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.

Variability in humpback whale songs reveals how individuals can be distinctive when sharing a complex vocal display.

Individually distinctive acoustic signals in animal vocal communication are taxonomically widespread, however, the investigation of these signal types in marine mammals has focused only on a few species. Humpback whale songs are a stereotyped, hierarchically structured vocal display performed by males, and hence thought to be sexually selected. Within a population, whales conform to a common version of the song despite the song constantly evolving. While humpback songs have been studied extensively at the population level, individual level variation has been rarely described, with inconclusive results. Here, we quantified inter- and intra-individual variability at different levels in the song hierarchy using songs from 25 singers across two song types from the eastern Australian population song of 2002 (12 singers), and the revolutionary song introduced in 2003 (13 singers). Inter-individual variability was found heterogeneously across all hierarchical levels of the song structure. In addition, distinct and individually specific patterns of song production were consistently recorded across song levels, with clear structural differences between the two song types. These results suggest that within the constraints of song conformity, males can produce individually distinctive patterns that could function as an advertisement to females to convey individual qualities.

A deepening understanding of animal culture suggests lessons for conservation.

A key goal of conservation is to protect biodiversity by supporting the long-term persistence of viable, natural populations of wild species. Conservation practice has long been guided by genetic, ecological and demographic indicators of risk. Emerging evidence of animal culture across diverse taxa and its role as a driver of evolutionary diversification, population structure and demographic processes may be essential for augmenting these conventional conservation approaches and decision-making. Animal culture was the focus of a ground-breaking resolution under the Convention on the Conservation of Migratory Species of Wild Animals (CMS), an international treaty operating under the UN Environment Programme. Here, we synthesize existing evidence to demonstrate how social learning and animal culture interact with processes important to conservation management. Specifically, we explore how social learning might influence population viability and be an important resource in response to anthropogenic change, and provide examples of how it can result in phenotypically distinct units with different, socially learnt behavioural strategies. While identifying culture and social learning can be challenging, indirect identification and parsimonious inferences may be informative. Finally, we identify relevant methodologies and provide a framework for viewing behavioural data through a cultural lens which might provide new insights for conservation management.

Short-range hunters: exploring the function and constraints of water shooting in dwarf gouramis.

Ballistic predation is a rare foraging adaptation: in fishes, most attention has focused on a single genus, the archerfish, known to manipulate water to shoot down prey above the water surface. However, several gourami species also exhibit apparently similar 'shooting' behaviour, spitting water up to 5 cm above the surface. In a series of experiments, we explored the shooting behaviour and aspects of its significance as a foraging ability in the dwarf gourami (Trichogaster lalius). We investigated sex differences in shooting abilities to determine whether gourami shooting is related to the sex-specific bubble nest manufacture where males mix air and water at the surface to form bubbles. We found that, actually, both sexes were equally able to shoot and could learn to shoot a novel target. In a second experiment, we presented untrained gouramis with opportunities to shoot at live prey and found they successfully shot down both fruit flies and crickets. Finally, we explored the effect of target height on shooting performance to establish potential constraints of shooting as a foraging ability. The frequency of attempted shots and success of hitting targets decreased with height, whereas latency to shoot increased. We also observed that repeatable individual differences account for variation in these measures of shooting performance. Together, our results provide evidence that gourami shooting has a foraging function analogous to that of archerfish. Gourami shooting may serve as an example of convergent evolution and provide opportunities for comparative studies into the, as yet unexplored, ecology and evolution of shooting in fishes.

Adaptation of sperm whales to open-boat whalers: rapid social learning on a large scale?

Animals can mitigate human threats, but how do they do this, and how fast can they adapt? Hunting sperm whales was a major nineteenth century industry. Analysis of data from digitized logbooks of American whalers in the North Pacific found that the rate at which whalers succeeded in harpooning ('striking') sighted whales fell by about 58% over the first few years of exploitation in a region. This decline cannot be explained by the earliest whalers being more competent, as their strike rates outside the North Pacific, where whaling had a longer history, were not elevated. The initial killing of particularly vulnerable individuals would not have produced the observed rapid decline in strike rate. It appears that whales swiftly learned effective defensive behaviour. Sperm whales live in kin-based social units. Our models show that social learning, in which naive social units, when confronted by whalers, learned defensive measures from grouped social units with experience, could lead to the documented rapid decline in strike rate. This rapid, large-scale adoption of new behaviour enlarges our concept of the spatio-temporal dynamics of non-human culture.

A boldly comparative approach will strengthen co-evolutionary accounts of musicality's origins.

Focus on the evolutionary origins of musicality has been neglected relative to attention on language, so these new proposals are welcome stimulants. We argue for a broad comparative approach to understanding how the elements of musicality evolved, and against the use of overly simplistic evolutionary accounts.

