Material & Methods

Maritime traffic map of the study area, the Strait of Gibraltar (circle in the middle). The two circles on the right indicate the core areas used year-round by resident pilot whales.

Data were collected in 2012, 2013, and 2015 in the Strait of Gibraltar, where six social groups of long-finned pilot whales were simultaneously tagged with DTAGs, digital tags that record both sound and movement. These bio-logging devices were attached using suction cups and programmed to stay on the animals for up to 24 hours.

The DTAGs captured fine-scale acoustic and behavioral data, enabling analysis of vocal output levels, dive behavior, and environmental noise. This setup allowed us to investigate how whales adjusted their vocalizations in response to varying noise levelsdepth, and behavioral context.

DTAGs during the test phase before deployment on whales.

The processing of the DTAG recordings began with manual auditing, where trained observers labeled each call by call type and the individual whale that produced it. These labeled calls were then passed through a machine learning model, which extracted acoustic features and grouped similar calls using unsupervised clustering techniques.

As a result, calls were categorized into four distinct types based on their acoustic structure:

  • Low-frequency calls
  • High-frequency calls
  • Short pulsed calls
  • Two-component calls

This classification allowed us to examine how vocal output levels varied not only with ambient noise and depth, but also across different call types.

The four call types distinguished in our study.

To analyze the influence of dive context, each call was linked to its corresponding dive phase using depth profiles generated for each tagged social group. We classified vocalizations into four dive contexts based on depth:

  • Surface: 0–80 m
  • Shallow dive: 0–300 m
  • Deep dive – descent phase: 0–800 m
  • Deep dive – ascent phase: 800–0 m

The dive plot (shown below) illustrates the diving behavior of three tagged individuals, with overlaid dots representing detected foraging events. Red dashed lines indicate the depth boundaries used to define each context.

Dive plot of a tagged subgroup of three individuals (gm13_220b, gm13_220d, gm13_220e). Dotted red lines indicate dive context categories. Yellow and orange dots mark foraging events.

We used linear mixed-effects models to estimate the magnitude of the Lombard response. Four models focused on the effect of call type, while five additional models incorporated depth and dive context as predictors to explore how these factors influenced vocal amplitude adjustments.