Population genetic structure and taxonomy of the common dolphin (Delphinus sp.) at its southernmost range limit: New Zealand waters

New Zealand is the southernmost limit of the common dolphin's (genus Delphinus) distribution in the Pacific Ocean. In this area, common dolphins occur in both coastal and oceanic habitats, exhibit seasonal and resident occurrence, and present high morphological variability. Here we investigated the population structure and the taxonomic identity of common dolphins (Delphinus sp.) within New Zealand waters using 14 microsatellite loci, 577 bp of the mtDNA control region, and 1,120 bp of the mtDNA cytochrome b gene across 90 individuals. We found high genetic variability and evidence of population expansion. Phylogenetic analyses conducted to clarify the taxonomic status of New Zealand common dolphins did not show any clustering reflecting geographic origin or morphotypes. The microsatellite analysis showed genetic differentiation between Coastal and Oceanic putative populations, while mtDNA revealed significant genetic differentiation only between the Hauraki Gulf and other putative groups. Our results suggest that differences in habitat choice and possible female site fidelity may play a role in shaping population structure of New Zealand common dolphins.     

Behavior of free-ranging common dolphins (Delphinus sp.) in the Hauraki Gulf, New Zealand

Here we present the first data describing the behavior of common dolphins ( Delphinus sp.) in the Hauraki Gulf, New Zealand. Activity budgets are used to assess the effects of diel, season, depth, sea surface temperature, group size, and composition on dolphin behavior. Additionally, the presence/absence of Bryde's whale ( Balaenoptera brydei ) and Australasian gannet ( Morus serrator ) is examined in relation to dolphin behavior. Behavioral data were collected from 686 independent dolphin groups during boat-based surveys conducted between February 2002 and January 2005. Foraging (46.7%) and social (7.2%) were the most and least frequently observed behaviors, respectively. Travel (28.9%), mill (9.5%), and rest (7.7%) accounted for the remainder of the activity budget. Behavior varied seasonally, with the highest proportion of foraging and resting groups observed during the spring and autumn, respectively. Behavior also varied with water depth, with foraging animals observed in the deepest and resting groups recorded in the shallowest regions of the Hauraki Gulf. A correlation between group size and behavior was evident, although behavior did not vary with the composition of dolphin groups. Resting, milling, and socializing animals were more frequently observed in smaller group sizes. Foraging behavior was prevalent in both small and large group sizes, suggesting foraging plasticity exists within this population. Behavior differed between single- and multispecies groups, with foraging more frequent in multispecies groups. Resting, milling, or socializing was rarely observed in the presence of any associated species, indicating the primary mechanism for association is likely prey related.     

Hematological,biochemical, and morphological parameters as prognostic indicators for stranded common dolphins (Delphinus delphis) from Cape Cod,Massachusetts, U.S.A.

The current paucity of published blood values and other clinically relevant data for short‐beaked common dolphins, Delphinus delphis, hinders the ability of veterinarians and responders to make well‐informed diagnoses and disposition decisions regarding live strandings of this species. This study examined hematologic, clinical chemistry, and physical parameters from 26 stranded common dolphins on Cape Cod, Massachusetts, in light of their postrelease survival data to evaluate each parameter's efficacy as a prognostic indicator. Statistically and clinically significant differences were found between failed and survived dolphins, including lower hematocrit, hemoglobin, TCO₂, and bicarbonate and higher blood urea nitrogen, uric acid, and length‐to‐girth ratios in animals that failed. In general when compared to survivors, failed dolphins exhibited acidosis, dehydration, lower PCVs, and decreased body condition. Additionally, failed dolphins had the highest ALT, AST, CK, LDH, GGT, and lactate values. These blood values combined with necropsy findings indicate that there are likely a variety of factors affecting postrelease survival, including both preexisting illness and stranding‐induced conditions such as capture myopathy. Closer evaluation of these parameters for stranded common dolphins on point of care analyzers in the field may allow stranding personnel to make better disposition decisions in the future.     