Song hybridization events during revolutionary song change provide insights into cultural transmission in humpback whales.

Cultural processes occur in a wide variety of animal taxa, from insects to cetaceans. The songs of humpback whales are one of the most striking examples of the transmission of a cultural trait and social learning in any nonhuman animal. To understand how songs are learned, we investigate rare cases of song hybridization, where parts of an existing song are spliced with a new one, likely before an individual totally adopts the new song. Song unit sequences were extracted from over 9,300 phrases recorded during two song revolutions across the South Pacific Ocean, allowing fine-scale analysis of composition and sequencing. In hybrid songs the current and new songs were spliced together in two specific ways: (i) singers placed a single hybrid phrase, in which content from both songs were combined, between the two song types when transitioning from one to the other, and/or (ii) singers spliced complete themes from the revolutionary song into the current song. Sequence analysis indicated that both processes were governed by structural similarity rules. Hybrid phrases or theme substitutions occurred at points in the songs where both songs contained "similar sounds arranged in a similar pattern." Songs appear to be learned as segments (themes/phrase types), akin to birdsong and human language acquisition, and these can be combined in predictable ways if the underlying structural pattern is similar. These snapshots of song change provide insights into the mechanisms underlying song learning in humpback whales, and comparative perspectives on the evolution of human language and culture.

Vocal sequences in narwhals (Monodon monoceros).

Sequences are indicative of signal complexity in vocal communication. While vocal sequences are well-described in birds and terrestrial mammals, the extent to which marine mammals use them is less well understood. This study documents the first known examples of sequence use in the narwhal (Monodon monoceros), a gregarious Arctic cetacean. Eight female narwhals were fitted with animal-borne recording devices, resulting in one of the largest datasets of narwhal acoustic behaviour to date. A combination of visual and quantitative classification procedures was used to test whether subjectively defined vocalization patterns were organized into sequences. Next, acoustic characteristics were analyzed to assess whether sequences could disclose group or individual identity. Finally, generalized linear models were used to investigate the behavioural context under which sequences were produced. Two types of sequences, consisting of "paired" patterns and "burst pulse series," were identified. Sequences of burst pulse series were typically produced in periods of high vocal activity, whereas the opposite was true for sequences of paired patterns, suggesting different functions for each. These findings extend the set of odontocetes which are known to use vocal sequences. Inquiry into vocal sequences in other understudied marine mammals may provide further insights into the evolution of vocal communication.

A framework for auralization of boundary element method simulations including source and receiver directivity.

The Boundary Element Method (BEM) is a proven numerical prediction tool for computation of room acoustic transfer functions, as are required for auralization of a virtual space. In this paper, it is validated against case studies drawn from the "Ground Truth for Room Acoustical Simulation" database within a framework that includes source and receiver directivity. These aspects are often neglected but are respectively important to include for auralisation applications because source directivity is known to affect how a room is excited and because the human auditory system is sensitive to directional cues. The framework uses weighted-sums of spherical harmonic functions to represent both the source directivity to be simulated and the pressure field predicted in the vicinity of the receiver location, the coefficients of the former being fitted to measured directivity and those of the latter computed directly from the boundary data by evaluating a boundary integral. Three validation cases are presented, one of which includes a binaural receiver. The computed results match measurements closely for the two cases conducted in anechoic conditions but show some significant differences for the third room scenario; here, it is likely that uncertainty in boundary material data limited modelling accuracy.

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.

Short first click intervals in echolocation trains of three species of deep diving odontocetes.

All odontocetes produce echolocation clicks as part of their vocal repertoire. In this paper the authors analysed inter-click-intervals in recordings from suction cup tags with a focus on the first inter-click interval of each click train. The authors refer to shorter first inter-click intervals as short first intervals (SFIs). The authors found that the context of SFI occurrence varies across three deep-diving species. In Blainville's beaked whales, 87% of click trains that were preceded by a terminal buzz started with SFIs. In Cuvier's beaked whales, only sub-adult animals produced notable amounts of SFIs. In contrast, sperm whales were much more likely to produce SFIs on the first click train of a dive. While the physiological and/or behavioural reasons for SFI click production are unknown, species differences in their production could provide a window into the evolution of odontocete echolocation.

The devil is in the detail: Quantifying vocal variation in a complex, multi-levelled, and rapidly evolving display.

