Manatees display diel trends in acoustic activity at two microhabitats in Belize.

Many marine mammals exhibit diel trends in vocal production, which can provide information on habitat use and behavioral activity. In Belize, Antillean manatees (Trichechus manatus manatus) commonly inhabit small depressions in the substrate or deep-water coves known as "resting holes". Determining if manatees exhibit diel temporal trends in their call production rate and call types between microhabitats can provide insights into their diurnal and nocturnal activity patterns. Here, we investigate the diel vocalization patterns of wild Antillean manatees in two adjacent resting holes off of St. George's Caye, Belize. Recordings of manatees were made using a bottom-mounted hydrophone located near a reef barrier reef for nine days in July of 2017 and ten days in January of 2018. To explore if and how manatee acoustic activity differs between sites, we compared the number of calls per hour, the number of manatee positive hours, the number of tonal and atonal sounds, and the number of boats detected across sites. A total of 370 hours of acoustic recordings were analyzed resulting in the detection of 3,262 calls. There were no significant differences in the number of manatee calls produced per hour between sites. The average number of calls produced by manatees decreased over the course of several days. The proportion of tonal calls decreased with hours after sunset and increased in boat presence. These results suggest manatees in this region may exhibit different diel activity patterns which appear to be influenced by the characteristics of the environment. These findings can support ongoing conservation and management efforts to safeguard species in Belize.

Differences in the whistles of two ecotypes of bottlenose dolphins from the Gulf of California.

Intraspecific whistle variation in dolphins has been explained in the context of geographical isolation, soundscape, and social structure. Whistles of two ecotypes of bottlenose dolphins from La Paz Bay, Gulf of California, were analyzed. Both ecotypes showed similar whistle contours. However, they could be identified by contour maximum frequency which was predominantly above 15 kHz for oceanic dolphins and below 15 kHz for coastal dolphins. The different whistle frequencies of the two ecotypes could result from differences in group size and acoustic characteristics of their respective habitats, suggesting potential future passive acoustic monitoring applications.

Tour boats affect the activity patterns of bottlenose dolphins (Tursiops truncatus) in Bocas del Toro, Panama.

Bottlenose dolphins (Tursiops truncatus) of the Bocas del Toro archipelago are targeted by the largest boat-based cetacean watching operation in Panama. Tourism is concentrated in Dolphin Bay, home to a population of resident dolphins. Previous studies have shown that tour boats elicit short-term changes in dolphin behavior and communication; however, the relationship of these responses to the local population's biology and ecology is unclear. Studying the effects of tour boats on dolphin activity patterns and behavior can provide information about the biological significance of these responses. Here, we investigated the effects of tour boat activity on bottlenose dolphin activity patterns in Bocas del Toro, Panama over 10 weeks in 2014. Markov chain models were used to assess the effect of tour boats on dolphin behavioral transition probabilities in both control and impact scenarios. Effect of tour boat interactions was quantified by comparing transition probabilities of control and impact chains. Data were also used to construct dolphin activity budgets. Markov chain analysis revealed that in the presence of tour boats, dolphins were less likely to stay socializing and were more likely to begin traveling, and less likely to begin foraging while traveling. Additionally, activity budgets for foraging decreased and traveling increased as an effect of tour boat presence. These behavioral responses are likely to have energetic costs for individuals which may ultimately result in population-level impacts. Boat operator compliance with Panamanian whale watching regulations is urgently needed to minimize potential long-term impacts on this small, genetically distinct population and to ensure the future viability of the local tourism industry.

Foraging habits and levels of mercury in a resident population of bottlenose dolphins (Tursiops truncatus) in Bocas del Toro Archipelago, Caribbean Sea, Panama.

A small and genetically isolated bottlenose dolphin (Tursiops truncatus) population resides year-round in the Bocas del Toro Archipelago-Panama (BDT). Photo-identification and genetic data showed that this dolphin population is highly phylopatric and is formed exclusively by individuals of the "inshore form". This study aimed to investigate the trophic ecology and mercury concentrations of bottlenose dolphins in BDT to assess their coastal habits. We collected muscle samples (n = 175) of 11 potential fish prey species, and skin samples from free-ranging dolphins in BDT (n = 37) and La Guajira-Colombia (n = 7) to compare isotopic niche width. Results showed that BDT dolphins have a coastal feeding habit, belong to the "inshore form" (δ13C = -13.05 ±â€¯1.89‰), and have low mercury concentrations (mean = 1637 ±â€¯1387 ng g-1dw). However, this element is biomagnified in the BDT food chain, showing a marginal dolphins health risk (RQ = 1.00). We call for a monitoring pollutant program and conservation strategies aimed to protect the dolphin population at BDT.

