian Sea during the studied interval. Palaeotemperatures calculated for ammonites, which are assumed to have lived in near-surface waters, are considerably higher (average 13 °C). The presented data show a vertical thermal gradient in the Middle Russian Sea. The belemnite oxygen isotope record and the relative abundances of ammonite families in the Dubki section do not correlate with each other probably as a result of different depth habitats of ammonites and belemnites. A review of literature isotope data shows the climatic zonation in European seas at the Middle–Late Jurassic transition. Despite the flux of cold polar waters to the Middle Russian Sea and the area of Scotland there is no evidence for glaciation at the Middle–Late Jurassic transition. Changes in water circulation during a sea-level highstand were likely a source of spreads of cold bottom waters and cardioceratid ammonite fauna in this time period.The belemnite isotope record of the Callovian–Oxfordian boundary in the Russian Platform is characterized by significant scatter of δ13C values. No temporal carbon isotope trend is observed. The δ13C values of Russian belemnite rostra average 2.6‰ VPDB being 1 to 2‰ higher than the values of coeval Lower Oxfordian belemnites from the area of the Submediterranean ammonite province. Higher (than Submediterranean) δ13C values of Russian belemnite rostra are likely related to high biologic productivity and/or high organic matter burial in semi-isolated Boreal–Subboreal marine basins.  相似文献   

5.
The higher rank systematics of coleoid cephalopods are primarily based on the total number of arms. Completely preserved arm crows are, therefore, the most important character complex by which fossil representatives can be assigned to higher taxa. Whereas belemnoids—similar to modern Decabrachia—are well-known to have ten hooklet-bearing arms, recent studies have shown that Middle-Late Jurassic and Late Cretaceous gladius-bearing coleoids belong to the Vamyropoda, owing to having only eight hooklet-free arms. The arm number of Early Jurassic gladius-bearing coleoids was unclear for a long time. This paper, however, reports the first unequivocally complete arm crown from the Early Jurassic (Toarcian). It consists of eight strongly phosphatised arms of equal shape. Affinity with decabrachiate coleoids is therefore unlikely. The mantle sack of the specimen, which was identified as Loligosepia aalensis (Schübler in Zieten, 1832), has, furthermore, an unusual arrangement of numerous angular pieces. The irregular scattering of the mantle fragments was most probably caused by a scavenger. Finally, hooklets in the food residues in the posterior mantle indicate that Loligosepia preyed upon belemnites.  相似文献   

6.
A morphological comparison of shell‐muscle contacts in coleoid cephalopods mainly from the Early Jurassic (Toarcian) Posidonia Shales of Holzmaden (Germany), the Middle Jurassic (Callovian) Oxford Clay of Christian Malford (UK), Late Jurassic (Kimmeridgian‐Tithonian) plattenkalks of Solnhofen (Germany), and the Late Cretaceous (Cenomanian) of Hâdjoula and Hâkel (Lebanon) provides new and meaningful insights into their locomotion systems. The study shows that both pro‐ostracum‐ and gladius‐bearing coleoids are typified by a marginal mantle attachment and by distinctly separated fins, which usually insert (indirectly via the shell sac and basal fin cartilages) to posterior shell parts. While absent in gladius‐bearing forms, mantle‐locking cartilages might have existed already in pro‐ostracum‐bearing belemnoids. Similar to ectocochleate ancestors, funnel‐ and cephalic retractors are generally attached to the internal (ventral) shell surface. A comparison of Mesozoic and Recent gladius‐bearing coleoids shows that the locomotion system (most significantly the dorsal mantle configuration, and the presence of nuchal‐ and funnel‐locking cartilages) is fundamentally different. This does not support the concept of ‘fossil teuthids’, but suggests, owing to similarities with Recent Vampyroteuthis, placement of Mesozoic gladius‐bearing coleoids within the Octobrachia (Octopoda + Vampyromorpha). Classification of Mesozoic gladius‐bearing coleoids as octobrachians implies that: (1) unambiguous teuthids are still unknown in the fossil record and (2) the similarity between Recent and some fossil gladiuses represents a matter of homoplasy.  相似文献   

