Categorizing click trains to increase taxonomic precision in echolocation click loggers.

Passive acoustic monitoring is an efficient way to study acoustically active animals but species identification remains a major challenge. C-PODs are popular logging devices that automatically detect odontocete echolocation clicks. However, the accompanying analysis software does not distinguish between delphinid species. Click train features logged by C-PODs were compared to frequency spectra from adjacently deployed continuous recorders. A generalized additive model was then used to categorize C-POD click trains into three groups: broadband click trains, produced by bottlenose dolphin (Tursiops truncatus) or common dolphin (Delphinus delphis), frequency-banded click trains, produced by Risso's (Grampus griseus) or white beaked dolphins (Lagenorhynchus albirostris), and unknown click trains. Incorrect categorization rates for broadband and frequency banded clicks were 0.02 (SD 0.01), but only 30% of the click trains met the categorization threshold. To increase the proportion of categorized click trains, model predictions were pooled within acoustic encounters and a likelihood ratio threshold was used to categorize encounters. This increased the proportion of the click trains meeting either the broadband or frequency banded categorization threshold to 98%. Predicted species distribution at the 30 study sites matched well to visual sighting records from the region.

Post-glacial habitat release and incipient speciation in the genus Delphinus.

The role of ecological and changing environmental factors in the radiation of species diversity is a fundamental question in evolutionary biology. Of particular interest is the potential for these factors to determine the boundary between what we would consider differentiation among populations and incipient speciation. Dolphins in the genus Delphinus provide a useful test case, exhibiting morphological variation in beak length, coloration and body size across their wide geographic distribution, and in particular among coastal and more pelagic habitats. Two species have been proposed, D. delphis and D. capensis, but morphologically similar allopatric populations are not monophyletic, indicating that the mostly coastal 'long-beaked' D. capensis form is not a single globally distributed species. However, the sympatric populations in the Eastern North Pacific currently designated as these two species are both morphologically and genetically differentiated. Here we use microsatellite DNA and mitochondrial DNA markers to investigate the evolutionary mechanisms that led to this incipient speciation event. We used coalescent and assignment methods to investigate the timing and extent of reproductive isolation. Our data indicate that although there is some level of on-going gene flow, the putative species found in the Eastern North Pacific are reciprocally monophyletic. The timing of isolation appears to be associated with regional changes in paleoceanographic conditions within the Holocene timeframe.

Combining whistle acoustic parameters to discriminate Mediterranean odontocetes during passive acoustic monitoring.

Acoustic observation can complement visual observation to more effectively monitor occurrence and distribution of marine mammals. For effective acoustic censuses, calibration methods must be determined by joint visual and acoustic studies. Research is still needed in the field of acoustic species identification, particularly for smaller odontocetes. From 1994 to 2012, whistles of four odontocete species were recorded in different areas of the Mediterranean Sea to determine how reliably these vocalizations can be classified to species. Recordings were attributed to species by simultaneous visual observation. The results of this study highlight that the frequency parameters, which are linked to physical features of animals, show lower variability than modulation parameters, which are likely to be more dependent on complex eco-ethological contexts. For all the studied species, minimum and maximum frequencies were linearly correlated with body size. DFA and Classification Tree Analysis (CART) show that these parameters were the most important for classifying species; however, both statistical methods highlighted the need for combining them with the number of contour minima and contour maxima for correct classification. Generally, DFA and CART results reflected both phylogenetic distance (especially for common and striped dolphins) and the size of the species.

Sounds in common: Time-frequency as the classification parameters for pulsed sounds produced by Delphinus delphis.

Sounds produced by dolphins can be grouped into tonal (whistles) and pulsed sounds (e.g., echolocation clicks and burst sounds). Clicks are broadband pulses temporarily spaced to allow echo processing between the sound source and the object. Echolocation is related mainly to prey detection and environmental recognition. Echolocation click trains tend to present a decreasing inter-click interval due to a continuous changing of the target's location when the animal approaches to capture the prey. In addition to foraging and feeding contexts, burst pulsed sounds have been associated with short social communication. Although echolocation clicks are relatively well documented, there is no consensus regarding the broad variety of the burst pulsed signals. The present study analyzed time-frequency characteristics by conducting a clustering and discrimination analysis to classify pulsed sounds. A total of 64 click trains were analyzed from short-beaked common dolphins recorded in the slope region of the western South Atlantic Ocean. Three analyses (time; frequency; and combined time-frequency parameters) were compared through k-means clustering and posterior cluster validation using Random forest analysis. The k-means clustering resulted in four clusters for all groups of analysis. The time parameters were the most accurate among the comparisons, with the first two-dimensional axis corresponding to 87 % (Dim1 = 70.2 % and Dim2 = 17.2 %). The random forest analysis showed that the time-frequency dataset was the best classification of pulsed sounds in D. delphis (Accuracy = 84.6 %; confidence interval CI = 65.1 %-95.6 %; p < 0.01). This result considers the animal an acoustical identity, emphasizing the importance of certain parameters that influence this identity and thus reflecting the energy-cost optimization for sound production.