Song structure and singing activity of two separate humpback whales populations wintering off the coast of Caño Island in Costa Rica.

Central American (CA) and Breeding Stock-G (BSG) humpback whales are known to winter off Caño Island, Costa Rica at different times of the year. To study their singing behavior, autonomous underwater recorders were used to record the whales. Song detection for BSG whales was higher than CA whales, and song structure was distinct for each population. No strong evidence for cross-equatorial connectivity was found. This study provides the first humpback whale song reference for both populations in Costa Rica, which can help advance understanding of CA and BSG whale song rate of change and connectivity with other wintering areas.

High genetic structure and low mitochondrial diversity in bottlenose dolphins of the Archipelago of Bocas del Toro, Panama: A population at risk?

The current conservation status of the bottlenose dolphin (Tursiops truncatus) under the IUCN is 'least concern'. However, in the Caribbean, small and localized populations of the 'inshore form' may be at higher risk of extinction than the 'worldwide distributed form' due to a combination of factors including small population size, high site fidelity, genetic isolation, and range overlap with human activities. Here, we study the population genetic structure of bottlenose dolphins from the Archipelago of Bocas del Toro in Panama. This is a small population characterized by high site fidelity and is currently heavily-impacted by the local dolphin-watching industry. We collected skin tissue samples from 25 dolphins to study the genetic diversity and structure of this population. We amplified a portion of the mitochondrial Control Region (mtDNA-CR) and nine microsatellite loci. The mtDNA-CR analyses revealed that dolphins in Bocas del Toro belong to the 'inshore form', grouped with the Bahamas-Colombia-Cuba-Mexico population unit. They also possess a unique haplotype new for the Caribbean. The microsatellite data indicated that the Bocas del Toro dolphin population is highly structured, likely due to restricted movement patterns. Previous abundance estimates obtained with mark-recapture methods reported a small population of 80 dolphins (95% CI = 72-87), which is similar to the contemporary effective population size estimated in this study (Ne = 73 individuals; CI = 18.0 - ∞; 0.05). The combination of small population size, high degree of genetic isolation, and intense daily interactions with dolphin-watching boats puts the Bocas del Toro dolphin to at high risk of extinction. Despite national guidelines to regulate the dolphin-watching industry in Bocas del Toro and ongoing educational programs for tour operators, only in 2012 seven animals have died due to boat collisions. Our results suggest that the conservation status of bottlenose dolphins in Bocas del Toro should be elevated to 'endangered' at the national level, as a precautionary measure while population and viability estimates are conducted.

Islands within islands: Diversification of tailless whip spiders (Amblypygi, Phrynus) in Caribbean caves.

Islands have played a key role in understanding species formation ever since Darwin's work on the Galapagos and Wallace's work in the Malay Archipelago. Like oceanic islands, habitat 'islands', such as mountaintops and caves similarly may drive diversification. Here we examine patterns of diversification in the tailless whip spider genus Phrynus Larmarck, 1809 (Amblypygida: Phrynidae) a system that shows evidence of diversification under the influence of 'islands within islands'. We estimate phylogeographic history and measure genetic diversity among representatives of three nominal Phrynus species from epigean and cave systems of Puerto Rico and nearby islands. Data from five loci (mitochondrial 12S, 16S, Cox1; nuclear H3, 28S) were used to generate phylogenetic hypotheses and to assess species monophyly and phylogeographic relationships. Genetic divergences and population limits were estimated and assessed using the Geneious barcoding plugin and the genealogical sorting index. We find that mtDNA sequence divergences within each of the three Phrynus species range between 15% and 20%. Genetic divergence is structured at three spatial scales: among islands in a manner consistent with the GAARlandia hypothesis, among bedrock formations within Puerto Rico, and among caves within these formations. Every isolated cave system contains a unique mtDNA genetic lineage of Phrynus, with divergence among cave systems far exceeding that within. In some localities epigean specimens nest among cave taxa, in others caves are monophyletic. Remarkably, clades that show up to 20% mtDNA sequence divergence show little or no variation in the nuclear markers. We interpret this pattern as resulting from extreme conservation of our nuclear markers rather than male sex-biased dispersal, based on high conservation of 28S and H3 between our individuals and other amblypygid genera that are restricted to Africa. While this study includes but a tiny fraction of Caribbean caves, our findings suggest Phrynus may be much more diverse than hitherto thought, at least in terms of mtDNA diversity, and that the arthropod fauna of caves may represent a dimension of biodiversity that is yet to be discovered in the Caribbean biodiversity hotspot.