Physical maturity in common bottlenose dolphins (Tursiops truncatus) from Sarasota Bay,Florida

Cetacean physical maturity is defined by growth cessation and complete fusion of epiphyses to vertebral bodies indicated by invisible sutures. Many studies have shown epiphyseal fusion is highly variable among individuals. In-depth examinations into fusion variability are lacking. We analyzed vertebrae of 37 (n = 21 female, n = 16 male) stranded common bottlenose dolphins (Tursiops truncatus) from the well-studied Gulf of Mexico, Sarasota Bay community. For each specimen, vertebrae were examined by vertebral region for degree of fusion anteriorly and posteriorly of each centrum and categorized from unfused to fused in five degrees. An ordinal logistic regression was used to estimate degree of fusion probability for each epiphysis. The model had fixed effects for age, number of offspring, sex, sexual maturity, and a random effect for epiphysis. Results show that age/reproductive status significantly explains an individual's degree of fusion. Adult females with fewer calves had more fusion than those with more reproductive experience across multiple ages. Access to long-term observational and sample data on the dolphins residing in the area served by Mote Marine Laboratory's Stranding Investigations Program offers a unique opportunity to examine the relationship between energetic demands of reproduction (calcium production/reproductive output) versus preconceived definitions of physical maturity (skeletal fusion) more closely.     

Cranial morphology and taxonomic resolution of some dolphin taxa (Delphinidae) in Australian waters,with a focus on the genus Tursiops

Phylogenetic relationships in the family Delphinidae have been widely debated. We examined 347 skulls of Tursiops, Stenella, Delphinus, Steno, Lagenodelphis, and Sousa in order to resolve the phylogenetic position of Australian species of Tursiops. Five Tursiops type specimens were included. Cranial morphology was described using 2‐dimensional (2‐D) and 3‐dimensional geometric morphometrics (3‐GM), counts and categorical data. Analyses showed a clear morphological separation of Tursiops, including type specimens, from other genera. The three Stenella species did not cluster together. Stenella attenuata clustered with Delphinus delphis, and Stenella coeruleoalba with Lagenodelphis hosei. Length and width of the skull and rostrum were important discriminators in both methods. For 3‐D data, round vs. angular posterior skull shape distinguished some genera. Taxa that overlapped in the multivariate analyses had different mean tooth counts. Our study challenges genetic studies that identified Tursiops as polyphyletic, with T. aduncus closer to S. attenuata.     

Whistle characteristics of common dolphins (Delphinus sp.) in the Hauraki Gulf,New Zealand

Quantifying the vocal repertoire of a species is critical for subsequent analysis of signal functionality, geographic variation, and social relevance. However, the vocalizations of free‐ranging common dolphins (Delphinus sp.) have not previously been described from New Zealand waters. We present the first quantitative analysis of whistle characteristics to be undertaken on the New Zealand population. Acoustic data were collected in the Hauraki Gulf, North Island from 28 independent dolphin group encounters. A total of 11,715 whistles were collected from 105.1 min of recordings. Seven whistle contours were identified containing 29 subtypes. Vocalizations spanned from 3.2 to 23 kHz, with most whistles occurring between 11 and 13 kHz. Whistle duration ranged from 0.01 to 4.00 s (mean ± SD; 0.27 ± 0.32). Of the 2,663 whistles analyzed, 82% have previously been identified within U.K. populations. An additional six contours, apparently unique to New Zealand Delphinus were also identified. Data presented here offer a first insight into the whistle characteristics of New Zealand Delphinus. Comparisons with previously studied populations reveal marked differences in the whistle frequency and modulation of the New Zealand population. Interpopulation differences suggest behavior and the local environment likely play a role in shaping the vocal repertoire of this species.     

