Evidence from sperm whale clans of symbolic marking in non-human cultures.

Culture, a pillar of the remarkable ecological success of humans, is increasingly recognized as a powerful force structuring nonhuman animal populations. A key gap between these two types of culture is quantitative evidence of symbolic markers-seemingly arbitrary traits that function as reliable indicators of cultural group membership to conspecifics. Using acoustic data collected from 23 Pacific Ocean locations, we provide quantitative evidence that certain sperm whale acoustic signals exhibit spatial patterns consistent with a symbolic marker function. Culture segments sperm whale populations into behaviorally distinct clans, which are defined based on dialects of stereotyped click patterns (codas). We classified 23,429 codas into types using contaminated mixture models and hierarchically clustered coda repertoires into seven clans based on similarities in coda usage; then we evaluated whether coda usage varied with geographic distance within clans or with spatial overlap between clans. Similarities in within-clan usage of both "identity codas" (coda types diagnostic of clan identity) and "nonidentity codas" (coda types used by multiple clans) decrease as space between repertoire recording locations increases. However, between-clan similarity in identity, but not nonidentity, coda usage decreases as clan spatial overlap increases. This matches expectations if sympatry is related to a measurable pressure to diversify to make cultural divisions sharper, thereby providing evidence that identity codas function as symbolic markers of clan identity. Our study provides quantitative evidence of arbitrary traits, resembling human ethnic markers, conveying cultural identity outside of humans, and highlights remarkable similarities in the distributions of human ethnolinguistic groups and sperm whale clans.

Population ecology and the management of whale watching operations on a data-deficient dolphin population.

Whale watching is a popular commercial activity, producing socio-ecological benefits but also potential long-term effects on the targeted cetacean population. This industry is currently developing in data-deficient contexts in a largely unregulated fashion. Management schemes should adopt precaution and be informed by the relevant literature, but would be more effective if the assessment of the target population vulnerability, biological impacts, and management implications was drawn from site-specific data.This paper focuses on a reef-associated, data-deficient population of spinner dolphins in the Egyptian Red Sea. In Satayah Reef, new information on population size and dynamic parameters were documented using visual observation and photo-identification-based capture-recapture methods (Cormack-Jolly-Seber time-since-marking model).Dolphins occurred on 98% of the survey days. Average school size was 66 individuals (±42.1 SE), with most groups including calves. The population was equally divided into recurrent and transient individuals. An "emigration + mortality" model best described residence at the site. Five recurrent males (5% of the Satayah population) provided connectivity between this and the geographically close population of Samadai Reef.Average annual survival probability was 0.83 (±0.06 SE) in the year following first capture and 0.99 (±0.06 SE) for recurrent individuals. Mean yearly population sizes ranged 143-207 individuals.The study had the power to detect a 30% decline in the population, but not the rate of change in abundance estimated from the data (r = 0.018 ± 0.04), which would have required a 3- to 5-times longer study. Synthesis and application: These findings advance the assessment of the Satayah population's intrinsic vulnerability and have three major management applications: (a) the delineation of management units; (b) the identification of key indicators for future impact monitoring and assessment; and (c) realistic estimates of the statistical power for trend detection. Based on our results, we recommend supporting future research, devising site-specific time-area closure plans, and integrating them in a regional scheme. Approaches employed in this case study can inform the management of whale watching industries targeting other data-deficient populations.

Behavioural responses of spinner dolphins to human interactions.

There is increasing evidence that whale and dolphin watching activities have detrimental effects on targeted cetacean populations. In Egypt, spinner dolphins regularly occur in the resting areas of Samadai, Satayah and Qubbat'Isa reefs. In-water human interactions with dolphins are regulated with a time-area closure system at Samadai, unregulated at Satayah and non-existent at Qubbat'Isa. This provided an ideal experimental setting to advance our understanding of the effects of tourism on a species highly sensitive to disturbances. Our study confirmed that the intensity and duration of interactions, and therefore, dolphin exposure to tourism, differed among the study sites. Compared with the Qubbat'Isa control site, behavioural reactions to boats and swimmers at the two tourism sites suggested that dolphin rest was disrupted, especially around the middle of the day and especially at Satayah, where dolphin tourism is unregulated. Our results indicate also that the dolphin protection measures at Samadai reduce the level of disturbance. We recommend that similar measures be implemented at other dolphin tourism locations, and that no new operations be initiated until the long-term impacts on dolphin populations are better understood. Our experience emphasizes the need to adopt precautionary approaches in research and management of whale and dolphin watching.

