Abundance and residency dynamics of the Indo-Pacific humpback dolphin,Sousa chinensis,in the Dafengjiang River Estuary,China

Robust population size estimates are essential for informing population conservation status. Residency dynamics show population habitat use through time. Population size of Indo-Pacific humpback dolphins (Sousa chinensis) has been extensively investigated in Chinese waters, but their residency dynamics are rarely known. Mark-recapture analysis based on photo-identification records was applied to humpback dolphins in the Dafengjiang River Estuary habitat, one of the key habitats in the northern Beibu Gulf, China. Movement analyses based on lagged identification rate indicated the humpback dolphins spent, on average, 78.5 days inside and 46.9 days outside the survey area. Within the study area, the humpback dolphin abundance was 83 identifiable dolphins. A total of 353–430 humpback dolphins, estimated by POPAN modeling, were involved in this fluid habitat-use dynamic. Robust Design analysis showed strong seasonality in humpback dolphin abundance and emigration probability, implying a movement- and habitat-use pattern likely associated with spatiotemporal distribution of oceanographic characteristics and prey occurrences. Population surveys and conservation measures currently conducted in Chinese waters seldom consider seasonality in movements between habitat patches, which can be addressed by genetic analyses across habitats and cross-matching photo-identification records among neighboring habitats.     

Passive Acoustic Monitoring the Diel,Lunar, Seasonal and Tidal Patterns in the Biosonar Activity of the Indo-Pacific Humpback Dolphins (Sousa chinensis) in the Pearl River Estuary,China

A growing demand for sustainable energy has led to an increase in construction of offshore windfarms. Guishan windmill farm will be constructed in the Pearl River Estuary, China, which sustains the world’s largest known population of Indo-Pacific humpback dolphins (Sousa chinensis). Dolphin conservation is an urgent issue in this region. By using passive acoustic monitoring, a baseline distribution of data on this species in the Pearl River Estuary during pre-construction period had been collected. Dolphin biosonar detection and its diel, lunar, seasonal and tidal patterns were examined using a Generalized Linear Model. Significant higher echolocation detections at night than during the day, in winter-spring than in summer-autumn, at high tide than at flood tide were recognized. Significant higher echolocation detections during the new moon were recognized at night time. The diel, lunar and seasonal patterns for the echolocation encounter duration also significantly varied. These patterns could be due to the spatial-temporal variability of dolphin prey and illumination conditions. The baseline information will be useful for driving further effective action on the conservation of this species and in facilitating later assessments of the effects of the offshore windfarm on the dolphins by comparing the baseline to post construction and post mitigation efforts.     

Multiple lines of evidence for an Australasian geographic boundary in the Indo-Pacific humpback dolphin (<Emphasis Type="Italic">Sousa chinensis</Emphasis>): population or species divergence?

The taxonomic status of humpback dolphins (genus Sousa, sub-family Delphininae) is unresolved. While the classification of this genus ranges from a single to three nominal species, the International Union for Conservation of Nature and the International Whaling Commission only recognise a ‘two-species’ taxonomy (S. teuszii in west Africa, and S. chinensis in the Indo-Pacific). Under the IUCN (2008), S. chinensis is listed as ‘near threatened’, but is only considered as a ‘migratory’ species in Australia. Taxonomic resolution of the genus Sousa is needed to define particular conservation status and develop appropriate management actions. Using phylogenetic analyses of 1,082 bp of mitochondrial and 1,916 bp of nuclear DNA, we provide multiple lines of genetic evidence for the genetic distinction of S. chinensis in China and Indonesia from S. chinensis in Australia. The separation of Australian Sousa from Sousa of Southeast Asia requires a review of their current conservation status and respective management actions.     

