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1.
It is generally assumed that species with low population sizes have lower genetic diversities than larger populations and vice versa. However, this would not be the case for long‐lived species with long generation times, and which populations have declined due to anthropogenic effects, such as the blue whale (Balaenoptera musculus). This species was intensively decimated globally to near extinction during the 20th century. Along the Chilean coast, it is estimated that at least 4288 blue whales were hunted from an apparently pre‐exploitation population size (k) of a maximum of 6200 individuals (Southeastern Pacific). Thus, here, we describe the mtDNA (control region) and nDNA (microsatellites) diversities of the Chilean blue whale aggregation site in order to verify the expectation of low genetic diversity in small populations. We then compare our findings with other blue whale aggregations in the Southern Hemisphere. Interestingly, although the estimated population size is small compared with the pre‐whaling era, there is still considerable genetic diversity, even after the population crash, both in mitochondrial (N = 46) and nuclear (N = 52) markers (Hd = 0.890 and Ho = 0.692, respectively). Our results suggest that this diversity could be a consequence of the long generation times and the relatively short period of time elapsed since the end of whaling, which has been observed in other heavily‐exploited whale populations. The genetic variability of blue whales on their southern Chile feeding grounds was similar to that found in other Southern Hemisphere blue whale feeding grounds. Our phylogenetic analysis of mtDNA haplotypes does not show extensive differentiation of populations among Southern Hemisphere blue whale feeding grounds. The present study suggests that although levels of genetic diversity are frequently used as estimators of population health, these parameters depend on the biology of the species and should be taken into account in a monitoring framework study to obtain a more complete picture of the conservation status of a population.  相似文献   

2.
Historical harvesting pushed many whale species to the brink of extinction. Although most Southern Hemisphere populations are slowly recovering, the influence of future climate change on their recovery remains unknown. We investigate the impacts of two anthropogenic pressures—historical commercial whaling and future climate change—on populations of baleen whales (blue, fin, humpback, Antarctic minke, southern right) and their prey (krill and copepods) in the Southern Ocean. We use a climate–biological coupled “Model of Intermediate Complexity for Ecosystem Assessments” (MICE) that links krill and whale population dynamics with climate change drivers, including changes in ocean temperature, primary productivity and sea ice. Models predict negative future impacts of climate change on krill and all whale species, although the magnitude of impacts on whales differs among populations. Despite initial recovery from historical whaling, models predict concerning declines under climate change, even local extinctions by 2100, for Pacific populations of blue, fin and southern right whales, and Atlantic/Indian fin and humpback whales. Predicted declines were a consequence of reduced prey (copepods/krill) from warming and increasing interspecific competition between whale species. We model whale population recovery under an alternative scenario whereby whales adapt their migratory patterns to accommodate changing sea ice in the Antarctic and a shifting prey base. Plasticity in range size and migration was predicted to improve recovery for ice‐associated blue and minke whales. Our study highlights the need for ongoing protection to help depleted whale populations recover, as well as local management to ensure the krill prey base remains viable, but this may have limited success without immediate action to reduce emissions.  相似文献   

3.
Many aspects of blue whale biology are poorly understood. Some of the gaps in our knowledge, such as those regarding their basic taxonomy and seasonal movements, directly affect our ability to monitor and manage blue whale populations. As a step towards filling in some of these gaps, microsatellite and mtDNA sequence analyses were conducted on blue whale samples from the Southern Hemisphere, the eastern tropical Pacific (ETP) and the northeast Pacific. The results indicate that the ETP is differentially used by blue whales from the northern and southern eastern Pacific, with the former showing stronger affinity to the region off Central America known as the Costa Rican Dome, and the latter favouring the waters of Peru and Ecuador. Although the pattern of genetic variation throughout the Southern Hemisphere is compatible with the recently proposed subspecies status of Chilean blue whales, some discrepancies remain between catch lengths and lengths from aerial photography, and not all blue whales in Chilean waters can be assumed to be of this type. Also, the range of the proposed Chilean subspecies, which extends to the Galapagos region of the ETP, at least seasonally, perhaps should include the Costa Rican Dome and the eastern North Pacific as well.  相似文献   