Guyana dolphins (Sotalia guianensis) from Costa Rica emit whistles that vary with surface behaviors.

Guyana dolphins show remarkable intraspecific whistle variation. This variation has been largely explained in terms of distance among populations; however, other factors such as behavior may also be important. A broadband recording system recorded the whistles of Guyana dolphins under three behavioral states. A discriminant analysis found that during social and travel events, dolphins emit whistles with high delta and minimum frequency, respectively. Whistle duration was also important in discriminating behaviors. This study indicates that behavior is an important factor contributing to whistle variation of Guyana dolphins. Understanding how dolphin whistles vary with behavioral context will advance our understanding of dolphin communication and enable appropriate comparative studies.

Small-scale estimation of relative abundance for the coastal spotted dolphins (Stenella attenuata) in Costa Rica: the effect of habitat and seasonality / Estimación a pequeña escala de la abundancia relativa del delfín manchado costero (Stenella attenuata) en Costa Rica: el efecto del hábitat y la estacionalidad

The coastal spotted dolphin (Stenella attenuata graffmani) is one of the most common species of dolphin in inshore Pacific waters of Costa Rica. We conducted surveys in protected waters of the Papagayo Gulf, Costa Rica, to determine relative abundance of dolphins in relation to environmental variables. We used Generalized Additive Models to investigate the influence of a particular set of environmental factors and determine inter-annual trends in relative abundance. School sizes ranged from 1 to 50 individuals ( mean 9.95, SD=10.28). The number of dolphins increased linearly with water depth and transparency, and non-linearly with the dissolved oxygen concentration. High variability in the relative abundance occurred during the dry season (January-April). A previous study on this population found that high number of groups are involved in foraging activities during the dry season. Seasonal changes in relative abundance probably are associated with food availability, a variable that we did not measure. Understanding local resident populations may have important implications for conservation and management strategies. Large-scale studies may overlook variables affecting the abundance of local resident populations that may be detected with studies on a smaller scale such as this one.

El delfín manchado costero (Stenella attenuata graffmani) es una de las especies de delfines mas comunes de las aguas costeras del Pacifico de Costa Rica. En este estudio realizamos muestreos dentro de las aguas protegidas del Golfo de Papagayo para determinar su abundancia relativa en relación a características físico-químicas de su hábitat. Usamos modelos aditivos generalizados para investigar la influencia de un juego de variables ambientales y determinar tendencias inter-anuales en la abundancia relativa. El tamaño de los grupos varió de 1 a 50 individuos (promedio 9.95, SD=10.28). La cantidad de delfines aumentó linealmente con la profundidad y claridad del agua, y de forma no lineal con el oxígeno disuelto. Durante la época seca (enero-abril) encontramos la mayor variabilidad en la abundancia relativa. Un estudio anterior sugiere que una cantidad importante de delfines se alimenta aquí en la época seca. Ello sugiere que los cambios estacionales en la abundancia relativa de delfines manchados costeros podría estar asociada con la disponibilidad de alimento, una variable que no medimos pero que podría explicar la variación observada debido a la naturaleza productiva estacional del área. Los estudios de poblaciones locales y residentes pueden tener un mayor impacto en estrategias de conservación y manejo. Por lo general, los estudios a mayor escala geográfica pueden pasar por alto variables importantes que afectan la abundancia local de poblaciones residentes, las cuales pueden ser detectadas en estudios de menor escala como el presente.