Comparison of reproductive parameters for populations of eastern North Pacific common dolphins: Delphinus capensis and D. delphis

Reproductive parameters were estimated and compared for eastern North Pacific populations of common dolphins using specimen and photogrammetric data. Age and length data for Delphinus capensis and D. delphis specimens recovered as bycatch or strandings were used to estimate the postnatal growth rates needed to estimate age for calves measured in aerial photographs. Bayesian methods propagated uncertainty among models and revealed that the 2009 cohort of calves had birth dates centered on 6 March 2009 for D. capensis and 12 December 2008 for D. delphis. The evidence for discrete calving seasons suggests a mechanism of reproductive isolation has evolved between species. Photogrammetric data and Bayesian methods were also used to estimate the average length at which calves swim independently: 145.1 cm (≈ 11.1 mo) in D. capensis and 140.1 cm (≈ 14.0 mo) in D. delphis, and the proportion of calves (calves/dolphins counted): 0.045 in D. capensis and 0.069 in D. delphis. The latter parameter was converted to an index of calf production (calf/female dolphin) that was >50% lower than pregnancy rates suggesting few births occurred during the study year. Comparisons of regional differences in calf production suggest variability in habitat use patterns within the study area.     

Standardization and application of metrics to quantify human‐interaction behaviors by the bottlenose dolphin (Tursiops spp.)

The conditioning of dolphins to human‐interaction behaviors has been documented in several areas worldwide. However, the metrics used to report human‐interaction behaviors vary among studies, making comparison across study areas difficult. The purpose of this study was to develop standard metrics for reporting human‐interaction behaviors and utilize these metrics to quantify the prevalence of human‐interaction behaviors by common bottlenose dolphins (Tursiops truncatus) near Savannah, Georgia. The four metrics used were percentage of days with human‐interaction behaviors, percentage of sightings with human‐interaction behaviors, percentage of the catalog that interacted with humans, and spatial extent of human‐interaction behaviors. Human‐interaction behaviors were observed on 69.6% of days and 23.5% of sightings near Savannah. In addition, 20.1% of the animals in the catalog were observed interacting with humans. These rates are much higher than those found in other areas with known issues with human‐interaction behaviors. These behaviors were observed across an area of 272.6 km², which is larger than other reported areas. The four metrics used in this study proved to be a valuable way to report human‐interaction behaviors, and their use is recommended for future studies to allow for comparison among areas.     

Isolation and characterization of novel microsatellite loci for the short‐beaked common dolphin (Delphinus delphis) and cross‐amplification in other cetacean species

Seven tetranucleotide and three dinucleotide microsatellite loci were isolated from the short‐beaked common dolphin (Delphinus delphis). Number of alleles and observed heterozygosity ranged from three to twelve (mean 8.4) and from 0.087 to 0.978 (mean 0.741), respectively. All loci were tested in four other odontocete species and showed polymorphisms in most locus–species combinations. This panel of nuclear markers will be useful for the investigation of population ecology and social interactions in common dolphins and other cetacean species.     

Isotopic niche width differentiation between common bottlenose dolphin ecotypes and sperm whales in the Gulf of California

World populations or stock distinction of Tursiops truncatus has been difficult to assess, because of large variations in morphology, habitat, feeding habits, and social structure among areas, which may reflect phylogenetic segregation or ecological plasticity. In the Gulf of California, Mexico, two common bottlenose dolphin ecotypes (inshore and offshore) have been reported. The offshore ecotype is frequently observed in association with sperm whales (Physeter macrocephalus) but the reason for this is still unknown. To explore the degree of resource partitioning/overlap between these species and stocks, we used skin stable isotope values (δ¹³C, δ¹⁵N) to estimate quantitative metrics of isotopic niche width (Bayesian standard ellipse areas, SEA_B) and estimated their diet composition using Bayesian isotopic mixing models. The inshore ecotype in different regions (north, central, and south) of the Gulf of California exhibited distinct δ¹⁵N values and SEA_B, suggesting a latitudinal gradient in nitrogen sources of coastal localities. The SEA_B of inshore and offshore bottlenose dolphin ecotypes was completely distinct, indicating resource partitioning. Associated offshore ecotype and sperm whales had overlapping SEA_B. The isotopic mixing model indicates that a considerable proportion of both species’ diet is large Humbolt squid. Our results suggest that resource partitioning and species association are two strategies that bottlenose dolphin ecotypes use in this zone.     