Whistle characteristics of a newly recorded Indo-Pacific humpback dolphin (Sousa chinensis) population in waters southwest of Hainan Island,China, differ from other humpback dolphin populations

Indo-Pacific humpback dolphins (Sousa chinensis) use whistles to communicate with their conspecifics. Little is known about the acoustic repertoire of Indo-Pacific humpback dolphins in waters southwest of Hainan Island, a newly recorded population in 2014. In this study, whistles of Hainan humpback dolphin population were collected by using autonomous acoustic recorders. The fundamental frequencies and durations of whistles were in ranges of 0.71–21.35 kHz and 0.06–2.22 s, respectively. Significant intraspecific differences in duration and frequency of whistles were found between the Hainan population and the other geographically neighboring populations (in Chinese waters) or the population in Malaysia waters. Compared with other Sousa species, significant interspecific differences were also observed. Based on clustering analysis, the whistle parameters of neighboring populations were likely similar to each other. Significant differences were found between humpback dolphins in waters southwest of Hainan Island and those dolphins in the neighboring areas, supporting the hypothesis that this population may be independent. Ambient noise measurements in waters of Hainan Island, Zhanjiang, and Sanniang Bay showed that humpback dolphin populations may use whistles with longer duration, lower frequency, and fewer inflection points for more effective communication to adapt to a noisier environment.     

A preliminary investigation on genetic diversity of Sousa chinensis in the Pearl River Estuary and Xiamen of Chinese waters

In this study, the mitochondrial DNA (mtDNA) control region and the mitochondrial cytochrome b gene of stranded Indo-Pacific humpback dolphin (Sousa chinensis) samples from the Pearl River Estuary and Xiamen waters were sequenced and analyzed. The result of mtDNA control region revealed 34 variable sites and four unique haplotypes (named as A, B, C and D) identified among the total samples from these two water areas, and the most common haplotype (A) was shared by 75% of the dolphins sampled from the two water areas. The haplotypic diversity (h) was 0.455 and the nucleotide diversity (π) was 0.0088. The phylogenetic analysis showed that the haplotype A, C, and D were closely related, but the haplotype B (unique for XM01 from Xiamen) was far from the other three. By scanning cytochrome b fragments, two haplotypes (A and B) were identified in these two water areas, and the most common haplotype (A) was shared by 91.67% individuals, while XM01 from Xiamen as the only exception. The date suggest that there is a possibility of gene exchange between the two populations in the Pearl River Estuary and Xiamen waters, and there possibly exists a unique maternal lineage in Xiamen waters.     

Fine‐scale habitat use in Indo‐Pacific humpback dolphins,Sousa chinensis,may be more influenced by fish rather than vessels in the Pearl River Estuary,China

Shifts in habitat use and distribution patterns in dolphins are often concerns that can result from habitat degradation. We investigated how potential changes to a habitat from human activity may alter dolphin distributions within Lingding Bay in the Pearl River Estuary, China, by studying the relationship between fish choruses, vessel presence and Indo‐Pacific humpback dolphin (Sousa chinensis) detection rates. Analyses revealed temporal and spatial variation within fish choruses, vessel presence and dolphin detection rates. After accounting for any temporal autocorrelation, correlations between fish choruses and dolphin detection rates were also found; however, no relationship between fish choruses and vessel presence or dolphin detection rates and vessel presence were observed. Furthermore, fewer dolphins were detected at sites where fish activity was less intense. Thus fish activity, rather than vessels, may be a key factor influencing the distribution of the dolphins within the estuary. These findings emphasize the risk of potential shifts in habitat use for Indo‐Pacific humpback dolphins due to detrimental changes to prey availability and dolphin feeding grounds from human activity, such as overfishing and coastal developments, within the estuary. This is a critical conservation issue for this dolphin population that is facing intense anthropogenic pressure.     

Population Differentiation and Hybridisation of Australian Snubfin (Orcaella heinsohni) and Indo-Pacific Humpback (Sousa chinensis) Dolphins in North-Western Australia

