Rui Prieto

High gene flow in oceanic bottlenose dolphins ( Tursiops truncatus ) of the North Atlantic

Despite the openness of the oceanic environment, limited dispersal and tight social structure often induce genetic structuring in marine organisms, even in large animals such as cetaceans. In the bottlenose dolphin, mitochondrial and nuclear DNA analyses have revealed the existence of genetic differentiation between pelagic (or offshore) and coastal (or nearshore) ecotypes in the western North Atlantic, as well as between coastal populations. Because previous studies concentrated on continental margins, we analysed the population structure of bottlenose dolphins in two of the most isolated archipelagos of the North Atlantic: the Azores and Madeira. We analysed 112 samples collected on live animals in the two archipelagos, and nine samples collected on stranded animals in Madeira and mainland Portugal. Genetic analyses consisted in molecular sexing, sequencing of part of the mitochondrial hyper-variable region, and screening of ten microsatellite loci. We predicted that: (1) there is at least one pelagic and one or more coastal populations in each archipelago; (2) populations are differentiated between and possibly within archipelagos. Contrary to these predictions, results indicated a lack of population structure in the study area. In addition, comparison with published sequences revealed that the samples from the Azores and Madeira were not significantly differentiated from samples of the pelagic population of the western North Atlantic. Thus, bottlenose dolphins occurring in the pelagic waters of the North Atlantic belong to a large oceanic population, which should be regarded as a single conservation unit. Unlike what is known for coastal populations, oceanic bottlenose dolphins are able to maintain high levels of gene flow.

North Atlantic blue and fin whales suspend their spring migration to forage in middle latitudes: building up energy reserves for the journey?

The need to balance energy reserves during migration is a critical factor for most long-distance migrants and an important determinant of migratory strategies in birds, insects and land mammals. Large baleen whales migrate annually between foraging and breeding sites, crossing vast ocean areas where food is seldom abundant. How whales respond to the demands and constraints of such long migrations remains unknown. We applied a behaviour discriminating hierarchical state-space model to the satellite tracking data of 12 fin whales and 3 blue whales tagged off the Azores, to investigate their movements, behaviour (transiting and area-restricted search, ARS) and daily activity cycles during the spring migration. Fin and blue whales remained at middle latitudes for prolonged periods, spending most of their time there in ARS behaviour. While near the Azores, fin whale ARS behaviour occurred within a restricted area, with a high degree of overlap among whales. There were noticeable behavioural differences along the migratory pathway of fin whales tracked to higher latitudes: ARS occurred only in the Azores and north of 56°N, whereas in between these areas whales travelled at higher overall speeds while maintaining a nearly direct trajectory. This suggests fin whales may alternate periods of active migration with periods of extended use of specific habitats along the migratory route. ARS behaviour in blue whales occurred over a much wider area as whales slowly progressed northwards. The tracks of these whales terminated still at middle latitudes, before any behavioural switch was detected. Fin whales exhibited behavioural-specific diel rhythms in swimming speed but these varied significantly between geographic areas, possibly due to differences in the day-night cycle across areas. Finally, we show a link between fin whales seen in the Azores and those summering in eastern Greenland-western Iceland along a migratory corridor located in central Atlantic waters.

Occurrence and distribution of cetaceans in the waters around the Azores (Portugal), Summer and Autumn 1999- 2000

Boat-based surveys were conducted during the summer and autumn months of 1999 and 2000 around the islands of the Archipelago of the Azores to determine occurrence, distribution, and relative abundance of cetaceans. A total of 222 cetacean schools, corresponding to 11 species, were sighted over the two years. Cetaceans were widely distributed in the area surveyed, but the central group of islands recorded the highest relative abundance of cetaceans and the greater species diversity. Within each group of islands, cetaceans were more abundant in the coastal area (to 9 km from shore) than in the oVshore area (9 to 28 km). Atlantic spotted dolphins (Stenella frontalis ), short-beaked common dolphins ( Delphinus delphis ), and bottlenose dolphins (Tursiops truncatus ) were the most frequently sighted species and were found in all the islands surveyed. Although there was considerable overlap in spatial distribution among these species, common dolphins and bottlenose dolphins occurred in coastal areas, while Atlantic spotted dolphins were more common in o Vshore and deeper waters.