Decline in local abundance of bottlenose dolphins (Tursiops truncatus) in the Bay of Islands,New Zealand

Regional populations of bottlenose dolphins (Tursiops truncatus) around New Zealand are genetically isolated from each other and the species was recently classified as nationally endangered based on relatively small population sizes and reports of high calf mortality. Here, we estimate the abundance and trends in one of these regional populations, the Bay of Islands, using a photo‐identification database collected from 1997 to 1999 and from 2003 to 2006, containing a total of 3,841 records of 317 individual dolphins. Estimates of abundance obtained with the robust design fluctuated widely but showed a significant decline in the number of dolphins present in the bay over time (7.5% annual rate of decline). Temporary emigration was random and fluctuated considerably (γ  =  0.18, SE = 0.07 to γ  =  0.84, SE = 0.06). Apparent survival was estimated at 0.928 (CI = 0.911–0.942). Seasonal estimates (26 seasons) obtained in POPAN also showed a significant decline in abundance (5.8% annual rate of decline). Despite the decline observed in local abundance, dolphins continue to be found regularly in the Bay of Islands, suggesting that fewer dolphins use the bay on regular basis. Consequently, it seems that a change in habitat use, mortality and possibly low recruitment could underlie the apparent local decline.     

Likely future extirpation of another Asian river dolphin: The critically endangered population of the Irrawaddy dolphin in the Mekong River is small and declining

The population of Irrawaddy dolphins that occupies the Mekong River in southern Lao People's Democratic Republic and Cambodia is classified as Critically Endangered by the IUCN. Based on capture‐recapture of photo‐identified individuals, we estimated that the total population numbered 93 ±  SE 3.90 individuals (95% CI 86–101), as of April 2007. The combined photo‐identification and carcass recovery program undertaken from 2001 to 2007 established that the Irrawaddy dolphin population inhabiting the Mekong River has reached a critical point with regards to its continued survival, where immediate research and management actions are required to greatly reduce adult mortality, and establish the cause of newborn mortality. In addition, community consultation is required to initiate, and evaluate, urgently required conservation measures. An ongoing well‐designed combined program of abundance estimation (i.e., photo‐identification) and carcass recovery is required to monitor total population size and mortality rates, to inform and evaluate management initiatives. The conclusions of this paper are likely generic to river dolphin populations, particularly where photo‐identification is possible.     

Occurrence of false killer whales (Pseudorca crassidens) and their association with common bottlenose dolphins (Tursiops truncatus) off northeastern New Zealand

On a global scale, false killer whales (Pseudorca crassidens) remain one of the lesser‐known delphinids. The occurrence, site fidelity, association patterns, and presence/absence of foraging in waters off northeastern New Zealand are examined from records collected between 1995 and 2012. The species was rarely encountered; however, of the 61 distinctive, photo‐identified individuals, 88.5% were resighted, with resightings up to 7 yr after initial identification, and movements as far as 650 km documented. Group sizes ranged from 20 to ca. 150. Results indicate that all individuals are linked in a single social network. Most observations were recorded in shallow (<100 m) nearshore waters. Occurrence in these continental shelf waters is likely seasonal, coinciding with the shoreward flooding of a warm current. During 91.5% of encounters, close interspecific associations with common bottlenose dolphins (Tursiops truncatus) were observed. Photo‐identification reveals repeat inter‐ and intraspecific associations among individuals with 34.2% of common bottlenose dolphins resighted together with false killer whales over 1,832 d. While foraging was observed during 39.5% of mixed‐species encounters, results suggest that social and antipredatory factors may also play a role in the formation of these mixed‐species groups.     