4.
Killer whales, Orcinus orca, are top predators occupying key ecological roles in a variety of ecosystems and are one of the most widely distributed mammals on the planet. In consequence, there has been significant interest in understanding their basic biology and ecology. Long‐term studies of Northern Hemisphere killer whales, particularly in the eastern North Pacific (ENP), have identified three ecologically distinct communities or ecotypes in that region. The success of these prominent ENP studies has led to similar efforts at clarifying the role of killer whale ecology in other regions, including Antarctica. In the Southern Hemisphere, killer whales present a range of behavioural, social and morphological characteristics to biologists, who often interpret this as evidence to categorize individuals or groups, and draw general ecological conclusions about these super‐predators. Morphologically distinct forms (Type A, B, C, and D) occur in the Southern Ocean and studies of these different forms are often presented in conjunction with evidence for specialised ecology and behaviours. Here we review current knowledge of killer whale ecology and ecotyping globally and present a synthesis of existing knowledge. In particular, we highlight the complexity of killer whale ecology in the Southern Hemisphere and examine this in the context of comparatively well‐studied Northern Hemisphere populations. We suggest that assigning erroneous or prefatory ecotypic status in the Southern Hemisphere could be detrimental to subsequent killer whale studies, because unsubstantiated characteristics may be assumed as a result of such classification. On this basis, we also recommend that ecotypic status classification for Southern Ocean killer whale morphotypes be reserved until more evidence‐based ecological and taxonomic data are obtained.  相似文献   

5.
Understanding the degree of genetic exchange between subspecies and populations is vital for the appropriate management of endangered species. Blue whales (Balaenoptera musculus) have two recognized Southern Hemisphere subspecies that show differences in geographic distribution, morphology, vocalizations and genetics. During the austral summer feeding season, the Antarctic blue whale (B. m. intermedia) is found in polar waters and the pygmy blue whale (B. m. brevicauda) in temperate waters. Here, we genetically analyzed samples collected during the feeding season to report on several cases of hybridization between the two recognized blue whale Southern Hemisphere subspecies in a previously unconfirmed sympatric area off Antarctica. This means the pygmy blue whales using waters off Antarctica may migrate and then breed during the austral winter with the Antarctic subspecies. Alternatively, the subspecies may interbreed off Antarctica outside the expected austral winter breeding season. The genetically estimated recent migration rates from the pygmy to Antarctic subspecies were greater than estimates of evolutionary migration rates and previous estimates based on morphology of whaling catches. This discrepancy may be due to differences in the methods or an increase in the proportion of pygmy blue whales off Antarctica within the last four decades. Potential causes for the latter are whaling, anthropogenic climate change or a combination of these and may have led to hybridization between the subspecies. Our findings challenge the current knowledge about the breeding behaviour of the world's largest animal and provide key information that can be incorporated into management and conservation practices for this endangered species.  相似文献   

6.
A clear understanding of population structure is essential for assessing conservation status and implementing management strategies. A small, non-migratory population of humpback whales in the Arabian Sea is classified as “Endangered” on the IUCN Red List of Threatened Species, an assessment constrained by a lack of data, including limited understanding of its relationship to other populations. We analysed 11 microsatellite markers and mitochondrial DNA sequences extracted from 67 Arabian Sea humpback whale tissue samples and compared them to equivalent datasets from the Southern Hemisphere and North Pacific. Results show that the Arabian Sea population is highly distinct; estimates of gene flow and divergence times suggest a Southern Indian Ocean origin but indicate that it has been isolated for approximately 70,000 years, remarkable for a species that is typically highly migratory. Genetic diversity values are significantly lower than those obtained for Southern Hemisphere populations and signatures of ancient and recent genetic bottlenecks were identified. Our findings suggest this is the world''s most isolated humpback whale population, which, when combined with low population abundance estimates and anthropogenic threats, raises concern for its survival. We recommend an amendment of the status of the population to “Critically Endangered” on the IUCN Red List.  相似文献   