Expensive Brains: "Brainy" Rodents have Higher Metabolic Rate.

Brains are the centers of the nervous system of animals, controlling the organ systems of the body and coordinating responses to changes in the ecological and social environment. The evolution of traits that correlate with cognitive ability, such as relative brain size is thus of broad interest. Brain mass relative to body mass (BM) varies among mammals, and diverse factors have been proposed to explain this variation. A recent study provided evidence that energetics play an important role in brain evolution (Isler and van Schaik, 2006). Using composite phylogenies and data drawn from multiple sources, these authors showed that basal metabolic rate (BMR) correlates with brain mass across mammals. However, no such relationship was found within rodents. Here we re-examined the relationship between BMR and brain mass within Rodentia using a novel species-level phylogeny. Our results are sensitive to parameter evaluation; in particular how species mass is estimated. We detect no pattern when applying an approach used by previous studies, where each species BM is represented by two different numbers, one being the individual that happened to be used for BMR estimates of that species. However, this approach may compromise the analysis. When using a single value of BM for each species, whether representing a single individual, or available species mean, our findings provide evidence that brain mass (independent of BM) and BMR are correlated. These findings are thus consistent with the hypothesis that large brains evolve when the payoff for increased brain mass is greater than the energetic cost they incur.

Phylogenetic analysis of conservation priorities for aquatic mammals and their terrestrial relatives, with a comparison of methods.

BACKGROUND: Habitat loss and overexploitation are among the primary factors threatening populations of many mammal species. Recently, aquatic mammals have been highlighted as particularly vulnerable. Here we test (1) if aquatic mammals emerge as more phylogenetically urgent conservation priorities than their terrestrial relatives, and (2) if high priority species are receiving sufficient conservation effort. We also compare results among some phylogenetic conservation methods. METHODOLOGY/PRINCIPAL FINDINGS: A phylogenetic analysis of conservation priorities for all 620 species of Cetartiodactyla and Carnivora, including most aquatic mammals. Conservation priority ranking of aquatic versus terrestrial species is approximately proportional to their diversity. However, nearly all obligated freshwater cetartiodactylans are among the top conservation priority species. Further, ∼74% and 40% of fully aquatic cetartiodactylans and carnivores, respectively, are either threatened or data deficient, more so than their terrestrial relatives. Strikingly, only 3% of all 'high priority' species are thought to be stable. An overwhelming 97% of these species thus either show decreasing population trends (87%) or are insufficiently known (10%). Furthermore, a disproportional number of highly evolutionarily distinct species are experiencing population decline, thus, such species should be closely monitored even if not currently threatened. Comparison among methods reveals that exact species ranking differs considerably among methods, nevertheless, most top priority species consistently rank high under any method. While we here favor one approach, we also suggest that a consensus approach may be useful when methods disagree. CONCLUSIONS/SIGNIFICANCE: These results reinforce prior findings, suggesting there is an urgent need to gather basic conservation data for aquatic mammals, and special conservation focus is needed on those confined to freshwater. That evolutionarily distinct--and thus 'biodiverse'--species are faring relatively poorly is alarming and requires further study. Our results offer a detailed guide to phylogeny-based conservation prioritization for these two orders.

Dogs, cats, and kin: a molecular species-level phylogeny of Carnivora.