Identifying and quantifying variation in vocalizations is fundamental to advancing our understanding of processes such as speciation, sexual selection, and cultural evolution. The song of the humpback whale (Megaptera novaeangliae) presents an extreme example of complexity and cultural evolution. It is a long, hierarchically structured vocal display that undergoes constant evolutionary change. Obtaining robust metrics to quantify song variation at multiple scales (from a sound through to population variation across the seascape) is a substantial challenge. Here, the authors present a method to quantify song similarity at multiple levels within the hierarchy. To incorporate the complexity of these multiple levels, the calculation of similarity is weighted by measurements of sound units (lower levels within the display) to bridge the gap in information between upper and lower levels. Results demonstrate that the inclusion of weighting provides a more realistic and robust representation of song similarity at multiple levels within the display. This method permits robust quantification of cultural patterns and processes that will also contribute to the conservation management of endangered humpback whale populations, and is applicable to any hierarchically structured signal sequence.

Archerfish foraging success varies with immediate competition level but not group size.

Group living can lead to kleptoparasitism, the theft of resources by competitors. Under such conditions, foragers may alter their behavior to minimize competition. However, it is unclear how such behavioral changes impact foraging performance. Archerfish (Toxotes spp.) are a good model for investigating the behavioral responses to kleptoparasitism, as their hunting method (shooting waterjets at insects perched above the water) leaves them vulnerable to theft. They must hit the target prey with sufficient force to dislodge it; thus, the prey may land some distance away from the shooter. Kleptoparasitism rates increase with group size in archerfish, and individuals alter their behavior around conspecifics. We investigated whether group size affected shooting success, using 7-spot archerfish T. chatareus. We considered a fish's shot to be successful if it knocked a fly, placed on a transparent platform above the tank, into the water. The probability of shooting success was modeled as a function of group size, aiming duration, nearest neighbor distance and position, and trial number. We found no effect of group size, aiming duration, or nearest neighbor distance or position on shooting success. Shooting success increased as trials progressed, likely due to the fish becoming more familiar with the task. We also found no change in the kleptoparasitism rate between group sizes. Instead, the likelihood of the shooter consuming the prey depended on the types of competition present at the time of shooting. We suggest that archerfish shooting behavior can be influenced by the presence of conspecifics in ways not previously considered.

Can genetic differences explain vocal dialect variation in sperm whales, Physeter macrocephalus?

Sperm whale social groups can be assigned to vocal clans based on their production of codas, short stereotyped patterns of clicks. It is currently unclear whether genetic variation could account for these behavioural differences. We studied mitochondrial DNA (mtDNA) variation among sympatric vocal clans in the Pacific Ocean, using sequences extracted from sloughed skin samples. We sampled 194 individuals from 30 social groups belonging to one of three vocal clans. As in previous studies of sperm whales, mtDNA control region diversity was low (π = 0.003), with just 14 haplotypes present in our sample. Both hierarchical AMOVAs and partial Mantel tests showed that vocal clan was a more important factor in matrilineal population genetic structure than geography, even though our sampling spanned thousands of kilometres. The variance component attributed to vocal dialects (7.7%) was an order of magnitude higher than those previously reported in birds, while the variance component attributed to geographic area was negligible. Despite this, the two most common haplotypes were present in significant quantities in each clan, meaning that variation in the control region cannot account for behavioural variation between clans, and instead parallels the situation in humans where parent-offspring transmission of language variation has resulted in correlations with neutral genes. Our results also raise questions for the management of sperm whale populations, which has traditionally been based on dividing populations into geographic 'stocks', suggesting that culturally-defined vocal clans may be more appropriate management units.

Ocean nomads or island specialists? Culturally driven habitat partitioning contrasts in scale between geographically isolated sperm whale populations.

The sperm whale (Physeter macrocephalus) is a deep-diving cetacean with a global distribution and a multi-leveled, culturally segregated, social structure. While sperm whales have previously been described as 'ocean nomads', this might not be universal. We conducted surveys of sperm whales along the Lesser Antilles to document the acoustic repertoires, movements and distributions of Eastern Caribbean (EC) sperm whale cultural groups (called vocal clans). In addition to documenting a potential third vocal clan in the EC, we found strong evidence of fine-scale habitat partitioning between vocal clans with scales of horizontal movements an order of magnitude smaller than from comparable studies on Eastern Tropical Pacific sperm whales. These results suggest that sperm whales can display cultural ecological specialization and habitat partitioning on flexible spatial scales according to local conditions and broadens our perception of the ecological flexibility of the species. This study highlights the importance of incorporating multiple temporal and spatial scales to understand the impact of culture on ecological adaptability, as well as the dangers of extrapolating results across geographical areas and cultural groups.

Distinctive, fine-scale distribution of Eastern Caribbean sperm whale vocal clans reflects island fidelity rather than environmental variables.