Ontogenetic allometry and body composition of the common bottlenose dolphin (Tursiops truncatus) from the U.S. mid‐Atlantic

Growth in common bottlenose dolphins (Tursiops truncatus) was investigated through examination of sex‐specific, ontogenetic changes in the mass of 38 discrete body compartments, utilizing stranded dolphins in good body condition (n = 145). Ontogenetic allometry and the body composition technique were used to quantitatively describe growth patterns. Although adult males were significantly larger than adult females in total body mass (TBM) and total length, overall patterns of growth were remarkably similar between sexes. The integument, locomotor muscle, and vertebral column together represented 50%–58% of TBM across all life history categories, although their relative contributions varied ontogenetically. Young dolphins invested the greatest percentage of TBM in integument, while locomotor muscle was the single largest body component in adults. In both sexes (1) most muscle groups displayed positive allometry, (2) most skeletal elements displayed negative allometric or isometric growth, (3) most abdominal viscera associated with digestion displayed positive allometry, and (4) the brain displayed negative allometric growth. Reproductive tissues exhibited the highest rates of growth in both sexes, and increased as a percentage of TBM with maturity. This study provides an integrated view of bottlenose dolphin growth and a quantitative baseline of body composition for future monitoring of this sentinel species of ecosystem health.     

Fission‐fusion dynamics of Guiana dolphin (Sotalia guianensis) groups at Pipa Bay,Rio Grande do Norte,Brazil

Fission‐fusion dynamics seem to reflect individual decisions as well as temporal and spatial variations in the organization of groups of the same species. To understand the group dynamics of the Guiana dolphin, Sotalia guianensis, at Pipa Bay, Brazil, we investigated the three dimensions of a fission‐fusion social system: (1) variation in spatial cohesion, (2) variation in party size, and (3) variation in party composition. Sampling took place from December 2007 to February 2009 over 176 d and we analyzed the behavioral patterns of 658 groups. Within subgroups, animals remained cohesive, particularly in groups of adults and calves. Greater cohesion was also observed during resting and fission‐fusion rates were higher during milling and feeding. Groups composed of adults and juveniles showed a higher dynamics index (group size variation as a function of time) than groups composed only of adults and the fission‐fusion rate was higher during dry periods. Guiana dolphin groups frequently changed their group size and composition every 20 min on average. Taking these factors into consideration, we suggest that the Guiana dolphin demonstrates fission‐fusion dynamics, a pattern of behavior similar to what has been observed in other coastal odontocete species, such as Tursiops spp. and Lagenorhynchus obscurus.     

Blood mercury concentrations in common bottlenose dolphins from the Indian River Lagoon,Florida: Patterns of social distribution

Social network analysis has been shown to be effective in studying the social structure of cetacean populations. Common bottlenose dolphins (Tursiops truncatus) inhabiting the Indian River Lagoon (IRL), Florida, have among the highest concentrations of total mercury (THg) in blood reported worldwide. The purpose of this study was to examine the relationship between THg concentrations in IRL dolphins and their social affiliations. Whole blood samples from 98 dolphins with photo‐identification sighting histories were collected between 2003–2007 and 2010–2012. Dolphins were categorized into approximate tertiles of low (mean 199.7 μg/L), medium (mean 366.8 μg/L), and high (mean 990.5 μg/L) THg exposure. Social associations between individual dolphins were defined by the proportion of sightings documented with another known individual. Social network measures of individuals and associations between dyads were examined to determine differences among THg categories. Strong social affiliations of individuals within the highest category of THg were found (P = 0.04), suggesting shared exposures among dolphins foraging in specific areas of the estuary. Network measures of strength and affinity were significantly higher in the highest exposure category. This report used social network analysis as a novel way to examine patterns of exposure to an environmental contaminant in a cetacean population.     

Bottlenose dolphin (Tursiops truncatus) social structure in the Shannon Estuary,Ireland, is distinguished by age‐ and area‐related associations

Social and network analyses that incorporate information on individuals within a population enhance our knowledge of complex species. In this study, the social structure of wild bottlenose dolphins in the Shannon Estuary, Ireland, was analyzed by examining the dynamics of the whole population and then of individuals classed by sex, age, and area. One hundred and twenty‐one dolphins were identified during 522 sightings between 2012 and 2015. The mean half‐weight association index (HWI) of the population was 0.07 ± 0.02. The highest HWIs for individuals of known sex were for female‐male pairs. Mean within‐class HWI was significantly higher than mean between‐class HWI for both age and area classes. Ordinations and sociograms were used to visualize social networks. Permutation tests revealed nonrandom associations for the population overall and both between and within classes. Temporal analyses showed associations persisting for >1,000 d. The whole population's best fit model was for two levels of casual acquaintances. Movement analyses demonstrated the use of the inner estuary by only 25% of the population revealing a potential community division by area. The difference between mean HWI when socializing (0.09 ± 0.03) compared to foraging (0.06 ± 0.03) was significant. These results highlight the importance of localized research, reflecting the complexity found in bottlenose dolphin societies globally.     