7.
To explore the spatio-temporal dynamics of endangered fin whales (Balaenoptera physalus) within the baleen whale (Mysticeti) lineages, we analyzed 148 published mitochondrial genome sequences of baleen whales. We used a Bayesian coalescent approach as well as Bayesian inferences and maximum likelihood methods. The results showed that the fin whales had a single maternal origin, and that there is a significant correlation between geographic location and evolution of global fin whales. The most recent common female ancestor of this species lived approximately 9.88 million years ago (Mya). Here, North Pacific fin whales first appeared about 7.48 Mya, followed by a subsequent divergence in Southern Hemisphere approximately 6.63 Mya and North Atlantic about 4.42 Mya. Relatively recently, approximately 1.76 and 1.42 Mya, there were two additional occurrences of North Pacific populations; one originated from the Southern Hemisphere and the other from an uncertain location. The evolutionary rate of this species was 1.002?×?10?3 substitutions/site/My. Our Bayesian skyline plot illustrates that the fin whale population has the rapid expansion event since ~?2.5 Mya, during the Quaternary glaciation stage. Additionally, this study indicates that the fin whale has a sister group relationship with humpback whale (Meganoptera novaeangliae) within the baleen whale lineages. Of the 16 genomic regions, NADH5 showed the most powerful signal for baleen whale phylogenetics. Interestingly, fin whales have 16 species-specific amino acid residues in eight mitochondrial genes: NADH2, COX2, COX3, ATPase6, ATPase8, NADH4, NADH5, and Cytb.  相似文献   

8.
In the Southwestern Atlantic Ocean, humpback whales migrate every winter to the Brazilian coast for breeding and calving in the Abrolhos Bank. This breeding stock represents the remnants of a larger population heavily exploited during the beginning of the 20th century. Despite its relevance to conservation efforts, the degree of current genetic variation and the migratory relationship with Antarctic feeding areas for this population are still largely unknown. To examine these questions, we sequenced ∼400 bp of the mitochondrial DNA control region from samples taken off the Brazilian coast (n = 171) and near the Antarctic Peninsula (n = 77). The genetic variability of the Brazilian humpback whale breeding population was high and similar to that found in other Southern Hemisphere breeding grounds. Phylogenetic analysis suggested the existence of a new mitochondrial clade that exists at low frequency among Southern Hemisphere populations. Direct comparison between the Brazilian and the Colombia breeding populations and the Antarctic Peninsula feeding population showed no genetic differentiation between this feeding region and the Colombian breeding area or between feeding Areas I and II near the Antarctic Peninsula. In contrast, these populations were genetically distinct from the Brazilian population. Two humpback whales sampled off South Georgia Islands, in the Scotia Sea, shared identical haplotypes to whales from Brazil. Our results, supported by photo-identification and satellite telemetry data, suggest that the main feeding area of the Southern Hemisphere humpback whale population is likely to be located near the South Georgia/South Sandwich Islands area and not in the Antarctic Peninsula.  相似文献   

9.
In the Southern Hemisphere, blue whales are currently divided into two subspecies, Antarctic blue whales (Balaenoptera musculus intermedia) and pygmy blue whales (B. m. brevicauda), but there is some debate about whether Chilean blue whales should also be considered as a separate subspecies. Here, we provide novel morphometric data to directly address this taxonomic question from a biological survey of 60 blue whales taken during the 1965/1966 Chilean whaling season. The data show that maximum body length and mean body length of both sexually mature females and males for Chilean blue whales are intermediate between pygmy and Antarctic blue whales; and that fluke-anus lengths of Chilean blue whales are significantly different from pygmy blue whales, but not necessarily from Antarctic blue whales. There is also some support from the data that snout-eye measurements are different among all three groups. These data provide further confirmation that Chilean blue whales are a distinct population requiring separate management from other blue whale populations, and are also consistent with suggestions that Chilean blue whales are not the same subspecies as pygmy blue whales.  相似文献   

10.
Understanding the seasonal movements and distribution patterns of migratory species over ocean basin scales is vital for appropriate conservation and management measures. However, assessing populations over remote regions is challenging, particularly if they are rare. Blue whales (Balaenoptera musculus spp) are an endangered species found in the Southern and Indian Oceans. Here two recognized subspecies of blue whales and, based on passive acoustic monitoring, four “acoustic populations” occur. Three of these are pygmy blue whale (B.m. brevicauda) populations while the fourth is the Antarctic blue whale (B.m. intermedia). Past whaling catches have dramatically reduced their numbers but recent acoustic recordings show that these oceans are still important habitat for blue whales. Presently little is known about the seasonal movements and degree of overlap of these four populations, particularly in the central Indian Ocean. We examined the geographic and seasonal occurrence of different blue whale acoustic populations using one year of passive acoustic recording from three sites located at different latitudes in the Indian Ocean. The vocalizations of the different blue whale subspecies and acoustic populations were recorded seasonally in different regions. For some call types and locations, there was spatial and temporal overlap, particularly between Antarctic and different pygmy blue whale acoustic populations. Except on the southernmost hydrophone, all three pygmy blue whale acoustic populations were found at different sites or during different seasons, which further suggests that these populations are generally geographically distinct. This unusual blue whale diversity in sub-Antarctic and sub-tropical waters indicates the importance of the area for blue whales in these former whaling grounds.  相似文献   

