Photographic mark-recapture analysis of clustered mammal-eating killer whales around the Aleutian Islands and Gulf of Alaska

We used photographic mark-recapture methods to estimate the number of mammal-eating “transient” killer whales using the coastal waters from the central Gulf of Alaska to the central Aleutian Islands, around breeding rookeries of endangered Steller sea lions. We identified 154 individual killer whales from 6,489 photographs collected between July 2001 and August 2003. A Bayesian mixture model estimated seven distinct clusters (95% probability interval = 7–10) of individuals that were differentially covered by 14 boat-based surveys exhibiting varying degrees of association in space and time. Markov Chain Monte Carlo methods were used to sample identification probabilities across the distribution of clusters to estimate a total of 345 identified and undetected whales (95% probability interval = 255–487). Estimates of covariance between surveys, in terms of their coverage of these clusters, indicated spatial population structure and seasonal movements from these near-shore waters, suggesting spatial and temporal variation in the predation pressure on coastal marine mammals.     

Estimating abundance of killer whales in the nearshore waters of the Gulf of Alaska and Aleutian Islands using line-transect sampling

Killer whale (Orcinus orca Linnaeus, 1758) abundance in the North Pacific is known only for a few populations for which extensive longitudinal data are available, with little quantitative data from more remote regions. Line-transect ship surveys were conducted in July and August of 2001–2003 in coastal waters of the western Gulf of Alaska and the Aleutian Islands. Conventional and Multiple Covariate Distance Sampling methods were used to estimate the abundance of different killer whale ecotypes, which were distinguished based upon morphological and genetic data. Abundance was calculated separately for two data sets that differed in the method by which killer whale group size data were obtained. Initial group size (IGS) data corresponded to estimates of group size at the time of first sighting, and post-encounter group size (PEGS) corresponded to estimates made after closely approaching sighted groups. ‘Resident’-type (fish-eating) killer whales were more abundant than the ‘transient’-type (mammal-eating). Abundance estimates of resident killer whales (991 [95% CI = 379–2,585] [IGS] and 1,587 [95% CI = 608–4,140] [PEGS]), were at least four times greater than those of the transient killer whales (200 [95% CI = 81–488] [IGS] and 251 [95% CI = 97–644] whales [PEGS]). The IGS estimate of abundance is preferred for resident killer whales because the estimate based on PEGS data may show an upward bias. The PEGS estimate of abundance is likely more accurate for transients. Residents were most abundant near Kodiak Island in the northern Gulf of Alaska, around Umnak and Unalaska Islands in the eastern Aleutians, and in Seguam Pass in the central Aleutians. This ecotype was not observed between 156 and 164°W, south of the Alaska Peninsula. In contrast, transient killer whale sightings were found at higher densities south of the Alaska Peninsula between the Shumagin Islands and the eastern Aleutians. Only two sightings of ‘offshore’-type killer whales were recorded during the surveys, one northeast of Unalaska Island and the other south of Kodiak Island. These are the first estimates of abundance of killer whale ecotypes in the Aleutian Islands and Alaska Peninsula area and provide a baseline for quantifying the role of these top predators in their ecosystem. Electronic Supplementary Material  Supplementary material is available in the online version of this article at and is accessible for authorized users.

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Vocalizations of the North Atlantic pilot whale (Globicephala melas) as related to behavioral contexts

Summary Vocalizations of free-ranging North Atlantic pilot whales were studied in different behavioral contexts to gain insight into the function and biological significance of different sound types. Simple whistles (with no frequency inflections) were heard more frequently when whales were milling, a restful behavior type. During surface active behavior, energetic, often coordinated activity probably representing feeding, many sound types, especially complex whistles (with more frequency inflections) and pulsed sounds, occurred with greater frequency than when this behavior was absent. Greater numbers of most whistle types were produced when whales were spread over a larger area and when more subgroups were present. Thus, in pilot whales, there is a significant relationship between their sounds and their behavior, with vocalizations possibly serving to maintain contact and coordinate movements of the herd. Offprint requests to: L.S. Weilgart at the current address     

Signature whistles of free-ranging bottlenose dolphins Tursiops truncatus: stability and mother-offspring comparisons

Summary Mother-calf whistle exchanges were recorded from temporarily captured free-ranging bottlenose dolphins from 1975 to 1989. This is part of a long-term research project studying social structure and behavior of a community of approximately 100 dolphins in waters near Sarasota, Florida. Analysis of whistle exchanges from 12 mothercalf pairs shows that signature whistles can remain stable for periods up to at least 12 years. We looked for effects of vocal learning on the development of the signature whistle by comparing whistles of calves to those of their mothers. Eight female calves produced whistles distinct from those of their mothers, while four male calves produced whistles similar to those of their mothers. Male calves appeared to produce a greater proportion of whistles other than the signature whistle (termed variants). We hypothesize that these sex differences in whistle vocalizations may reflect differences in the roles males and females play in the social structure of the community.     