Little is known about the Australian snubfin (Orcaella heinsohni) and Indo-Pacific humpback (Sousa chinensis) dolphins (‘snubfin’ and ‘humpback dolphins’, hereafter) of north-western Australia. While both species are listed as ‘near threatened’ by the IUCN, data deficiencies are impeding rigorous assessment of their conservation status across Australia. Understanding the genetic structure of populations, including levels of gene flow among populations, is important for the assessment of conservation status and the effective management of a species. Using nuclear and mitochondrial DNA markers, we assessed population genetic diversity and differentiation between snubfin dolphins from Cygnet (n = 32) and Roebuck Bays (n = 25), and humpback dolphins from the Dampier Archipelago (n = 19) and the North West Cape (n = 18). All sampling locations were separated by geographic distances >200 km. For each species, we found significant genetic differentiation between sampling locations based on 12 (for snubfin dolphins) and 13 (for humpback dolphins) microsatellite loci (F _ST = 0.05–0.09; P<0.001) and a 422 bp sequence of the mitochondrial control region (F _ST = 0.50–0.70; P<0.001). The estimated proportion of migrants in a population ranged from 0.01 (95% CI 0.00–0.06) to 0.13 (0.03–0.24). These are the first estimates of genetic diversity and differentiation for snubfin and humpback dolphins in Western Australia, providing valuable information towards the assessment of their conservation status in this rapidly developing region. Our results suggest that north-western Australian snubfin and humpback dolphins may exist as metapopulations of small, largely isolated population fragments, and should be managed accordingly. Management plans should seek to maintain effective population size and gene flow. Additionally, while interactions of a socio-sexual nature between these two species have been observed previously, here we provide strong evidence for the first documented case of hybridisation between a female snubfin dolphin and a male humpback dolphin.     

厦门湾中华白海豚潜在生态廊道识别及人类活动干扰评估

生态廊道具有维持或恢复生态连通性的功能,对于连接生物栖息地、保护物种多样性具有重要意义。现有的生态廊道研究主要集中于陆地,而海洋生态系统具有水体广泛连通、缺乏直观的景观斑块等特点,导致海洋生态廊道的研究成为长期以来的科学难题。以栖息地位于厦门湾的国家一级保护动物中华白海豚(Sousa chinensis)为对象,尝试基于物种分布模型和最小成本路径分析法建立海洋生态廊道的识别方法。研究采用物种分布模型识别厦门湾内中华白海豚的适宜生境分布区和节点,并利用模型产出的生境适宜性结果生成海洋中的阻力表面,模拟计算节点与节点间在阻力表面上的最小成本路径,从而生成物种扩散网络。研究结果显示,厦门湾中华白海豚的分布主要受到航道距离、到岸线距离和叶绿素浓度三项因素的影响,主要适宜生境位于西海域至九龙江口和大嶝海域。潜在的核心生态廊道面积93.19km²,次级生态廊道面积170.41km²,九龙江口-鼓浪屿南侧-黄厝-大小嶝岛沿线可能是厦门湾中华白海豚的主要迁移路线。在此基础上,从用海空间重叠和桥梁影响两方面开展了人类活动对廊道的干扰评估。评估结果显示旅游活动和...     

Integrating multiple lines of evidence to better understand the evolutionary divergence of humpback dolphins along their entire distribution range: a new dolphin species in Australian waters?

The conservation of humpback dolphins, distributed in coastal waters of the Indo‐West Pacific and eastern Atlantic Oceans, has been hindered by a lack of understanding about the number of species in the genus (Sousa) and their population structure. To address this issue, we present a combined analysis of genetic and morphologic data collected from beach‐cast, remote‐biopsied and museum specimens from throughout the known Sousa range. We extracted genetic sequence data from 235 samples from extant populations and explored the mitochondrial control region and four nuclear introns through phylogenetic, population‐level and population aggregation frameworks. In addition, 180 cranial specimens from the same geographical regions allowed comparisons of 24 morphological characters through multivariate analyses. The genetic and morphological data showed significant and concordant patterns of geographical segregation, which are typical for the kind of demographic isolation displayed by species units, across the Sousa genus distribution range. Based on our combined genetic and morphological analyses, there is convincing evidence for at least four species within the genus (S. teuszii in the Atlantic off West Africa, S. plumbea in the central and western Indian Ocean, S. chinensis in the eastern Indian and West Pacific Oceans, and a new as‐yet‐unnamed species off northern Australia).     

Distribution and abundance of Indo‐Pacific humpback dolphins in Leizhou Bay,China

Abstract

The Indo‐Pacific humpback dolphin (Sousa chinensis) is under severe threat in Chinese waters. Only four small populations of this dolphin are known to survive in the East China Sea and South China Sea. We conducted exploratory surveys from June to September 2005, and found a population of humpback dolphins in Leizhou Bay. Thirty‐eight individuals were identified and catalogued, of which 26 were photographed and re‐identified on more than one occasion, and nine were sighted on five or more occasions each. Our preliminary estimate of the size of the Leizhou population is about 237 individuals, second only to the Pearl River estuary population. We suggest that Leizhou Bay has the potential to serve as a “humpback dolphin sanctuary” in Chinese waters.     