11.
The distribution patterns of many fishes between the three continents (Africa, Australia, and South America) in the Southern Hemisphere have been uncovered to be influenced by mostly vicariance or historical dispersal. Although some demersal fishes with intercontinental distribution are suggested to be more influenced by current/recent dispersal, few genetic studies have been made for demersal fishes so far. To provide more information for such fishes, genetic divergence was analyzed for two pairs of gadiform species and subspecies distributed around Australasia and South America: the blue grenadier, Macruronus novaezelandiae (from New Zealand) and the Patagonian grenadier, M. magellanicus (from South America) as well as two subspecies of the southern blue whiting, Micromesistius australis pallidus (from New Zealand) and M. a. australis (from South America). The sequence analyses of two mitochondrial DNA regions showed no divergence between Australasian and South American populations of the grenadiers and the southern blue whiting. The microsatellite DNA analysis also indicated significant but very minimal genetic differentiation between the two geographic populations of each pair. These results imply rather recent separation of the two geographic populations. Current/recent dispersal may be an important common factor for determining the distribution of demersal fishes in the Southern Hemisphere. Nonetheless, low but significant genetic differentiation observed requires treating the two populations of the economically important grenadiers and southern blue whiting, respectively, as different stocks for proper resource management.  相似文献   

12.
New blue whale ovarian corpora data from illegal Soviet catches in the Southern Hemisphere and northern Indian Ocean were recovered from the original logbooks. Catches north of 52°S were assumed to be pygmy blue whales ( Balaenoptera musculus brevicauda , n = 1,272); those south of 56°S were assumed to be Antarctic (true) blue whales ( B. m. intermedia , n = 153). Three probable Antarctic blue whales north of 52°S were excluded. Lengths at which 50% and 95% of females become sexually mature ( L 50 and L 95) were estimated from a Bayesian logistic model. These estimates are more precise than previous Japanese estimates because Soviet catches below the legal minimum of 70 ft (21.3 m) were 32 times greater. For pygmy blue whales L 50 was 19.2 m (95% interval 19.1–19.3 m) and L 95 was 20.5 m (95% interval 20.4–20.7 m). Antarctic L 50 (23.4 m, 95% interval 22.9–23.9 m) was much longer than L 50 for pygmy blue whale regions (18.4–19.9 m). The median L 50 for the northern Indian Ocean was 0.5–0.6 m shorter than for pygmy blue whales from other regions; although statistically significant, these small length differences provide little support for northern Indian Ocean blue whales being a separate subspecies, B. m. indica .  相似文献   

13.
Seasonal and spatial variations of blue ( Balaenoptera musculus ) and fin whale ( B. physalus ) calls were analyzed from recordings collected with Acoustic Recording Packages (ARPs) deployed between January 2003 and July 2004 at four circumpolar locations: the Western Antarctic Peninsula (WAP), the Scotia Sea (SS), Eastern Antarctica (EA), and the Ross Sea (RS). Call characteristics were compared among sites using the average pressure spectrum levels from 1 month of data at each location. Presence of calls was analyzed using automatic call detection and acoustic power analysis methods. Blue whale calls were recorded year-round, with the highest detections in February–May and November. This suggests that the blue whale population may not migrate synchronously, and may indicate long duration calls are more common during migrations. Fin whale calls were detected only during February–July. Two distinct fin whale call types were recorded, suggesting a possible separation into two populations. The calls at the EA site had a secondary frequency peak in the pressure spectrum at 99 Hz and the calls at the WAP and the SS sites had a peak at 89 Hz. No fin whale calls were detected at the RS site. Acoustics are a good tool to monitor large whales in the Southern Ocean.  相似文献   