Assessing performance of Bayesian state-space models fit to Argos Satellite telemetry locations processed with Kalman Filtering

Argos recently implemented a new algorithm to calculate locations of satellite-tracked animals that uses a Kalman filter (KF). The KF algorithm is reported to increase the number and accuracy of estimated positions over the traditional Least Squares (LS) algorithm, with potential advantages to the application of state-space methods to model animal movement data. We tested the performance of two Bayesian state-space models (SSMs) fitted to satellite tracking data processed with KF algorithm. Tracks from 7 harbour seals (Phoca vitulina) tagged with ARGOS satellite transmitters equipped with Fastloc GPS loggers were used to calculate the error of locations estimated from SSMs fitted to KF and LS data, by comparing those to “true” GPS locations. Data on 6 fin whales (Balaenoptera physalus) were used to investigate consistency in movement parameters, location and behavioural states estimated by switching state-space models (SSSM) fitted to data derived from KF and LS methods. The model fit to KF locations improved the accuracy of seal trips by 27% over the LS model. 82% of locations predicted from the KF model and 73% of locations from the LS model were <5 km from the corresponding interpolated GPS position. Uncertainty in KF model estimates (5.6±5.6 km) was nearly half that of LS estimates (11.6±8.4 km). Accuracy of KF and LS modelled locations was sensitive to precision but not to observation frequency or temporal resolution of raw Argos data. On average, 88% of whale locations estimated by KF models fell within the 95% probability ellipse of paired locations from LS models. Precision of KF locations for whales was generally higher. Whales’ behavioural mode inferred by KF models matched the classification from LS models in 94% of the cases. State-space models fit to KF data can improve spatial accuracy of location estimates over LS models and produce equally reliable behavioural estimates.

Spatial and temporal distribution of cetaceans in the mid-Atlantic waters around the Azores

Abstract Cetaceans living in offshore waters are under increasing pressure from anthropogenic activities. Yet, due to the lack of survey effort, relatively little is known about the demography or ecology of these populations. Spatial and temporal distribution of cetaceans in mid-Atlantic waters were investigated using a long term dataset collected from boat surveys and land-based observations around the Azores. From 1999 to 2009, 7307 cetacean schools were sighted during 271,717 km of survey effort. In 4944 h of land-based observations, 2,968 cetacean groups were detected. Twenty-four species were recorded: seven baleen whales, six beaked whales, eight dolphin species, Physeter macrocephalus, Kogia breviceps and Kogia sima. Overall, Delphinus delphis was the most frequently sighted species, but its encounter rate decreased in June–November, coinciding with the presence of Stenella frontalis in the region. Tursiops truncatus, P. macrocephalus and Grampus griseus were frequently encountered year-round, whereas large baleen whales showed a distinct peak in encounter rates in March–May. Mesoplodonts were fairly common and appear to be present throughout the year. These findings fill in a significant gap in the knowledge of cetaceans occurring in a poorly studied region of the North Atlantic, providing much needed data to inform management initiatives.

Geographic variability in the acoustic parameters of striped dolphin's (Stenella coeruleoalba) whistles

Geographic variation in the acoustic features of whistles emitted by the striped dolphin (Stenella coeruleoalba) from the Atlantic Ocean (Azores and Canary Islands) and the Mediterranean was investigated. Ten parameters (signal duration, beginning, end, minimum and maximum frequency, the number of inflection points, of steps, of minima and maxima in the contour and the frequency range) were extracted from each whistle. Discriminant function analysis correctly classified 73% of sounds between Atlantic Ocean and Mediterranean Sea. A cline in parameters was apparent from the Azores to the Mediterranean, with a major difference between the Canaries and the Mediterranean than between Azores and Canaries. Signal duration, maximum frequency, and frequency range measured in the Mediterranean sample were significantly lower compared to those measured in the Atlantic. Modulation parameters played a considerable role in area discrimination and were the only parameters contributing to highlight the differences within the Atlantic Ocean. Results suggest that the acoustic features constrained by structural phenotype, such as whistles frequency parameters, have a major effect on the Atlantic and Mediterranean separation while behavioral context, social, and physical environment may be among the main factors contributing to local distinctiveness of Atlantic areas. These results have potential passive acoustic monitoring applications.