Signature whistle variations in a bottlenosed dolphin,Tursiops truncatus

To examine whether context-specific information is superimposed upon the individual cues present in the whistling of the bottlenosed dolphin, Tursiops truncatus, parameter variations within the two most frequently emitted whistle types of a captive individual were investigated in three different behavioural contexts. The study concentrated on comparing signal features of spontaneously occurring vocalizations in two possible phases following the performance of a trained discrimination task and those occurring during isolation. Phases of the discrimination task differed according to whether the animal showed correct (reward given) or incorrect performance (no reward). Signature whistles were most common in isolation, but also represented just over half of the whistles following a choice task. Of 14 signature whistle frequency and time parameters measured 9 differed significantly between isolation and at least one of the phases following a choice task (Table 1). Three parameters also varied according to whether performance was correct or incorrect. In contrast, only one out of four parameters (start frequency) measured from the second most frequent whistle type varied significantly between contexts (isolation vs. phase following correct choice). The results indicate that not only identity but also context-related information is available in the whistles of a bottlenosed dolphin.     

Use of signature whistles during separations and reunions by wild bottlenose dolphin mothers and infants

We examine the contexts and patterns of signature whistle production by wild bottlenose dolphin mother-infant pairs (Tursiops spp.) to gain insight into the functional significance of whistles. Results are based on focal observations and simultaneous recordings of underwater vocalizations. Whistles occur primarily when mother-infant pairs are separated, and the probability of whistles increases with distance of separation. The timing of whistles during separations varies, but whistles tend to be produced in repetitive series and are generally concentrated toward the later stages of the separation, i.e., during the process of reunion. Although we focused on infants, mothers do not appear to whistle during separations as frequently as infants. Infant whistles may function to facilitate reunions by conveying information to the mother concerning the infant's motivation to reunite and/or its location. Infant whistles could induce a cooperative response from the mother including approach, slowing to allow the infant to catch up or whistling. Highly individualized signature whistles may be particularly useful in a fission-fusion society in which individuals (mothers and infants as well as adults) join and leave temporary parties in a fluid manner, yet maintain consistent, long-term associations with particular individuals. Correspondence to: R.A. Smolker     

Pups crying bass: vocal adaptation for avoidance of age-dependent predation risk in ground squirrels?

In most mammals, larger adult body size correlates with lower fundamental frequency and more closely spaced formants in vocalizations relative to juveniles. In alarm whistles of two free-living rodents, the speckled ground squirrel Spermophilus suslicus and yellow ground squirrel S. fulvus, these cues to body size were absent despite prominent differences in body weight and skull and larynx sizes between juveniles and adults. No significant correlations were found between the individual maximum fundamental frequency and body weight, both within age classes and for pooled samples of all animals within species. Furthermore, the mean alarm whistle maximum fundamental frequencies did not differ significantly between age classes (juvenile versus adult) in the speckled squirrel and were even significantly lower in juvenile yellow squirrels. We discuss the hypothesis that the obfuscation of vocal differences between juvenile and adult squirrels may represent a special adaptation of pup vocal behaviour—a form of “vocal mimicry,” resulting in imitation of adult vocal pattern to avoid infanticide and age-dependent predation risk.     

Territory ownership and familiarity status affect how much male root voles (Microtus oeconomus) invest in territory defence

Neighbour–stranger discrimination occurs when individuals respond with more aggression to strangers than to territorial neighbours—a phenomenon termed the “dear enemy phenomenon” (DEP). We investigated the DEP with male and female root voles (Microtus oeconomus Pallas 1776) using field dyadic arena tests conducted in enclosures where we could test for the effects of familiarity (familiar versus stranger), ownership (resident versus intruder status) and resource-holding potential (body mass) on territorial behaviours. The results showed that males put more effort into territorial defence than females, and males could discriminate between neighbours and strangers. In males, aggressiveness was influenced by a significant two-way interaction between treatment and ownership. Male residents were more aggressive towards stranger intruders than towards neighbour intruders, while male intruders were less aggressive towards stranger residents than towards neighbour residents. In females, neither treatment nor ownership status had a significant effect on aggressiveness. Familiar males performed more social behaviours but less non-social behaviours than stranger males. Furthermore, there was a clear dominance hierarchy between residents and intruders in stranger dyads, with the male territory holders dominating the intruder in pairwise interactions. To our knowledge, these results demonstrate for the first time DEP in a small mammal with a known pedigree and present the first evidence for “prior resident advantage” in voles. We argue that both ownership status and familiarity status affect how much an individual invests in territory defence. The benefits of neighbour–stranger discrimination for male root voles and the absence of neighbour–stranger discrimination in female root voles are discussed.     