14.
Human‐mediated biological transfers of species have substantially modified many ecosystems with profound environmental and economic consequences. However, in many cases, invasion events are very hard to identify because of the absence of an appropriate baseline of information for receiving sites/regions. In this study, use of high‐resolution genetic markers (single nucleotide polymorphisms – SNPs) highlights the threat of introduced Northern Hemisphere blue mussels (Mytilus galloprovincialis) at a regional scale to Southern Hemisphere lineages of blue mussels via hybridization and introgression. Analysis of a multispecies SNP dataset reveals hotspots of invasive Northern Hemisphere blue mussels in some mainland New Zealand locations, as well as the existence of unique native lineages of blue mussels on remote oceanic islands in the Southern Ocean that are now threatened by invasive mussels. Samples collected from an oil rig that has moved between South Africa, Australia, and New Zealand were identified as invasive Northern Hemisphere mussels, revealing the relative ease with which such non‐native species may be moved from region to region. In combination, our results highlight the existence of unique lineages of mussels (and by extension, presumably of other taxa) on remote offshore islands in the Southern Ocean, the need for more baseline data to help identify bioinvasion events, the ongoing threat of hybridization and introgression posed by invasive species, and the need for greater protection of some of the world's last great remote areas.  相似文献   

15.
Geographic variation in tooth row counts among sand tiger sharks Carcharias taurus (Chondrichthyes), from the SW Atlantic, NW Atlantic and the East China Sea is analyzed in this paper. We found significant differences between sand tigers from the SW Atlantic (Southern Hemisphere population) and each of the other two (Northern Hemisphere) regions in the number of upper lateral tooth rows, and between individuals from the SW Atlantic and the East China Sea in the total number of upper tooth rows. Sand tiger sharks from the two Northern Hemisphere populations did not differ in any of the studied variables. Our results agree with comparisons of vertebral counts between sand tiger sharks from Southern and Northern Hemispheres. Both lines of evidence suggest that Southern and Northern Hemisphere populations of C. taurus were isolated to a larger extent than populations of the Northern Hemisphere. The fossil record of the genus Carcharias begins in the Early Cretaceous and C. taurus is certainly known since the Late Miocene. During the Miocene, the Tethys Sea separating northern and southern land masses was still present and it provided a continuous temperate shallow sea that could allow dispersal of sand tiger sharks along Northern Hemisphere seas. Independent observations on the distribution and evolutionary history of the genera Myripristis , Neoniphon , Sargocentron and Aphanius , and genetic studies on the temperate shark genus Mustelus that indicate a close relationship between the Indo-Pacific M. manazo and the Mediterranean M. asterias suggest that this hypothesis is plausible and deserves to be tested.  相似文献   

16.
Studies of social differentiation between populations of killer whales (Orcinus orca) are important due to the cosmopolitan nature of the species, both in terms of distribution and feeding habits. The following research provides preliminary findings describing the social structure of the killer whale, Orcinus orca, population at subantarctic Marion Island. We provide evidence for consistent, observable patterns of social interactions with animals associating and disassociating in nonrandom patterns. We show that the social structure of this population may follow a new pattern of association, displaying a blend of the traditional resident/transient model displayed in the Northern Hemisphere. However, we emphasize the critical need for further studies related to the sociality, biology, and life history of Southern Ocean killer whales.  相似文献   

17.
Global phylogeographic patterns in Sanionia uncinata are addressed based on information in internal transcribed spacer (ITS) (214 specimens) and the plastid markers trnLtrnF (221) and rpl16 (217). ITS suggests a monophyletic Sanionia and a paraphyletic S. uncinata; this was neither supported nor rejected by plastid data. Northern or Eastern Eurasia and Alaska appear important in the early evolution of Sanionia and some populations dispersed into the Southern Hemisphere relatively early. Some haplotypes or groups of haplotypes are morphologically and ecologically distinct, biologically meaningful units that correspond with S. orthothecioides, S. symmetrica and S. georgicouncinata s.l. The latter includes two species that are indistinguishable by morphology, S. georgicouncinata s.s. (Southern Hemisphere) and S. nivalis (Northern Hemisphere). Tropical African and South American S. uncinata populations have separate origins and the Southern Hemisphere was colonized at least twice. In the northern circum‐Arctic region, the haplotype composition differs between the North Atlantic and Beringian areas. Eastern Eurasia has a higher S. uncinata haplotype diversity than other Holarctic regions, implying less devastating effects of recurrent glacial periods. For Eastern and Western Eurasia, North America and the Southern Hemisphere, most of the haplotype variation was found within the regions, but 14–18% can be referred to among region variation. Plastid haplotype diversity was lower in the Southern Hemisphere than in the Arctic to subarctic, possibly attributable to founder effects. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 19–42.  相似文献   

