Population structure of pygmy (Kogia breviceps) and dwarf (Kogia sima) sperm whales in the Southern Hemisphere may reflect foraging ecology and dispersal patterns.

Little is known about the biology of pygmy (Kogia breviceps) and dwarf (K. sima) sperm whales as these animals are difficult to observe in the wild. However, both species strand frequently along the South African, Australian and New Zealand coastlines, providing samples for these otherwise inaccessible species. The use of DNA samples from tissue and DNA extracted from historical material, such as teeth and bone, allowed a first analysis of the population structure of both species in the Southern Hemisphere. A 279 base pair consensus region of the mitochondrial cytochrome b gene was sequenced for 96 K. breviceps (53 tissue and 43 teeth or bone samples) and 29 K. sima (3 tissue and 26 teeth or bone samples), and 26 and 12 unique haplotypes were identified, respectively. K. breviceps showed a higher nucleotide diversity of 0.82% compared to 0.40% in K. sima. Significant genetic differentiation was detected in the Southern Hemisphere between K. breviceps from South Africa and New Zealand (ФST = 0.042, p < 0.05). Mitochondrial control region sequences (505 bp) were available for 44 individuals (41 K. breviceps and 3 K. sima) for comparative purposes. A comprehensive global phylogenetic analysis (maternal lineage) of our sequences together with all available Kogia mtDNA sequences largely supported previously published phylogenetic findings, but highlighted some changed inferences about oceanic divergences within both species. The higher nucleotide diversity and low population differentiation observed in K. breviceps may result from its broad foraging ecology and wide distribution, which may indicate a more opportunistic feeding behaviour and tolerance towards a larger range of water temperatures than K. sima.

Inter- and intra-specific trophic interactions of coastal delphinids off the eastern coast of South Africa inferred from stable isotope analysis.

Dietary tracers, such as bulk stable carbon (δ13C) and nitrogen (δ15N) isotopes, can be used to investigate the trophic interactions of marine predators, which is useful to assess their ecological roles within communities. These tracers have also been used to elucidate population structure and substructure, which is critical for the better identification of management units for these species affected by a range of threats, particularly bycatch in fishing gears. Off eastern South Africa, large populations of Indo-Pacific bottlenose (Tursiops aduncus) and common dolphins (Delphinus delphis) co-occur and are thought to follow the pulses of shoaling sardines (Sardinops sagax) heading north-east in the austral winter. Here we used δ13C and δ15N to investigate the trophic interactions and define ecological units of these two species along a ≈800 km stretch of the east coast of South Africa, from Algoa Bay to the coast of KwaZulu-Natal. Common and bottlenose dolphin dietary niche overlapped by 39.7% overall in our study area, with the highest overlap occurring off the Wild Coast (40.7% at Hluleka). Both stable isotopes were significantly enriched in bottlenose dolphins sampled in the western part of our study area (i.e., Algoa Bay and Amathole) compared to eastern animals (i.e., from Hluleka, Pondoland, and KZN). In areas where genetic information is not available or is insufficient, food web tracers (such as stable isotopes) can be used to group individuals based on trophic ecology, which can provide ecological units for management of populations. The distinct isotope signatures found here for bottlenose dolphins can, therefore, be used as management units for conservation efforts in the future.

Histomorphological stratification of blubber of three dolphin species from subtropical waters.