Mercury and Selenium in Stranded Indo-Pacific Humpback Dolphins and Implications for Their Trophic Transfer in Food Chains

As top predators in the Pearl River Estuary (PRE) of China, Indo-Pacific humpback dolphins (Sousa chinensis) are bioindicators for examining regional trends of environmental contaminants in the PRE. We examined samples from stranded S. chinensis in the PRE, collected since 2004, to study the distribution and fate of total mercury (THg), methylmercury (MeHg) and selenium (Se) in the major tissues, in individuals at different ages and their prey fishes from the PRE. This study also investigated the potential protective effects of Se against the toxicities of accumulated THg. Dolphin livers contained the highest concentrations of THg (32.34±58.98 µg g⁻¹ dw) and Se (15.16±3.66 µg g⁻¹ dw), which were significantly different from those found in kidneys and muscles, whereas the highest residue of MeHg (1.02±1.11 µg g⁻¹ dw) was found in dolphin muscles. Concentrations of both THg and MeHg in the liver, kidney and muscle of dolphins showed a significantly positive correlation with age. The biomagnification factors (BMFs) of inorganic mercury (Hg_inorg) in dolphin livers (350×) and MeHg in muscles (18.7×) through the prey fishes were the highest among all three dolphin tissues, whereas the BMFs of Se were much lower in all dolphin tissues. The lower proportion of MeHg in THg and higher Se/THg ratios in tissues were demonstrated. Our studies suggested that S. chinensis might have the potential to detoxify Hg via the demethylation of MeHg and the formation of tiemannite (HgSe) in the liver and kidney. The lower threshold of hepatic THg concentrations for the equimolar accumulation of Se and Hg in S. chinensis suggests that this species has a greater sensitivity to THg concentrations than is found in striped dolphins and Dall’s porpoises.     

POPULATION ANALYSES OF INDO-PACIFIC HUMPBACK DOLPHINS SOUSA CHINENSIS IN ALGOA BAY,EASTERN CAPE,SOUTH AFRICA1

Mark-recapture analyses were performed on photo-identification data for Indo-Pacific humpback dolphins (Sousa chinensis) inhabiting Algoa Bay, on the Eastern Cape coast of South Africa. Boat-based photo-ID surveys were undertaken between May 1991 and May 1994. The rate of discovery of newly identified dolphins, distribution of sightings and frequency of resightings of known individuals indicate a high level of seasonal immigration of humpback dolphins into, and emigration from, the Algoa Bay region in summer. Consequently, humpback dolphins from Algoa Bay appear to be part of a substantially larger population that uses a considerable length of the coastal zone. The minimum population size is estimated to be about 466 dolphins.     

High genetic differentiation of Indo‐Pacific humpback dolphins (Sousa chinensis) along the Asian Coast of the Pacific Ocean

The Indo‐Pacific humpback dolphin (Sousa chinensis) is a vulnerable marine mammal species that inhabits shallow, coastal waters from Southeast China, southward throughout Southeast Asia, and westward around the Bay of Bengal to eastern India. Polymorphic microsatellites are useful for elucidating ecological and population genetics‐related questions. Here, 18 new polymorphic microsatellites were developed from S. chinensis genomic DNA by Illumina MiSeq sequencing. Population genetic analyses were conducted on 42 S. chinensis individuals from three geographic locations, including the Xiamen Bay of China, the Western Gulf of Thailand, and Andaman Sea. Our microsatellite data revealed a strong and significant population structure among the three sampling regions (overall F _ST = 0.371, p = .001). Pairwise mutual information index also demonstrated high levels of genetic differentiation between different region pairs (values range from 0.272 to 0.339, p < .001). Moreover, Structure analysis inferred three genetic clusters, with the high assignment probabilities of 95.92%, 99.47%, and 99.68%, respectively. Principal coordinate analysis plots of individuals divided entire genotypes into three clusters, indicating high level of genetic differentiation. Our results indicated the strong genetic structure in S. chinensis populations is a result of geographic distances. Other factors such as environmental variables, anthropogenic interference, and social behavior may also have contributed to population differentiation.     