7.
A ten armed fossil from the Pennsylvanian Francis Creek Shale and belonging to the famous Mazon Creek biota is described as a cephalopod. Apart from a superficial resemblance to the coleoids, there is little basis for an assessment of the animal's affinities. A mineralized Bask shaped structure preserved in calcite and seen in both part and counterpart in fragmentary form may represent the fossilized remains of an internal organ such as the stomach or ink sac A small, dark circular spot positioned anterior to the appendages could be the animal's eye although mere is no internal structure visible. The extremely rare occurrence of Palaeozoic cephalopods with soft parts has so far been Ended to the Francis Creek Shale and the Devonian Hunsrückschiefer of Germany. The Mazon Creek biota includes many fossils which show a remarkable standard of soft part preservation which in some cases includes the fossilization of eye spots and stomach traces. Previously deserted Mazon Creek cephalopods have included the tentacular impression of a squid complete with arm hooks, and the radulae and shells of both coleoids and nautiloids. The specimen described here is the first from the Palaeozoic to show any hint of organ preservation.  相似文献   

8.
E. F. v. Schlotheim described 46 species of Jurassic and Cretaceous belemnites in 1813,1820, and 1832. Their type specimens are in part housed at the Humboldt University of Berlin. The holotype ofBelemnites irregularis is here refigured. Lectotypes forB. acuarius, B. canaliculatus, B. clavatus, B. giganteus, andB. tripartitus are designated and figured for the first time. The lectotype ofBelemnites giganteus is designated to be the neotype ofBelemnites suevicus Klein 1773. 36 species are nomina nuda and nomina dubia, because none of their type specimens could be detected, among them the invalidB. paxillosus v. Schlotheim 1820, which is, according to its lectotype designated herein, likeBelemnites giganteus, a junior synonym ofMegateuthis suevica (Klein 1773). An original type specimenof Belemnites mucronatusv. Schlotheim 1813, for which a neotype had already been ruled by the ICZN in 1985, was re-discovered in thev. Schlotheim collection in 1996.v. Schlotheim’s belemnites were up to now designated to be type species of 12 genera.  相似文献   

9.
Based on well‐preserved belemnites, the ontogenetic trajectories of septal spacing between succeeding chambers were analysed. In the examined species (Passaloteuthis laevigata, Parapassaloteuthis zieteni and Pseudohasitites longiformis) that come from Buttenheim, Germany, and Lixhausen, France, the ontogenetic trajectories of septal spacing follow exponentially increasing trends with no decreasing phase of septal crowding during the earliest ontogenetic stage. The absence of a decreasing trend at the earliest ontogenetic stage is a unique character in contrast with those in modern cuttlefish and ancient and modern nautiloids, in which the decreasing trends are related to hatching events. These ontogenetic septal spacing trends suggest that the belemnite hatchlings had only a protoconch with no chamber. These belemnite hatchlings with no chamber and therefore small embryonic shell diameter are similar to those of ammonoids. Significant difference in a statistical test that compared the protoconch size between the two localities, might suggest that there was limited transportation at the embryonic stage, although it could also just indicate differences in regional environmental conditions, age and/or degree of time averaging which might differ between the examined taxa.  相似文献   

10.
The gudgeons (subfamily Gobioninae) are a group of cyprinid fishes primarily distributed in East Asia. However, studies on their origins and divergence are scarce. Here the whole mitochondrial genome sequences of 27 gudgeon species (including one newly determined), 22 other cyprinid species, and two non‐cyprinids as outgroups are applied to infer the evolution of the gudgeons. Based on Bayesian and maximum likelihood phylogenetic analyses, the gudgeons were determined to be a monophyletic group which can be further subdivided into four monophyletic clades with strong supports. The divergence times of the gudgeons were estimated using a relaxed molecular clock method; the results indicate that these fishes originated in the early Paleocene (approx. 63.5 Mya) and that the basal Hemibarbus group diverged from the other gudgeon fishes (approx. 58.3 Mya). As an independent group the Coreius began to diverge from the remaining two groups (approx. 54.6 Mya); the most derived two groups diverged from each other (approx. 53.6 Mya). The divergences of the four gudgeon groups were within a relatively short time frame (approx. 58–53 Mya). Based on the reconstruction of evolutionary trends of gudgeon habitat, evidence is provided that supports the origin and differentiation of this fauna as being associated with some special paleo‐environmental events occurring from the early Paleocene to the Pliocene. The study represents comprehensive molecular dating and character evolution analyses of the gudgeons, and providing a valuable framework for future research in the evolution of the Gobioninae fishes.  相似文献   