Population genetic structure of the short-beaked common dolphin from the Black Sea and the Turkish Straits System.

Our study aims to assess the population connectivity, evolutionary history, and conservation status of the short-beaked common dolphin in the Black Sea and Turkish Straits System (TSS). We also include DNA sequences from the Atlantic Ocean and the Mediterranean Sea to provide a regional perspective to our localized study. Analysis of 366 base pairs of mitochondrial DNA D-loop fragments from 37 samples collected from short-beaked common dolphins in the Black Sea, TSS, and Aegean Sea revealed 13 haplotypes, eight of which have not been previously reported. While analysis of samples archived on GenBank revealed 89 different haplotypes across the region. The haplotype network contains two main peripheral groups that include individuals from all locations. Haplotypes from the Atlantic Ocean are scattered across the network and no obvious population separation was detected. Some shared haplotypes potentially indicate multi-directional colonization events of the Mediterranean Sea from the eastern Atlantic Ocean. Moreover, some less widely distributed haplotypes suggest some level of more recent genetic connectivity through the Strait of Gibraltar and the TSS and point out the importance of these straits in the dispersal of short-beaked common dolphins. The haplotype and nucleotide diversity values were lower in the Black Sea, TSS, and western Mediterranean Sea when compared to the Atlantic Ocean, supporting the expansion of Atlantic populations into the Mediterranean and the Black Seas. Differentiation was observed between the Atlantic Ocean, and the Mediterranean Sea, TSS and the Black Sea based on Фst but not between Mediterranean and the Black Seas. For common dolphins, which have high dispersal potential, the protection of open seas and narrow seaways to enhance connectivity may be crucial.

Range extension for the common dolphin (Delphinus sp.) to the Colombian Caribbean, with taxonomic implications from genetic barcoding and phylogenetic analyses.

The nearest known population of common dolphins (Delphinus sp.) to the Colombian Caribbean occurs in a fairly restricted range in eastern Venezuela. These dolphins have not been previously reported in the Colombian Caribbean, likely because of a lack of study of the local cetacean fauna. We collected cetacean observations in waters of the Guajira Department, northern Colombia (~11°N, 73°W) during two separate efforts: (a) a seismic vessel survey (December 2009-March 2010), and (b) three coastal surveys from small boats (May-July 2012, May 2013, and May 2014). Here we document ten sightings of common dolphins collected during these surveys, which extend the known range of the species by ~1000 km into the southwestern Caribbean. We also collected nine skin biopsies in 2013 and 2014. In order to determine the taxonomic identity of the specimens, we conducted genetic barcoding and phylogenetic analyses using two mitochondrial markers, the Control Region (mtDNA) and Cytochrome b (Cytb). Results indicate that these specimens are genetically closer to the short-beaked common dolphin (Delphinus delphis) even though morphologically they resemble a long-beaked form (Delphinus sp.). However, the specific taxonomic status of common dolphins in the Caribbean and in the Western Atlantic remains unresolved. It is also unclear whether the distribution of the species between northern Colombia and eastern Venezuela is continuous or disjoined, or whether they can be considered part of the same stock.

Molecular and Morphological Differentiation of Common Dolphins (Delphinus sp.) in the Southwestern Atlantic: Testing the Two Species Hypothesis in Sympatry.