Blubber is a highly specialized and dynamic tissue unique to marine mammals and presents a reflection of the individuals' nutrition, environment, and life history traits. Few studies have investigated the histomorphology of cetacean blubber in subtropical environments. The aim of this study was to investigate the blubber histomorphology of three different dolphin species off the subtropical KwaZulu-Natal coast, South Africa, using adipocyte cell size, number, and density. Blubber tissue samples from the saddle area of 43 incidentally bycaught animals (four Sousa plumbea, 36 Tursiops aduncus, and three Delphinus delphis) were used to compare cell parameters between blubber layers. Samples were divided into the upper third (corresponding to the superficial layer closest to the epidermis), middle third, and lower third (corresponding to the deep layer). For T. aduncus, factors potentially affecting blubber histomorphology, such as sex, age class, and season, were also assessed. Our results showed that no stratification was present in S. plumbea, which could be ascribed to the species' warmer inshore habitat, large body size, and apparent lower mobility. For T. aduncus and D. capensis, however, blubber stratification was determined, characterized by a gradual transition of cell size, number, and density between layers rather than clearly defined layers. Significant differences in adipocyte cell number and density were found for different sexes and age classes of T. aduncus. However, there were no significant differences between seasons, which was attributed to the small temperature differences between seasons. This study represents the first investigation of odontocete blubber histomorphology in subtropical waters. It is recommended that future studies investigate blubber lipid content, while also taking into consideration the reproductive status of the females and the temperature range of their study area. It is hoped that our results, in conjunction with histopathology and other health indicators, could assist in assessing health and body condition.

A drastic shift in the energetic landscape of toothed whale sperm cells.

Mammalian spermatozoa are a notable example of metabolic compartmentalization.1 Energy in the form of ATP production, vital for motility, capacitation, and fertilization, is subcellularly separated in sperm cells. While glycolysis provides a local, rapid, and low-yielding input of ATP along the flagellum fibrous sheath, oxidative phosphorylation (OXPHOS), far more efficient over a longer time frame, is concentrated in the midpiece mitochondria.2 The relative weight of glycolysis and OXPHOS pathways in sperm function is variable among species and sensitive to oxygen and substrate availability.3-5 Besides partitioning energy production, sperm cell energetics display an additional singularity: the occurrence of sperm-specific gene duplicates and alternative spliced variants, with conserved function but structurally bound to the flagellar fibrous sheath.6,7 The wider selective forces driving the compartmentalization and adaptability of this energy system in mammalian species remain largely unknown, much like the impact of ecosystem resource availability (e.g., carbohydrates, fatty acids, and proteins) and dietary adaptations in reproductive physiology traits.8 Here, we investigated the Cetacea, an iconic group of fully aquatic and carnivorous marine mammals, evolutionarily related to extant terrestrial herbivores.9 In this lineage, episodes of profound trait remodeling have been accompanied by clear genomic signatures.10-14 We show that toothed whales exhibit impaired sperm glycolysis, due to gene and exon erosion, and demonstrate that dolphin spermatozoa motility depends on endogenous fatty acid ß-oxidation, but not carbohydrates. Such unique energetic rewiring substantiates the observation of large mitochondria in toothed whale spermatozoa and emphasizes the radical physiological reorganization imposed by the transition to a carbohydrate-depleted marine environment.

Head adaptation for sound production and feeding strategy in dolphins (Odontoceti: Delphinida).

Head morphology in toothed whales evolved under selective pressures on feeding strategy and sound production. The postnatal development of the skull (n = 207) and mandible (n = 219) of six Delphinida species which differ in feeding strategy but exhibit similar sound emission patterns, including two narrow-band high-frequency species, were investigated through 3D morphometrics. Morphological changes throughout ontogeny were demonstrated based on the main source of variation (i.e., prediction lines) and the common allometric component. Multivariate trajectory analysis with pairwise comparisons between all species was performed to evaluate specific differences on the postnatal development of skulls and mandibles. Changes in the rostrum formation contributed to the variation (skull: 49%; mandible: 90%) of the entire data set and might not only reflect the feeding strategy adopted by each lineage but also represents an adaptation for sound production and reception. As an important structure for directionality of sound emissions, this may increase directionality in raptorial feeders. Phylogenetic generalized least squares analyses indicated that shape of the anterior portion of the skull is strongly dependent on phylogeny and might not only reflect feeding mode, but also morphological adaptations for sound production, particularly in raptorial species. Thus, postnatal development seems to represent a crucial stage for biosonar maturation in some raptorial species such as Pontoporia blainvillei and Sousa plumbea. The ontogeny of their main tool for navigation and hunting might reflect their natural history peculiarities and thus potentially define their main vulnerabilities to anthropogenic changes in the environment.