11.
Klug, C., Schweigert, G., Fuchs, D. & Dietl, G. 2009: First record of a belemnite preserved with beaks, arms and ink sac from the Nusplingen Lithographic Limestone (Kimmeridgian, SW Germany). Lethaia, 10.1111/j.1502‐3931.2009.00203.x A recent discovery of an unusually preserved belemnite from Nusplingen comprises the extraordinarily rare remains of beaks and nearly in situ arm hooks, as well as the ink sac and an incomplete phragmocone. So far, Hibolithes semisulcatus ( Münster, 1830 ) is the only belemnite known from the Nusplingen Lithographic Limestone (Upper Jurassic, Late Kimmeridgian, Beckeri Zone, Ulmense Subzone; SW Germany) that has the same phragmocone shape and size, and thus we assign the new specimen to this taxon. The rostrum was probably lost due to a lethal predation attempt in which the prey was killed but not entirely eaten. For the first time a specimen reveals details of the belemnite beak morphology, which we compare with the beaks of other Jurassic coleoids. This specimen presently represents the only known rostrum‐bearing belemnite of post‐Toarcian age with preserved non‐mineralized body parts. With the new discovery, Nusplingen now represents the only locality which has yielded complete beak apparatuses from all major Jurassic cephalopod groups. □Beaks, Belemnitida, Coleoidea, Germany, Late Jurassic, morphology, taphonomy.  相似文献   

12.
Twenty species belonging to fifteen genera of cyclostome and cheilostome bryozoans encrusting belemnite rostra are described from the late Campanian to Maastrichtian of the Aktolagay Plateau, in western Kazakhstan. Due to the moderate to poor preservation of the material, only four cheilostome species are identified down to the species level: Wilbertopora? besoktiensis (Voigt, 1967), ‘Aechmellinastenostoma Voigt, 1930, and two new species, ‘Aechmellinaviskovae and Cheethamia aktolagayensis. All remaining species are left in open nomenclature. Type material of Wilbertopora? besoktiensis from the early Maastrichtian of the Mangyshlak Peninsula in Kazakhstan, has been re-examined. Palaeobiogeographical and implications are discussed. Cheilostomes slightly dominated over cyclostomes in the Aktolagay Plateau fauna encrusting on belemnites in terms of diversity. The dominant colony forms observed were spots and sheets.  相似文献   

13.
Early Jurassic belemnites are of particular interest to the study of the evolution of skeletal morphology in Lower Carboniferous to the uppermost Cretaceous belemnoids, because they signal the beginning of a global Jurassic–Cretaceous expansion and diversification of belemnitids. We investigated potentially relevant, to this evolutionary pattern, shell features of Sinemurian–Bajocian Nannobelus, Parapassaloteuthis, Holcobelus and Pachybelemnopsis from the Paris Basin. Our analysis of morphological, ultrastructural and chemical traits of the earliest ontogenetic stages of the shell suggests that modified embryonic shell structure of Early–Middle Jurassic belemnites was a factor in their expansion and colonization of the pelagic zone and resulted in remarkable diversification of belemnites. Innovative traits of the embryonic shell of Sinemurian–Bajocian belemnites include: (1) an inorganic–organic primordial rostrum encapsulating the protoconch and the phragmocone, its non‐biomineralized component, possibly chitin, is herein detected for the first time; (2) an organic rich closing membrane which was under formation. It was yet perforated and possessed a foramen; and (3) an organic rich pro‐ostracum earlier documented in an embryonic shell of Pliensbachian Passaloteuthis. The inorganic–organic primordial rostrum tightly coating the protoconch and phragmocone supposedly enhanced protection, without increase in shell weight, of the Early Jurassic belemnites against explosion in deep‐water environment. This may have increased the depth and temperature ranges of hatching eggs, accelerated the adaptation of hatchlings to a nektonic mode of life and promoted increasing diversity of belemnoids. This study supports the hypothesis that belemnite hatchlings were ‘a miniature of the adults’.  相似文献   