Environmental variables are often the primary drivers of species' distributions as they define their niche. However, individuals, or groups of individuals, may sometimes adopt a limited range within this larger suitable habitat as a result of social and cultural processes. This is the case for Eastern Caribbean sperm whales. While environmental variables are reasonably successful in describing the general distribution of sperm whales in the region, individuals from different cultural groups have distinct distributions around the Lesser Antilles islands. Using data collected over 2 years of dedicated surveys in the Eastern Caribbean, we conducted habitat modeling and habitat suitability analyses to investigate the mechanisms responsible for such fine-scale distribution patterns. Vocal clan-specific models were dramatically more successful at predicting distribution than general species models, showing how a failure to incorporate social factors can impede accurate predictions. Habitat variation between islands did not explain vocal clan distributions, suggesting that cultural group segregation in the Eastern Caribbean sperm whale is driven by traditions of site/island fidelity (most likely maintained through conformism and homophily) rather than habitat type specialization. Our results provide evidence for the key role of cultural knowledge in shaping habitat use of sperm whales within suitable environmental conditions and highlight the importance of cultural factors in shaping sperm whale ecology. We recommend that social and cultural information be incorporated into conservation and management as culture can segregate populations on fine spatial scales in the absence of environmental variability.

From Beethoven to Beyoncé: Do Changing Aesthetic Cultures Amount to "Cumulative Cultural Evolution?"

Culture can be defined as "group typical behaviour patterns shared by members of a community that rely on socially learned and transmitted information" (Laland and Hoppitt, 2003, p. 151). Once thought to be a distinguishing characteristic of humans relative to other animals (Dean et al., 2014) it is now generally accepted to exist more widely, with especially abundant evidence in non-human primates, cetaceans, and birds (Rendell and Whitehead, 2001; Aplin, 2019; Whiten, 2021). More recently, cumulative cultural evolution (CCE) has taken on this distinguishing role (Henrich, 2015; Laland, 2018). CCE, it is argued, allows humans, uniquely, to ratchet up the complexity or efficiency of cultural traits over time. This "ratchet effect" (Tomasello, 1994) gives the capacity to accumulate beneficial modifications over time beyond the capacities of a single individual (Sasaki and Biro, 2017). Mesoudi and Thornton (2018) define a core set of criteria for identifying CCE in humans and non-human animals that places emphasis on some performance measure of traits increasing over time. They suggest this emphasis is also pertinent to cultural products in the aesthetic domain, but is this the case? Music, art and dance evolve over time (Savage, 2019), but can we say they gain beneficial modifications that increase their aesthetic value? Here we bring together perspectives from philosophy, musicology and biology to build a conceptual analysis of this question. We summarise current thinking on cumulative culture and aesthetics across fields to determine how aesthetic culture fits into the concept of CCE. We argue that this concept is problematic to reconcile with dominant views of aesthetics in philosophical analysis and struggles to characterise aesthetic cultures that evolve over time. We suggest that a tension arises from fundamental differences between cultural evolution in aesthetic and technological domains. Furthermore, this tension contributes to current debates between reconstructive and preservative theories of cultural evolution.

Measuring inter-pulse intervals in sperm whale clicks: consistency of automatic estimation methods.

Sperm whale clicks are characterized by a multi-pulsed structure. The time lag between consecutive pulses, i.e., the inter-pulse interval (IPI), is related to the size of the sound production organ such that its measurement provides a means to acoustically estimate the size of individual whales. Due to off-axis effects the identification of pulses is, however, not always straightforward, and automatic measurement methods provide not only more objective estimation, but may also facilitate IPI estimation in cases where single click measurements are ambiguous. In particular, averaging measurements over a time series of clicks from the same whale could enhance the discrimination of time invariant pulses. The authors developed two automatic methods of automatic IPI measurement based on waveform and autocorrelation averaging and compared their accuracy and consistency with other previously used methods. Manual measurement by an experienced operator provided the most self-consistent estimates. The autocorrelation averaging technique had the best overall performance of the automated methods achieving a very similar performance to manual measurement. On some recordings cepstrum averaging methods converged when autocorrelation did not. Therefore, applying both of these automated methods and choosing the best of the two are recommended.

Flexible learning, rather than inveterate innovation or copying, drives cumulative knowledge gain.

Human technology is characterized by cumulative cultural knowledge gain, yet researchers have limited knowledge of the mix of copying and innovation that maximizes progress. Here, we analyze a unique large-scale dataset originating from collaborative online programming competitions to investigate, in a setting of real-world complexity, how individual differences in innovation, social-information use, and performance generate technological progress. We find that cumulative knowledge gain is primarily driven by pragmatists, willing to copy, innovate, explore, and take risks flexibly, rather than by pure innovators or habitual copiers. Our study also reveals a key role for prestige in information transfer.