Cranial crassicaudiasis in two coastal dolphin species from South Africa is predominantly a disease of immature individuals.

Crassicauda spp. (Nematoda) infest the cranial sinuses of several odontocetes, causing diagnostic trabecular osteolytic lesions. We examined skulls of 77 Indian Ocean humpback dolphins Sousa plumbea and 69 Indo-Pacific bottlenose dolphins Tursiops aduncus, caught in bather-protecting nets off KwaZulu-Natal (KZN) from 1970-2017, and skulls of 6 S. plumbea stranded along the southern Cape coast in South Africa from 1963-2002. Prevalence of cranial crassicaudiasis was evaluated according to sex and cranial maturity. Overall, prevalence in S. plumbea and T. aduncus taken off KZN was 13 and 31.9%, respectively. Parasitosis variably affected 1 or more cranial bones (frontal, pterygoid, maxillary and sphenoid). No significant difference was found by gender for either species, allowing sexes to be pooled. However, there was a significant difference in lesion prevalence by age, with immature T. aduncus 4.6 times more likely affected than adults, while for S. plumbea, the difference was 6.5-fold. As severe osteolytic lesions are unlikely to heal without trace, we propose that infection is more likely to have a fatal outcome for immature dolphins, possibly because of incomplete bone development, lower immune competence in clearing parasites or an over-exuberant inflammatory response in concert with parasitic enzymatic erosion. Cranial osteolysis was not observed in mature males (18 S. plumbea, 21 T. aduncus), suggesting potential cohort-linked immune-mediated resistance to infestation. Crassicauda spp. may play a role in the natural mortality of S. plumbea and T. aduncus, but the pathogenesis and population level impact remain unknown.

Phylogeography of the dugong (Dugong dugon) based on historical samples identifies vulnerable Indian Ocean populations.

We investigated the phylogeography of the dugong (Dugong dugon) across its original range using museum material from 14 natural history museum and university collections. The mitochondrial DNA control region was successfully amplified from samples of bone or tooth powder from 162 individuals. These samples range from 1827 to 1996 and span the historical distribution range of the dugong. We were able to successfully amplify overlapping fragments of the D-loop region of the mitochondrial DNA (mtDNA) resulting in sequences of a 355 bp fragment for 162 individuals for the final analyses. This included a new sequence (189 bp) from a previously unidentified piece of skin of the extinct Steller's sea cow (Hydrodamalis gigas), as an outgroup. The resulting dugong sequences match those from previous studies of dugongs from Australia and Indonesia, but revealed several new and divergent mtDNA lineages in the Indian Ocean. One mtDNA lineage includes most specimens from the Western Indian Ocean, with another distinct lineage isolated to nearby Madagascar and Comores. There is little geographic structuring detectable among other populations in the Western Indian Ocean and all populations from that region appear to have historically contained comparatively low levels of genetic diversity. The genetic diversity of several Indian Ocean samples collected after 1950 was lower than that of the samples collected earlier from similar locations, a result coincident with the anecdotal reductions in population size. The new lineages and potential loss of diversity highlight the particular conservation importance and vulnerability of dugong populations in the Western Indian Ocean.

Halogenated and organophosphorus flame retardants in cetaceans from the southwestern Indian Ocean.

PBDEs, HBCD, DBDPE, PBEB and HBB, dechloranes and OPFRs, as well as natural MeO-PBDEs were monitored in muscle tissue of three dolphin species from the southwestern Indian Ocean (Delphinus delphis, Sousa plumbea and Tursiops aduncus) collected between 2012 and 2015. The mean PBDE concentration was 416 ±â€¯333 ng g-1 lw. BDE-47 was found in all samples and was almost half the total PBDE contamination. BDE-209, BDE-100 and BDE-99 were present in ≥85% of the samples. HBCD was detected in just two samples at 20 and 330 ng g-1 lw. PBEB and HBB were not detected, while DBDPE was in all samples but always below its limit of quantification. Dec 602 was the only quantifiable dechlorane at 232 ±â€¯549 ng g-1 lw. Mean OPFR concentration was 10452 ±â€¯11301 ng g-1 lw. TBOEP was found in all samples making up most of the total OPFR contamination. MeO-PBDEs were detected in all samples at 114 ±â€¯137 ng g-1 lw. Data on flame retardants in biota and environmental samples from the southwestern Indian Ocean are scarce and, as a result, comparisons are difficult. However, data from other marine predators in the region, such as penguins, suggest that further studies are needed to determine if these concentrations are the consequence of a high local contamination or widespread thoughout the Indian Ocean.