Assessing the Underwater Acoustics of the World's Largest Vibration Hammer (OCTA-KONG) and Its Potential Effects on the Indo-Pacific Humpbacked Dolphin (Sousa chinensis)

Anthropogenic noise in aquatic environments is a worldwide concern due to its potential adverse effects on the environment and aquatic life. The Hongkong-Zhuhai-Macao Bridge is currently under construction in the Pearl River Estuary, a hot spot for the Indo-Pacific humpbacked dolphin (Sousa chinensis) in China. The OCTA-KONG, the world''s largest vibration hammer, is being used during this construction project to drive or extract steel shell piles 22 m in diameter. This activity poses a substantial threat to marine mammals, and an environmental assessment is critically needed. The underwater acoustic properties of the OCTA-KONG were analyzed, and the potential impacts of the underwater acoustic energy on Sousa, including auditory masking and physiological impacts, were assessed. The fundamental frequency of the OCTA-KONG vibration ranged from 15 Hz to 16 Hz, and the noise increments were below 20 kHz, with a dominant frequency and energy below 10 kHz. The resulting sounds are most likely detectable by Sousa over distances of up to 3.5 km from the source. Although Sousa clicks do not appear to be adversely affected, Sousa whistles are susceptible to auditory masking, which may negatively impact this species'' social life. Therefore, a safety zone with a radius of 500 m is proposed. Although the zero-to-peak source level (SL) of the OCTA-KONG was lower than the physiological damage level, the maximum root-mean-square SL exceeded the cetacean safety exposure level on several occasions. Moreover, the majority of the unweighted cumulative source sound exposure levels (SSELs) and the cetacean auditory weighted cumulative SSELs exceeded the acoustic threshold levels for the onset of temporary threshold shift, a type of potentially recoverable auditory damage resulting from prolonged sound exposure. These findings may aid in the identification and design of appropriate mitigation methods, such as the use of air bubble curtains, “soft start” and “power down” techniques.     

Bycatch and strandings programs as ecological indicators for data-limited cetaceans

An integrated approach of using strandings and bycatch data may provide an indicator of long-term trends for data-limited cetaceans. Strandings programs can give a faithful representation of the species composition of cetacean assemblages, while standardised bycatch rates can provide a measure of relative abundance. Comparing the two datasets may also facilitate managing impacts by understanding which species, sex or sizes are the most vulnerable to interactions with fisheries gear. Here we apply this approach to two long-term datasets in East Australia, bycatch in the Queensland Shark Control Program (QSCP, 1992–2012) and strandings in the Queensland Marine Wildlife Strandings and Mortality Program (StrandNet, 1996–2012). Short-beaked common dolphins, Delphinus delphis, were markedly more frequent in bycatch than in the strandings dataset, suggesting that they are more prone to being incidentally caught than other cetacean species in the region. The reverse was true for humpback whales, Megaptera novaeangliae, bottlenose dolphins, Tursiops spp.; and species predominantly found in offshore waters. QSCP bycatch was strongly skewed towards females for short-beaked common dolphins, and towards smaller sizes for Australian humpback dolphins, Sousa sahulensis. Overall, both datasets demonstrated similar seasonality and a similar long-term increase from 1996 until 2008. Analysis on a species-by-species basis was then used to explore potential explanations for long-term trends, which ranged from a recovering stock (humpback whales) to a shift in habitat use (short-beaked common dolphins).     

Life history of the Indo‐Pacific humpback dolphin in the Pearl River Estuary,southern China

We studied life history characteristics of the Hong Kong/Pearl River Estuary population of Indo‐Pacific humpback dolphins (Sousa chinensis), based on data from 120 specimens stranded between 1995 and 2009, 40 individuals biopsied at sea, and a long‐term (14+ yr) photo‐identification study. Ages were determined for 112 specimens by thin‐sectioning teeth and counting growth layer groups. Estimated length at birth was 101 cm. Longevity was at least 38 yr, and there was little difference in growth patterns of males and females. Growth was described by a Bayesian two‐phase Gompertz model; asymptotic length was reached at 249 cm. The tooth pulp cavity filled at an average of 18.5 yr of age. Physical maturity was reached at between 14 and 17 yr of age, apparently a few years after attainment of sexual maturity. Maximum lengths and weights of about 268 cm and 240 kg were attained. Females appear to lose all their spots by 30 yr, although males may retain some spotting throughout life. Calving occurred throughout the year, with a broad peak from March to June. Of 60 females monitored at sea for >14 yr of the study, none were documented to have more than three calves, suggestive of low reproductive output or low calf survival.     