18.
The Indian Ocean is an area in which a rich suite of cetacean fauna, including at least two subspecies of blue whale, is found; yet little information beyond stranding data and short‐term surveys for this species is available. Pygmy blue whale (Balaenoptera musculus spp.) call data are presented that provide novel information on the seasonal and geographic distribution of these animals. Acoustic data were recorded from January 2002 to December 2003 by hydrophones at three stations of the International Monitoring System, including two near the subequatorial Diego Garcia Atoll and a third southwest of Cape Leeuwin, Australia. Automated spectrogram correlation methods were used to scan for call types attributed to pygmy blue whales. Sri Lanka calls were the most common and were detected year‐round off Diego Garcia. Madagascar calls were only recorded on the northern Diego Garcia hydrophone during May and July, whereas Australia calls were only recorded at Cape Leeuwin, between December and June. Differences in geographic and seasonal patterns of these three distinct call types suggest that they may represent separate acoustic populations of pygmy blue whales and that these “acoustic populations” should be considered when assessing conservation needs of blue whales in the Indian Ocean.  相似文献   

19.
The North Pacific right whale, Eubalaena japonica, is one of the most endangered species of whale in the world. On 10 August 2004, two right whales were located in the Bering Sea using headings to right whale calls provided by directional sonobuoys. A satellite-monitored radio tag attached to one of these whales functioned for 40 days. Over the 40-day period, this whale moved throughout a large part of the southeast Bering Sea shelf, including areas of the outer-shelf where right whales have not been seen in decades. In September, multiple right whales were acoustically located and subsequently sighted by another survey vessel approaching a near-real-time position from the tag. An analysis of photographs confirmed at least 17 individual whales (not including the tagged whales). Genetic analysis of biopsy samples identified 17 individuals: 10 males and 7 females. The discovery of seven females was significant, as only one female had been identified in the past. Genetics also confirmed the presence of at least two calves. Although the future of this population is highly uncertain, the discovery of additional females and calves gives some hope that this most critically endangered of all whale populations may still possess the capacity to recover.  相似文献   

20.
How do populations of highly mobile species inhabiting open environments become reproductively isolated and evolve into new species? We test the hypothesis that elevated ocean‐surface temperatures can facilitate allopatry among pelagic populations and thus promote speciation. Oceanographic modelling has shown that increasing surface temperatures cause localization and reduction of upwelling, leading to fragmentation of feeding areas critical to pelagic species. We test our hypothesis by genetic analyses of populations of two closely related baleen whales, the Antarctic minke whale (Balaenoptera bonaerensis) and common minke whale (Balaenoptera acutorostrata) whose current distributions and migration patterns extent are largely determined by areas of consistent upwelling with high primary production. Phylogeographic and population genetic analyses of mitochondrial DNA control‐region nucleotide sequences collected from 467 whales sampled in four different ocean basins were employed to infer the evolutionary relationship among populations of B. acutorostrata by rooting an intraspecific phylogeny with a population of B. bonaerensis. Our findings suggest that the two species diverged in the Southern Hemisphere less than 5 million years ago (Ma). This estimate places the speciation event during a period of extended global warming in the Pliocene. We propose that elevated ocean temperatures in the period facilitated allopatric speciation by disrupting the continuous belt of upwelling maintained by the Antarctic Circumpolar Current. Our analyses revealed that the current populations of B. acutorostrata likely diverged after the Pliocene some 1.5 Ma when global temperatures had decreased and presumably coinciding with the re‐establishment of the polar–equatorial temperature gradient that ultimately drives upwelling. In most population samples, we detected genetic signatures of exponential population expansions, consistent with the notion of increasing carrying capacity after the Pliocene. Our hypothesis that prolonged periods of global warming facilitate speciation in pelagic marine species that depend on upwelling should be tested by comparative analyses in other pelagic species.  相似文献   

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