Phylogenies underpin comparative biology as high-utility tools to test evolutionary and biogeographic hypotheses, inform on conservation strategies, and reveal the age and evolutionary histories of traits and lineages. As tools, most powerful are those phylogenies that contain all, or nearly all, of the taxa of a given group. Despite their obvious utility, such phylogenies, other than summary 'supertrees', are currently lacking for most mammalian orders, including the order Carnivora. Carnivora consists of about 270 extant species including most of the world's large terrestrial predators (e.g., the big cats, wolves, bears), as well as many of man's favorite wild (panda, cheetah, tiger) and domesticated animals (dog, cat). Distributed globally, carnivores are highly diverse ecologically, having occupied all major habitat types on the planet and being diverse in traits such as sociality, communication, body/brain size, and foraging ecology. Thus, numerous studies continue to address comparative questions within the order, highlighting the need for a detailed species-level phylogeny. Here we present a phylogeny of Carnivora that increases taxon sampling density from 28% in the most detailed primary-data study to date, to 82% containing 243 taxa (222 extant species, 17 subspecies). In addition to extant species, we sampled four extinct species: American cheetah, saber-toothed cat, cave bear and the giant short-faced bear. Bayesian analysis of cytochrome b sequences data-mined from GenBank results in a phylogenetic hypothesis that is largely congruent with prior studies based on fewer taxa but more characters. We find support for the monophyly of Carnivora, its major division into Caniformia and Feliformia, and for all but one family within the order. The only exception is the placement of the kinkajou outside Procyonidae, however, prior studies have already cast doubt on its family placement. In contrast, at the subfamily and genus level, our results indicate numerous problems with current classification. Our results also propose new, controversial hypotheses, such as the possible placement of the red panda (Ailuridae) sister to canids (Canidae). Our results confirm previous findings suggesting that the dog was domesticated from the Eurasian wolf (Canis lupus lupus) and are congruent with the Near East domestication of the cat. In sum, this study presents the most detailed species-level phylogeny of Carnivora to date and a much needed tool for comparative studies of carnivoran species. To demonstrate one such use, we perform a phylogenetic analysis of evolutionary distinctiveness (EDGE), which can be used to help establish conservation priorities. According with those criteria, and under one of the many possible sets of parameters, the highest priority Carnivora species for conservation of evolutionary diversity include: monk seals, giant and red panda, giant otter, otter civet, Owston's palm civet, sea otter, Liberian mongoose, spectacled bear, walrus, binturong, and the fossa.

A characterization of Guyana dolphin (Sotalia guianensis) whistles from Costa Rica: the importance of broadband recording systems.

Knowledge of the whistle structure in Guyana dolphins comes mostly from Brazilian populations where recordings have been made using limited bandwidth systems (18 and 24 kHz). In Brazil, Guyana dolphin whistle frequency span is 1.34-23.89 kHz, but authors have suggested that limits of their recording system may underestimate frequency span. Whistles of Guyana dolphins from Costa Rica were studied using a broadband recording system. How bandwidth limitations affect the understanding of whistle structure and species classification between sympatric dolphin species was evaluated. In addition, whistles were compared to Brazilian populations. Guyana dolphin whistle frequency span was 1.38 up to 48.40 kHz, greater than previously reported. Bandwidth limitations explained 89% of the whistle variation between studies, and increase in bandwidth improved the whistle classification of Guyana dolphins. Whistle duration and minimum frequency were the most important variables in dolphin species classification. Finally, after accounting for differences in recording systems, Costa Rican Guyana dolphins whistled with significantly higher frequency than Brazilian populations, providing evidence for a postulated increase in frequency from south to north. The study concludes that equipment with an upper frequency limit of at least 50 kHz (150 kHz for harmonics) is required to capture the entire whistle repertoire of the Guyana dolphin.

The freshwater dolphin Inia geoffrensis geoffrensis produces high frequency whistles.

Because whistles are most commonly associated with social delphinids, they have been largely overlooked, ignored, or presumed absent, in solitary freshwater dolphin species. Whistle production in the freshwater dolphin, the boto (Inia geoffrensis geoffrensis), has been controversial. Because of its sympatry with tucuxi dolphins (Sotalia fluviatilis), a whistling species, some presume tucuxi whistles might have been erroneously assigned to the boto. Using a broadband recording system, we recorded over 100 whistles from boto dolphins in the Yasunf River, Ecuador, where the tucuxi dolphins are absent. Our results therefore provide conclusive evidence for whistle production in Inia geoffrensis geoffrensis. Furthermore, boto whistles are significantly different from tucuxi whistles recorded in nearby rivers. The Ecuadorian boto whistle has a significantly greater frequency range (5.30-48.10 kHz) than previously reported in other populations (Peru and Colombia) that were recorded with more bandwidth limited equipment. In addition, the top frequency and the range are greater than in any other toothed whale species recorded to date. Whistle production was higher during resting activities, alone or in the presence of other animals. The confirmation of whistles in the boto has important implications for the evolution of whistles in Cetacea and their association with sociality.