The taxonomy of common dolphins (Delphinus sp.) has always been controversial, with over twenty described species since the original description of the type species of the genus (Delphinus delphis Linnaeus, 1758). Two species and four subspecies are currently accepted, but recent molecular data have challenged this view. In this study we investigated the molecular taxonomy of common dolphins through analyses of cytochrome b sequences of 297 individuals from most of their distribution. We included 37 novel sequences from the Southwestern Atlantic Ocean, a region where the short- and long-beaked morphotypes occur in sympatry, but which had not been well sampled before. Skulls of individuals from the Southwestern Atlantic were measured to test the validity of the rostral index as a diagnostic character and confirmed the presence of the two morphotypes in our genetic sample. Our genetic results show that all common dolphins in the Atlantic Ocean belong to a single species, Delphinus delphis. According to genetic data, the species Delphinus capensis is invalid. Long-beaked common dolphins from the Northeastern Pacific Ocean may constitute a different species. Our conclusions prompt the need for revision of currently accepted common dolphin species and subspecies and of Delphinus delphis distribution.

The use of carcasses for the analysis of cetacean population genetic structure: a comparative study in two dolphin species.

Advances in molecular techniques have enabled the study of genetic diversity and population structure in many different contexts. Studies that assess the genetic structure of cetacean populations often use biopsy samples from free-ranging individuals and tissue samples from stranded animals or individuals that became entangled in fishery or aquaculture equipment. This leads to the question of how representative the location of a stranded or entangled animal is with respect to its natural range, and whether similar results would be obtained when comparing carcass samples with samples from free-ranging individuals in studies of population structure. Here we use tissue samples from carcasses of dolphins that stranded or died as a result of bycatch in South Australia to investigate spatial population structure in two species: coastal bottlenose (Tursiops sp.) and short-beaked common dolphins (Delphinus delphis). We compare these results with those previously obtained from biopsy sampled free-ranging dolphins in the same area to test whether carcass samples yield similar patterns of genetic variability and population structure. Data from dolphin carcasses were gathered using seven microsatellite markers and a fragment of the mitochondrial DNA control region. Analyses based on carcass samples alone failed to detect genetic structure in Tursiops sp., a species previously shown to exhibit restricted dispersal and moderate genetic differentiation across a small spatial scale in this region. However, genetic structure was correctly inferred in D. delphis, a species previously shown to have reduced genetic structure over a similar geographic area. We propose that in the absence of corroborating data, and when population structure is assessed over relatively small spatial scales, the sole use of carcasses may lead to an underestimate of genetic differentiation. This can lead to a failure in identifying management units for conservation. Therefore, this risk should be carefully assessed when planning population genetic studies of cetaceans.

Gaussian mixture model classification of odontocetes in the Southern California Bight and the Gulf of California.

A method for the automatic classification of free-ranging delphinid vocalizations is presented. The vocalizations of short-beaked and long-beaked common (Delphinus delphis and Delphinus capensis), Pacific white-sided (Lagenorhynchus obliquidens), and bottlenose (Tursiops truncatus) dolphins were recorded in a pelagic environment of the Southern California Bight and the Gulf of California over a period of 4 years. Cepstral feature vectors are extracted from call data which contain simultaneous overlapping whistles, burst-pulses, and clicks from a single species. These features are grouped into multisecond segments. A portion of the data is used to train Gaussian mixture models of varying orders for each species. The remaining call data are used to test the performance of the models. Species are predicted based upon probabilistic measures of model similarity with test segment groups having durations between 1 and 25 s. For this data set, 256 mixture Gaussian mixture models and segments of at least 10 s of call data resulted in the best classification results. The classifier predicts the species of groups with 67%-75% accuracy depending upon the partitioning of the training and test data.

Phylogeography and alpha taxonomy of the common dolphin (Delphinus sp.).

The resolution of taxonomic classifications for delphinid cetaceans has been problematic, especially for species in the genera Delphinus, Tursiops and Stenella. The frequent lack of correspondence between morphological and genetic differentiation in these species raises questions about the mechanisms responsible for their evolution. In this study we focus on the genus Delphinus, and use molecular markers to address questions about speciation and the evolution of population structure. Delphinus species have a worldwide distribution and show a high degree of morphological variation. Two distinct morphotypes, long-beaked and short-beaked, have been considered different species named D. capensis and D. delphis, respectively. However, genetic differentiation between these two forms has only been demonstrated in the Pacific. We analysed samples from eight different geographical regions, including two morphologically defined long-beaked form populations, and compared these with the eastern North Pacific populations. We found high differentiation among the populations described as long-beaked instead of the expected monophyly, suggesting that these populations may have evolved from independent events converging on the same morphotype. We observed low genetic differentiation among the short-beaked populations across a large geographical scale. We interpret these phylogeographical patterns in the context of life history and population structure in related species.