A simulation study of acoustic-assisted tracking of whales for mark-recapture surveys.

Collecting enough data to obtain reasonable abundance estimates of whales is often difficult, particularly when studying rare species. Passive acoustics can be used to detect whale sounds and are increasingly used to estimate whale abundance. Much of the existing effort centres on the use of acoustics to estimate abundance directly, e.g. analysing detections in a distance sampling framework. Here, we focus on acoustics as a tool incorporated within mark-recapture surveys. In this context, acoustic tools are used to detect and track whales, which are then photographed or biopsied to provide data for mark-recapture analyses. The purpose of incorporating acoustics is to increase the encounter rate beyond using visual searching only. While this general approach is not new, its utility is rarely quantified. This paper predicts the "acoustically-assisted" encounter rate using a discrete-time individual-based simulation of whales and survey vessel. We validate the simulation framework using existing data from studies of sperm whales. We then use the framework to predict potential encounter rates in a study of Antarctic blue whales. We also investigate the effects of a number of the key parameters on encounter rate. Mean encounter rates from the simulation of sperm whales matched well with empirical data. Variance of encounter rate, however, was underestimated. The simulation of Antarctic blue whales found that passive acoustics should provide a 1.7-3.0 fold increase in encounter rate over visual-only methods. Encounter rate was most sensitive to acoustic detection range, followed by vocalisation rate. During survey planning and design, some indication of the relationship between expected sample size and effort is paramount; this simulation framework can be used to predict encounter rates and establish this relationship. For a case in point, the simulation framework indicates unequivocally that real-time acoustic tracking should be considered for quantifying the abundance of Antarctic blue whales via mark-recapture methods.

Acoustically derived growth rates of sperm whales (Physeter macrocephalus) in Kaikoura, New Zealand.

A non-invasive acoustic method for measuring the growth of sperm whales was developed based on estimating the length of individuals by measuring the inter-pulse interval (IPI) of their clicks. Most prior knowledge of growth in male sperm whales has come from from fitting growth curves to length data gained from whaling. Recordings made at Kaikoura, New Zealand, were used to estimate the length and growth of 32 photographically identified, resident whales that have been recorded repeatedly between 1991 and 2009. All whales recorded more than six months apart (n = 30) showed an increase in IPI. Using established relationships between IPI and total length, it was found that the average growth rate in the Kaikoura population is lower, especially for smaller whales (13-14.5 m), than that derived from historical whaling data from other populations. This difference may be due to ecological differences among populations but might also reflect upward bias in measurements gained in whaling. The ability to track growth of individuals through time is only possible via non-lethal means and offers a fundamentally different kind of data because differences among individuals can be measured.

Measuring body length of male sperm whales from their clicks: the relationship between inter-pulse intervals and photogrammetrically measured lengths.

Sperm whales (Physeter macrocephalus) emit short, broadband clicks which often include multiple pulses. The time interval between these pulses [inter-pulse interval (IPI)] represents the two-way time for a pulse to travel between the air sacs located at either end of the sperm whale's head. The IPI therefore, is a proxy of head length which, using an allometric relationship, can be used to estimate total body length. Previous studies relating IPI to an independent measure of length have relied on very small sample sizes and manual techniques for measuring IPI. Sound recordings and digital stereo photogrammetric measurements of 21 individuals were made off Kaikoura, New Zealand, and, in addition, archived recordings of whales measured with a previous photogrammetric system were reanalyzed to obtain a total sample size of 33 individuals. IPIs were measured automatically via cepstral analysis implemented via a software plug-in for pamguard, an open-source software package for passive acoustic monitoring. IPI measurements were highly consistent within individuals (mean CV=0.63%). The new regression relationship relating IPI (I) and total length (T) was found to be T=1.258I+5.736 (r(2)=0.77, p<0.001). This new regression provides a better fit than previous studies of large (> 11 m) sperm whales.