Comparative bioacoustics: a roadmap for quantifying and comparing animal sounds across diverse taxa.

Animals produce a wide array of sounds with highly variable acoustic structures. It is possible to understand the causes and consequences of this variation across taxa with phylogenetic comparative analyses. Acoustic and evolutionary analyses are rapidly increasing in sophistication such that choosing appropriate acoustic and evolutionary approaches is increasingly difficult. However, the correct choice of analysis can have profound effects on output and evolutionary inferences. Here, we identify and address some of the challenges for this growing field by providing a roadmap for quantifying and comparing sound in a phylogenetic context for researchers with a broad range of scientific backgrounds. Sound, as a continuous, multidimensional trait can be particularly challenging to measure because it can be hard to identify variables that can be compared across taxa and it is also no small feat to process and analyse the resulting high-dimensional acoustic data using approaches that are appropriate for subsequent evolutionary analysis. Additionally, terminological inconsistencies and the role of learning in the development of acoustic traits need to be considered. Phylogenetic comparative analyses also have their own sets of caveats to consider. We provide a set of recommendations for delimiting acoustic signals into discrete, comparable acoustic units. We also present a three-stage workflow for extracting relevant acoustic data, including options for multivariate analyses and dimensionality reduction that is compatible with phylogenetic comparative analysis. We then summarize available phylogenetic comparative approaches and how they have been used in comparative bioacoustics, and address the limitations of comparative analyses with behavioural data. Lastly, we recommend how to apply these methods to acoustic data across a range of study systems. In this way, we provide an integrated framework to aid in quantitative analysis of cross-taxa variation in animal sounds for comparative phylogenetic analysis. In addition, we advocate the standardization of acoustic terminology across disciplines and taxa, adoption of automated methods for acoustic feature extraction, and establishment of strong data archival practices for acoustic recordings and data analyses. Combining such practices with our proposed workflow will greatly advance the reproducibility, biological interpretation, and longevity of comparative bioacoustic studies.

Seasonal movements of Gulf of Mexico sperm whales following the Deepwater Horizon oil spill and the limitations of impact assessments.

As part of the Deepwater Horizon Oil Spill Natural Resource Damage Assessment in the Gulf of Mexico, we conducted a large passive acoustic survey across the eastern Gulf continental shelf edge to assess impacts to sperm whale population. In the months immediately after the spill, sperm whale occurrence was significantly higher in areas closest to the spill. Over the following seasons in 2010-2011, we documented cyclical patterns of decreased and increased occurrence suggesting that this population exhibits a seasonal occurrence pattern in the region, with seasonal movements to other regions, and not likely directly influenced by the oil spill. Unfortunately, a lack of adequately scaled, pre-spill data on sperm whales, along with limitations on the survey duration constrain our ability to infer spill-related changes in sperm whale occurrence. However, our study establishes post-disaster baseline data for continued monitoring, and an expanded study design could provide a model for continued monitoring.

Evolution of correlated complexity in the radically different courtship signals of birds-of-paradise.

Ornaments used in courtship often vary wildly among species, reflecting the evolutionary interplay between mate preference functions and the constraints imposed by natural selection. Consequently, understanding the evolutionary dynamics responsible for ornament diversification has been a longstanding challenge in evolutionary biology. However, comparing radically different ornaments across species, as well as different classes of ornaments within species, is a profound challenge to understanding diversification of sexual signals. Using novel methods and a unique natural history dataset, we explore evolutionary patterns of ornament evolution in a group-the birds-of-paradise-exhibiting dramatic phenotypic diversification widely assumed to be driven by sexual selection. Rather than the tradeoff between ornament types originally envisioned by Darwin and Wallace, we found positive correlations among cross-modal (visual/acoustic) signals indicating functional integration of ornamental traits into a composite unit-the "courtship phenotype." Furthermore, given the broad theoretical and empirical support for the idea that systemic robustness-functional overlap and interdependency-promotes evolutionary innovation, we posit that birds-of-paradise have radiated extensively through ornamental phenotype space as a consequence of the robustness in the courtship phenotype that we document at a phylogenetic scale. We suggest that the degree of robustness in courtship phenotypes among taxa can provide new insights into the relative influence of sexual and natural selection on phenotypic radiations.

Seasonal and geographical patterns of fin whale song in the western North Atlantic Ocean.

Male fin whales, Balaenoptera physalus, produce a song consisting of 20 Hz notes at regularly spaced time intervals. Previous studies identified regional differences in fin whale internote intervals (INI), but seasonal changes within populations have not been closely examined. To understand the patterns of fin whale song in the western North Atlantic, the seasonal abundance and acoustic features of fin whale song are measured from two years of archival passive acoustic recordings at two representative locations: Massachusetts Bay and New York Bight. Fin whale 20 Hz notes are detected on 99% of recorded days. In both regions, INI varies significantly throughout the year as two distinct periods: a "short-INI" season in September-January (9.6 s) and a "long-INI" season in March-May (15.1 s). February and June-August are transitional-INI months, with higher variability. Note abundance decreases with increasing INI, where note abundance is significantly lower in April-August than in September-January. Short-INI and high note abundance correspond to the fin whale reproductive season. The temporal variability of INI may be a mechanism by which fin whale individuals encode and communicate a variety of behaviorally relevant information.

Acoustically detected year-round presence of right whales in an urbanized migration corridor.

Species' conservation relies on understanding their seasonal habitats and migration routes. North Atlantic right whales (Eubalaena glacialis), listed as endangered under the U.S. Endangered Species Act, migrate from the southeastern U.S. coast to Cape Cod Bay, Massachusetts, a federally designated critical habitat, from February through May to feed. The whales then continue north across the Gulf of Maine to northern waters (e.g., Bay of Fundy). To enter Cape Cod Bay, right whales must traverse an area of dense shipping and fishing activity in Massachusetts Bay, where there are no mandatory regulations for the protection of right whales or management of their habitat. We used passive acoustic recordings of right whales collected in Massachusetts Bay from May 2007 through October 2010 to determine the annual spatial and temporal distribution of the whales and their calling activity. We detected right whales in the bay throughout the year, in contrast to results from visual surveys. Right whales were detected on at least 24% of days in each month, with the exception of June 2007, in which there were no detections. Averaged over all years, right whale calls were most abundant from February through May. During this period, calls were most frequent between 17:00 and 20:00 local time; no diel pattern was apparent in other months. The spatial distribution of the approximate locations of calling whales suggests they may use Massachusetts Bay as a conduit to Cape Cod Bay in the spring and as they move between the Gulf of Maine and waters to the south in September through December. Although it is unclear how dependent right whales are on the bay, the discovery of their widespread presence in Massachusetts Bay throughout the year suggests this region may need to be managed to reduce the probability of collisions with ships and entanglement in fishing gear.