14.
Summary New petrographic and isotopic data from inoceramid bivalve shells and belemnite rostra from the lower Campanian and belemnite rostra from the mid-upper Maastrichtian of the Marambio Group, James Ross Basin, Antarctica are presented. Most of the inoceramid data were processed from shell fragments of the large formAntarcticeramus rabotensis (Crame and Luther) at the stratigraphic level marking the extinction of the inoceramids in the James Ross Basin (uppermost early Campanian-basal late Campanian). Standard transmitted light microscopy and cathodoluminescence (CL) studies in thin sections ofA. rabotensis show clear evidence of environmental stress, which is reflected as marked growth breaks in the shell banding of this large inoceramid bivalve. At Redonda Point, CL and the mean oxygen isotopic value (δ18O=-3.11‰ 3 (PDB); n=11; t°=25.4°C) indicate a varied degree of diagenetic modification, but without any evidence of neomorphism along the prismatic microstructures. Early Campanian belemnite rostra are much less diagenetically modified (at the Brandy Bay section; and the Santa Marta section; δ18O=-0.50‰ (PDB); n=5; t°=14.0°C and 3 δ18O=-0.94‰ (PDB); n=21; t°=15.8°C) and are non luminescent 3 except for localized, organic-rich bands. The mean oxygen isotopic value for mid-late Maastrichtian belemnite rostra (at the Seymour Island section; δ18O=-0.11‰ 3 (PDB); n=5; t°=12.5°C) indicates a substantial drop in the sea-water paleotemperature, suggesting a causal relationship between the early extinction of the inoceramid bivalves in high latitudes of the Southern Hemisphere and the falling sea-water temperature.  相似文献   

15.
Nino Mariotti 《Geobios》2003,36(5):603-623
About three hundred belemnite rostra were collected from lower Kimmeridgian beds of a structural high sequence cropping out at Mt. Nerone (Central Apennines, Pesaro Province, Italy). The belemnite fauna is composed mainly of new species. Nine species were recognised, ascribed to five genera, that include Hibolithes semisulcatus M?NSTER, 1830; H. pignattii nov. sp.; Acutibelus sp. cf. acuariformis RIEGRAF, 1981; Belemnopsis neronensis nov. sp., Duvalia matteuccii nov. sp., D. nicosiai nov. sp., D. pallinii nov. sp., D. raymondi nov. sp. and Rhopaloteuthis massimoi nov. sp.; moreover a single specimen is treated in open nomenclature as Belemnopseidae incertae sedis. The stratigraphic and palaeobiogeographic significance of the new fauna is discussed. The taphonomy of the belemnite-rich level is described, with reference to borings found on the belemnite rostra.  相似文献   

16.
1. Fish assemblages and habitats were sampled annually at fixed sites in three tributaries of the Gila River catchment over a 21‐year span that included prolonged low‐ and high‐flow periods. Model selection was used to evaluate responses of seven native fishes with variable ecological traits (four small‐bodied cyprinids, one large‐bodied cyprinid, and two large‐bodied catostomids) to mean annual discharge and predacious non‐native fishes across the three sites. We also compared habitat use and overlap of native and non‐native fishes to identify potential for negative interactions among species. 2. Assemblage structure (species abundance and richness) and recruitment of native species was strongly and primarily affected by mean annual discharge and secondarily by location and densities of non‐native predators (mainly the centrarchid Micropterus dolomieui). 3. Densities of age‐0 catostomids and small‐bodied cyprinids were positively associated with discharge, and this pattern was strongest in the tributary with the lowest densities of non‐native predators. Absence or extreme low abundance of natives during low‐flow years was most pronounced at the sites where non‐native predators were comparatively common. Densities of adults of large‐bodied native species also varied by site, but often were positively associated with densities of non‐native predators. 4. Spatially variable responses of native fish assemblages indicated that the persistence of native fishes could be jeopardized if key habitats were lost or flow regimes unnaturally altered, particularly during low‐flow conditions when recruitment of native fishes is low and predation by non‐natives is high. Large‐bodied species may be less vulnerable to multiple years of poor conditions because adults are able to avoid predation by non‐natives and thus can rely on occasional high discharge years for successful recruitment. 5. As in other arid‐land streams, native fish assemblages of the Gila River Basin continue to decline. Our results indicate that conservation requires specific knowledge and consideration of physical influences as well as life‐history attributes of native and non‐native fishes.  相似文献   