Characterization of the cyanobacteria and associated bacterial community from an ephemeral wetland in New Zealand

New Zealand ephemeral wetlands are ecologically important, containing up to 12% of threatened native plant species and frequently exhibiting conspicuous cyanobacterial growth. In such environments, cyanobacteria and associated heterotrophs can influence primary production and nutrient cycling. Wetland communities, including bacteria, can be altered by increased nitrate and phosphate due to agricultural practices. We have characterized cyanobacteria from the Wairepo Kettleholes Conservation Area and their associated bacteria. Use of 16S rRNA amplicon sequencing identified several operational taxonomic units (OTUs) representing filamentous heterocystous and non‐heterocystous cyanobacterial taxa. One Nostoc OTU that formed macroscopic colonies dominated the cyanobacterial community. A diverse bacterial community was associated with the Nostoc colonies, including a core microbiome of 39 OTUs. Identity of the core microbiome associated with macroscopic Nostoc colonies was not changed by the addition of nutrients. One OTU was highly represented in all Nostoc colonies (27.6%–42.6% of reads) and phylogenetic analyses identified this OTU as belonging to the genus Sphingomonas. Scanning electron microscopy showed the absence of heterotrophic bacteria within the Nostoc colony but revealed a diverse community associated with the colonies on the external surface.     

Multi‐methods approach to characterizing the magnitude,impact, and spatial risk of Irrawaddy dolphin (Orcaella brevirostris) bycatch in small‐scale fisheries in Malampaya Sound,Philippines

Addressing the urgent conservation threat of marine mammal bycatch in small‐scale fisheries requires information on bycatch magnitude, gear types, population impacts, and risk. However, data on these proximate attributes are widely lacking in developing countries. This study used a multi‐methods approach integrating boat surveys and interviews with fishermen (n = 526) to assess proximate attributes of bycatch for the Critically Endangered subpopulation of Irrawaddy dolphins (Orcaella brevirostris) in Malampaya Sound, Philippines. Given an updated population estimate of 35 individuals (CV = 22.9%), the estimated rate of bycatch fatalities exceeds the potential biological removal rate. Spatial overlap scores were calculated to characterize overlap between dolphins and fisheries as a proxy of bycatch risk. These scores identified particularly high risk areas that could be prioritized for gear bans. However, completely reducing bycatch risk would require more widespread bans beyond these high risk areas. Implementing gear restrictions will be immensely challenging, given serious obstacles to fisheries management at this site. The multi‐methods approach describes the urgency of the bycatch problem and the changes required for mitigation, setting the stage for identifying and evaluating potential solutions. It can be readily applied to developing country sites to guide more efficient and complete data collection and conservation efforts.     

Dental anomaly in a fossil squalodont dolphin from New Zealand,and the evolution of polydonty in whales

Abstract

A fragment of mandible of an indeterminate squalodontid dolphin (Upper Oligocene or lowermost Miocene, New Zealand) has 2 anomalous single-rooted teeth intercalated between typically squalodontid anterior cheek-teeth. The anomalous teeth are considered to be truly supernumerary, and not homologous with any erupted teeth in phylogenetically earlier Cetacea. This type of anomaly appears not to have been reported previously for any fossil cetacean. Polydonty, a characteristic of extant odontocetes, was probably attained initially in primitive odontocetes by intercalation of permanent and deciduous teeth, and later was elaborated by the addition of supernumerary teeth. The present specimen represents a morphological (but not phylogenetic) stage between odontocetes with limited and advanced polydonty. Some fossil mysticetes and embryonic extant mysticetes are also polydont, but mysticete polydonty and that of odontocetes have probably evolved convergently.