Sex bias in biopsy samples collected from free-ranging dolphins

Biological samples of free-ranging dolphins are increasingly used to gain information on population structure and ecology. In small cetaceans, the gender of individuals usually cannot be determined at sea, and population sex ratio has to be inferred indirectly. We used molecular sexing to determine the gender of 340 biopsy samples of bottlenose dolphins, Tursiops truncatus, spotted dolphins, Stenella frontalis, and common dolphins, Delphinus delphis, collected around the Azores and Madeira. Sex ratio was globally skewed in favor of males, and differed between species and archipelagos. Skew was probably influenced by the selectivity of biopsy collectors and seasonal or year-round predominance of males in natural populations. Skew was also influenced by sampling duration and intensity. In the Azores, when several samples were successively collected within the same group, the proportion of female samples decreased as a function of sample order. This trend indicated a tendency for females to increasingly avoid the boat while samples were being collected. It showed that males and females reacted differently to the perturbation caused by the biopsy sampling process (i.e., sample collection and driving style).

Population genetics and social organization of the sperm whale (Physeter macrocephalus) in the Azores inferred by microsatellite analyses

In the northeast Atlantic Ocean, the archipelago of the Azores is frequented by female–offspring groups of sperm whales (Physeter macrocephalus L., 1758), as well as large males. The Azores apparently constitute both a feeding ground and a reproduction site. Little is known about the population and group structure of sperm whales in the area. We analysed 151 sloughed skin and biopsy samples collected from 2002 to 2004. Molecular analyses involved genetic tagging using 11 microsatellite loci and molecular sexing. Our objectives were to determine the population genetic structure, compare relatedness within and between social groups, infer kinship, and estimate the age of males at dispersal. Results suggest that individuals visiting the archipelago of the Azores belong to a single population. High genetic diversity and absence of inbreeding suggest that the population is recovering from whaling. Individuals sampled in close association are highly related, as well as those observed in the same area on the same day, suggesting that secondary social groups (i.e., the union of primary social units) are largely but not exclusively composed of relatives. Probable mother–offspring and full-sibling pairs were identified. Age of males at dispersal was estimated at 16.6 years, which was well above previous estimates for this species.

Using blubber biopsies to provide ecological information about bottlenose dolphins (Tursiops truncatus) around the Azores

Fatty acid profiles were used to investigate aspects of bottlenose dolphin populations around the Azores archipelago. Biopsy samples were obtained from 70 dolphins during the period 2002–2004. No statistically significant differences in profiles were found between different island groups, between sexes or between year of sampling. Thus no evidence was seen for island group fidelity, in contrast to bottlenose dolphins found around similar island groups such as the Hawaiian archipelago or the Bahamas. The findings are consistent with concurrent genetic and photo-identification studies on dolphins in the Azores.

Macro- and micro-geographic variation of short-beaked common dolphin’s whistles in the Mediterranean Sea and Atlantic Ocean

Genetic studies have shown that there are small but significant differences between the short-beaked common dolphin populations in the Atlantic Ocean and those in the Mediterranean Sea. The short-beaked common dolphin is a highly vocal species with a wide sound production repertoire including whistles. Whistles are continuous, narrowband, frequency-modulated signals that can show geographic variation in dolphin species. This study tests whether the differences, highlighted by genetic studies, are recognisable in the acoustic features of short-beaked common dolphin’s whistles in the two adjacent areas of the Atlantic Ocean and the Mediterranean Sea. From a selected sample of good quality whistles (514 recorded in the Atlantic and 193 in the Mediterranean) 10 parameters of duration, frequency and frequency modulation were measured. Comparing data among basins, differences were found for duration and all frequency parameters except for minimum frequency. Modulation parameters showed the highest coefficient of variation. Through discriminant analysis we correctly assigned 75.7% of sounds to their basins. Furthermore, micro-geographic analysis revealed similarity between the sounds recorded around the Azores and the Canary archipelagos and between the Bay of Biscay and the Mediterranean Sea. Results are in agreement with the hypothesis proposed by previous genetic studies that two distinct populations are present, still supposing a gene flow between the basins. This study is the first to compare short-beaked common dolphin’s whistles of the Atlantic Ocean and the Mediterranean areas.