Sound Generating Structures of the Humpback Dolphin Sousa plumbea (Cuvier, 1829) and the Directionality in Dolphin Sounds.

The macroscopic morphology of structures involved in sound generation in the Indian Ocean humpback dolphin (Sousa plumbea) were described for the first time using computed tomography imaging and standard gross dissection techniques. The Indian Ocean humpback dolphin may represent a useful comparative model to the bottlenose dolphin (Tursiops sp.) to provide insights into the functional anatomy of the sound production in dolphins, since these coastal dolphins exhibit similar body size and share similarities on acoustic behavior. The general arrangement of sound generating structures, that is, air sacs and muscles, was similar in both the bottlenose dolphin and the Indian Ocean humpback dolphin. The main difference between the two species existed in a small left posterior branch of the melon in the Indian Ocean humpback dolphin, which was not found in the bottlenose dolphin and might reflect an adaptation of directionality for high frequency communication sounds as seen in some other delphinids (e.g., Lagenorhynchus sp., Grampus griseus). Thus, this may be the main reason for the asymmetry of the sound production structures in dolphins. Additionally, the longer rostrum in Indian Ocean humpback dolphins might suggest a more directional echolocation beam compared to the Lahille's bottlenose dolphin. Anat Rec, 302:849-860, 2019. © 2018 Wiley Periodicals, Inc.

Profiling and Spatial Variation Analysis of Persistent Organic Pollutants in South African Delphinids.

The continuous disposal of persistent organic pollutants (POPs) in South Africa (SA) warrants concern about their detrimental effects on humans and wildlife. We surveyed six dolphin species (n = 90) incidentally captured in shark net installations or stranded off the SA east and south coast from 2005 to 2009 to study the POP exposure. Sousa plumbea, an inshore and estuarine species, was found to be the most contaminated by total POPs (21 100 ng g(-1) lw) of all the dolphins off SA, followed by Tursiops aduncus (19 800 ng g(-1) lw), Lagenodelphis hosei (13 600 ng g(-1) lw), and Delphinus capensis (5500 ng g(-1) lw), whereas POP levels in the offshore or pelagic delphinids were much lower. In all delphinids, dominant pollutants were dichlorodiphenyltrichloroethanes (DDTs), which represented more than 60% of the total concentration of total POPs, followed by polychlorinated biphenyls (PCBs, 30%). Concentrations of DDTs in S. plumbea and T. aduncus off SA were among the highest levels reported in delphinids globally. Approximately half of the adult T. aduncus had PCB concentrations above the effect threshold for impairment of immune functions. The concentrations of Mirex and Dieldrin in SA delphinids were higher than those found in species from other regions of the Southern Hemisphere.

The Natural History and Conservation of Indian Ocean Humpback Dolphins (Sousa plumbea) in South African Waters.