17.
Polarization sensitivity (PS), which arises from the orthogonal arrangement of microvilli in the retina, has long been known in shallow-water cephalopods. Micrographs presented herein signify that some deep-water cephalopods may also possess PS. Precise measurements of the angles of microvilli in the retinas of shallow-water octopus, squid and cuttlefish revealed neuroanatomical differences that may explain variation in the limits of polarization angular discrimination in different species and habitats. A question yet unanswered is whether cephalopods can discriminate between polarization and intensity. Recent behavioral experimentation has illustrated that one clear function of PS is enhanced predation: it enables better detection of transparent, opaque, or silvery-reflecting prey. The use of PS for navigation in cephalopods is still controversial, and our recent experiment on squids failed to support this notion. It is possible that cephalopods show polarization patterning produced in their skin as a mode of communication that cannot be detected by polarization-insensitive predators such as many fishes and marine mammals.  相似文献   

18.
A biostratigraphic and systematic study based on belemnites collected along with ammonites was performed on four sections in the Subalpine Basin (SE France): Lac du Castillon and La Baume (Castellane area), Galabrun and Grand Lara (Gap area). The specimens, originating from hemi-pelagic marl-limestone alternations in the lower part of the “Calcaires à Zoophycos” Formation, are dated from Middle Aalenian (Murchisonae Zone) to Lower Bajocian (Humphriesianum Zone). Five belemnite taxa (Megateuthis elliptica, Holcobelus munieri, Htrauthi, Pachybelemnopsis roettingensis, Hibolithes sp.) have been identified, and two more taxa are reported in an open nomenclature (Belemnitida incertae sedis sp. 1 and sp. 2). The biostratigraphic range of the belemnite fauna is established. The new findings contribute to a more detailed understanding of the paleobiogeography of holcobelid belemnites that flourished at the northern margin of the Tethys Ocean and formed a distinct sub-Mediterranean fauna. The association herein described is similar to the fauna of the Calabro-Peloritani Arc (Calabria, Italy), a further hint for the supposed paleogeographic position of the latter during the Middle Jurassic.  相似文献   

19.
Our understanding of how environmental change in the Southern Ocean will affect marine diversity, habitats and distribution remain limited. The habitats and distributions of Southern Ocean cephalopods are generally poorly understood, and yet such knowledge is necessary for research and conservation management purposes, as well as for assessing the potential impacts of environmental change. We used net-catch data to develop habitat suitability models for 15 of the most common cephalopods in the Southern Ocean. Using modeled habitat suitability, we assessed favorable areas for each species and examined the relationships between species distribution and environmental parameters. The results compared favorably with the known ecology of these species and with spatial patterns from diet studies of squid predators. The individual habitat suitability models were overlaid to generate a “hotspot” index of species richness, which showed higher numbers of squid species associated with various fronts of the Antarctic circumpolar current. Finally, we reviewed the overall distribution of these species and their importance in the diet of Southern Ocean predators. There is a need for further studies to explore the potential impacts of future climate change on Southern Ocean squid.  相似文献   

20.
Almany GR 《Oecologia》2004,141(1):105-113
Greater structural complexity is often associated with greater abundance and diversity, perhaps because high complexity habitats reduce predation and competition. Using 16 spatially isolated live-coral reefs in the Bahamas, I examined how abundance of juvenile (recruit) and adult (non-recruit) fishes was affected by two factors: (1) structural habitat complexity and (2) the presence of predators and interference competitors. Manipulating the abundance of low and high complexity corals created two levels of habitat complexity, which was cross-factored with the presence or absence of resident predators (sea basses and moray eels) plus interference competitors (territorial damselfishes). Over 60 days, predators and competitors greatly reduced recruit abundance regardless of habitat complexity, but did not affect adult abundance. In contrast, increased habitat complexity had a strong positive effect on adult abundance and a weak positive effect on recruit abundance. Differential responses of recruits and adults may be related to the differential effects of habitat complexity on their primary predators. Sedentary recruits are likely most preyed upon by small resident predators that ambush prey, while larger adult fishes that forage widely and use reefs primarily for shelter are likely most preyed upon by large transient predators that chase prey. Increased habitat complexity may have inhibited foraging by transient predators but not resident predators. Results demonstrate the importance of habitat complexity to community dynamics, which is of concern given the accelerated degradation of habitats worldwide.  相似文献   

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