Morphological and genetic differentiation of two loliginid squids, Uroteuthis (Photololigo) chinensis and Uroteuthis (Photololigo) edulis (Cephalopoda: Loliginidae), in Asia

The squids Uroteuthis (Photololigo) edulis and Uroteuthis (Photololigo) chinensis (family Loliginidae) are commercially important fishery species in many coastal regions of Asia. The morphologies of these two squids are very similar, and identification based on morphology has been inadequate. The occurrence of cryptic species in the family Loliginidae has been reported. The widely distributed U. (P.) chinensis and U. (P.) edulis are believed to comprise several cryptic species. In this study, the taxonomic status of the two species in East Asia was elucidated by morphological and genetic analyses. Analysis of U. (P.) chinensis from Hong Kong and Xiamen (China) and U. (P.) edulis from Yamaguchi (Japan) and Shanghai (China) was performed in order to determine the effectiveness of different morphometric variables in discriminating between the two species. Multivariate analysis of 27 morphometric indices revealed no new morphological characters for the taxonomic identification of the two taxa, which can be distinguished by the teeth shape and number on arm sucker rings, and the percentage of hectocotylized part of left arm IV of males. The morphometric differences between U. (P.) edulis individuals from the two localities is most probably due to differences in the maturity stages of the sampled individuals between the two localities. Genetic analysis based on the mitochondrial COI and 16S rRNA genes revealed a high divergence of 15.5% and 7.5% respectively, indicating that U. (P.) edulis and U. (P.) chinensis are distinct species.     

Molecular footprints of inshore aquatic adaptation in Indo-Pacific humpback dolphin (Sousa chinensis)

The Indo-Pacific humpback dolphin, Sousa chinensis, being a member of cetaceans, had fully adapted to inshore waters. As a threatened marine mammal, little molecular information available for understanding the genetic basis of ecological adaptation. We firstly sequenced and obtained the draft genome map of S. chinensis. Phylogenetic analysis in this study, based on the single copy orthologous genes of the draft genome, is consistent with traditional phylogenetic classification. The comparative genomic analysis indicated that S. chinensis had 494 species-specific gene families, which involved immune, DNA repair and sensory systems associated with the potential adaption mechanism. We also identified the expansion and positive selection genes in S. chinensis lineage to investigate the potential adaptation mechanism. Our study provided the potential insight into the molecular bases of ecological adaptation in Indo-Pacific humpback dolphin and will be also valuable for future understanding the ecological adaptation and evolution of cetaceans at the genomic level.     

Site fidelity and association patterns of Indo-Pacific humpback dolphins off the east coast of Zhanjiang,China

This study examined the population of Indo-Pacific humpback dolphins, Sousa chinensis, inhabiting the waters off the east coast of Zhanjiang, China. A total of 116 dolphins were identified during 147 boat-based surveys, completed between June 2005 and June 2007. Abundance estimates indicated that a small population of 268 dolphins (95% CI = 189–413) inhabited this coastal area. The sighting frequencies of identified dolphins varied between one and five when the 2006 and 2007 data were organized into six occasions. Twenty-three percent (27 individuals) of the cataloged dolphins were identified in more than one calendar year. The two animals (ZJ001, ZJ011) with the highest degree of site fidelity were present in the study area for a period of 23 months. The most frequently (15 times) sighted dolphin (ZJ046) was recorded in nine (56%) of the 16 months surveyed. This evidence indicates the interannual site fidelity of S. chinensis to the area off the eastern coast of Zhanjiang City. Range sizes of eight identified individuals calculated by the minimum convex polygon varied from 2.07 to 331.20 km². Associations between 34 individuals sighted at least three times and more than three times were measured by the half-weight index and SOCPROG program. The majority of these dolphins (80.57%) were not seen together during surveys and the mean association rate between dyads was low at 0.05. Temporal analyses for all the individuals were conducted using a lagged association rate. The results indicated a non-random social structure made by constant companions.