Although most knowledge on the biology of Sousa plumbea has primarily come from South African waters, a number of research gaps remain on the natural history and status of the species in the region. Research on two populations in South African waters for which some historical data exist may aid in highlighting long-term changes in the biology and natural history of this little known coastal delphinid. Recent studies on the age, growth and reproduction of animals incidentally caught in shark nets in Richards Bay, KwaZulu-Natal, yielded a lower maximum age estimate of 24 (previously 46) growth-layer-groups (GLGs), sexual maturity of 7.5 and 8 GLGs in males and females (previously 12-13 and 10 GLGs, respectively), an ovulation rate of 0.2 and a 5-year calving interval (previously 0.3 and 3-year calving interval) than previously reported. These differences may be due to a difference in the interpretation of GLGs between observers or a predominance of young males being caught in the shark nets. Stomach content analysis revealed a change in the relative proportions of the main prey items over the past 25 years, but no difference in species richness or diversity was found between the sexes. No change in trophic level was recorded between 1972 and 2009. Field studies in Algoa Bay, Eastern Cape, conducted 16 years apart indicated a decline in the mean group size (from 7 to 3 animals), a decline in the maximum group size (from 24 to 13 animals), an increase in solitary individuals (15.4-36%), and a change in behaviour from predominantly foraging (64-18%) to mainly travelling (24-49%). The observed changes are suggestive of a change in food availability, resulting in a range shift or a potential decline in numbers. These studies indicate the importance of long-term studies to monitor population changes and their possible causes. A number of threats, such as shark nets, pollution (noise and chemical), and coastal development and disturbance, to the humpback dolphin populations in South Africa have been identified. Urgent action is required to ensure continued existence of the species in South African waters.

A systematic health assessment of indian ocean bottlenose (Tursiops aduncus) and indo-pacific humpback (Sousa plumbea) dolphins incidentally caught in shark nets off the KwaZulu-Natal Coast, South Africa.

Coastal dolphins are regarded as indicators of changes in coastal marine ecosystem health that could impact humans utilizing the marine environment for food or recreation. Necropsy and histology examinations were performed on 35 Indian Ocean bottlenose dolphins (Tursiops aduncus) and five Indo-Pacific humpback dolphins (Sousa plumbea) incidentally caught in shark nets off the KwaZulu-Natal coast, South Africa, between 2010 and 2012. Parasitic lesions included pneumonia (85%), abdominal and thoracic serositis (75%), gastroenteritis (70%), hepatitis (62%), and endometritis (42%). Parasitic species identified were Halocercus sp. (lung), Crassicauda sp. (skeletal muscle) and Xenobalanus globicipitis (skin). Additional findings included bronchiolar epithelial mineralisation (83%), splenic filamentous tags (45%), non-suppurative meningoencephalitis (39%), and myocardial fibrosis (26%). No immunohistochemically positive reaction was present in lesions suggestive of dolphin morbillivirus, Toxoplasma gondii and Brucella spp. The first confirmed cases of lobomycosis and sarcocystosis in South African dolphins were documented. Most lesions were mild, and all animals were considered to be in good nutritional condition, based on blubber thickness and muscle mass. Apparent temporal changes in parasitic disease prevalence may indicate a change in the host/parasite interface. This study provided valuable baseline information on conditions affecting coastal dolphin populations in South Africa and, to our knowledge, constitutes the first reported systematic health assessment in incidentally caught dolphins in the Southern Hemisphere. Further research on temporal disease trends as well as disease pathophysiology and anthropogenic factors affecting these populations is needed.

Killer whale nuclear genome and mtDNA reveal widespread population bottleneck during the last glacial maximum.

Ecosystem function and resilience is determined by the interactions and independent contributions of individual species. Apex predators play a disproportionately determinant role through their influence and dependence on the dynamics of prey species. Their demographic fluctuations are thus likely to reflect changes in their respective ecological communities and habitat. Here, we investigate the historical population dynamics of the killer whale based on draft nuclear genome data for the Northern Hemisphere and mtDNA data worldwide. We infer a relatively stable population size throughout most of the Pleistocene, followed by an order of magnitude decline and bottleneck during the Weichselian glacial period. Global mtDNA data indicate that while most populations declined, at least one population retained diversity in a stable, productive ecosystem off southern Africa. We conclude that environmental changes during the last glacial period promoted the decline of a top ocean predator, that these events contributed to the pattern of diversity among extant populations, and that the relatively high diversity of a population currently in productive, stable habitat off South Africa suggests a role for ocean productivity in the widespread decline.