Whistle sharing in paired male bottlenose dolphins,<Emphasis Type="Italic"> Tursiops truncatus</Emphasis>

The signature whistle hypothesis states that dolphins produce highly stereotyped, individually distinctive whistles when in isolation. The presence of signature whistles has been called into question by recent studies proposing that dolphins produce a shared, simple upsweep whistle when in isolation, and that whistles produced by socializing dolphins are shared across individuals and social groups. This shared repertoire hypothesis suggests that when two animals produce the same whistle type, it is due to sharing the same common repertoire rather than one animal learning to produce the whistle of another. One difference between studies supporting or denying the existence of signature whistles is the method used to classify whistle types. We examined whistle production by 17 free-ranging bottlenose dolphins while temporarily restrained. We used both a quantitative comparison technique similar to that used to support the shared repertoire hypothesis and human judges to classify whistle types and quantify similarity between types. Contrary to recent studies that emphasize shared whistles, overall whistle sharing between isolated individuals was low (25%) and a simple upsweep did not account for the most common whistle type in half of the animals. Some species of birds, bats, and primates with stable social groups use vocal learning to converge over time to one common group distinctive call type. We examined whistle similarity between adult male dolphins that are partners in a close social alliance in order to test whether vocal learning may enable a similar vocal convergence. Whistle similarity was rated very high between partners and low between non-partners by both the quantitative technique and human observers. This suggests that as in songbirds and some other mammals, adult male bottlenose dolphins may use vocal learning to converge on similar whistles as they develop affiliative social relationships.Communicated by G. Wilkinson     

Whistle repertoires of two bottlenosed dolphins,Tursiops truncatus: mimicry of signature whistles?

Summary The whistle vocalizations of two bottlenosed dolphins, Tursiops truncatus, were recorded at the Sealand Aquarium in Brewster, Massachusetts. The identification of which dolphin within the group produced a vocalization was made possible by a telemetry device attached to the dolphin's head with a suction cup. 77% of the identified whistles (219 our of 284) fell into two primary categories, type 1 and type 2 (Table 1). The remaining 23% of whistles fell into five secondary categories. Of the primary whistles produced by one dolphin, 78% were of type 1 (22% type 2), while 69% of primary whistles from the other dolphin were of type 2 (31% type 1). The result that each of the dolphins favored a different primary whistle supports the findings of Caldwell and Caldwell (1965), that each dolphin produces an individually distinctive whistle. But in the present study, both dolphins produced both primary whistle types. This may represent mimicry of signature whistles.     

Spatial and social connectivity of fish-eating “Resident” killer whales (Orcinus orca) in the northern North Pacific

The productive North Pacific waters of the Gulf of Alaska, Aleutian Islands and Bering Sea support a high density of fish-eating “Resident” type killer whales (Orcinus orca), which overlap in distribution with commercial fisheries, producing both direct and indirect interactions. To provide a spatial context for these interactions, we analyzed a 10-year dataset of 3,058 whale photo-identifications from 331 encounters within a large (linear ~4,000 km) coastal study area to investigate the ranging and social patterns of 532 individually identifiable whales photographed in more than one encounter. Although capable of large-scale movements (maximum 1,443 km), we documented ranges generally <200 km, with high site fidelity across summer sampling intervals and also re-sightings during a winter survey. Bayesian analysis of pair-wise associations identified four defined clusters, likely representing groupings of stable matrilines, with distinct ranging patterns, that combined to form a large network of associated whales that ranged across most of the study area. This provides evidence of structure within the Alaska stock of Resident killer whales, important for evaluating ecosystem and fisheries impacts. This network included whales known to depredate groundfish from longline fisheries, and we suggest that such large-scale connectivity has facilitated the spread of depredation.     

Individual dispersal status influences space use of conspecific residents in the common lizard, Lacerta vivipara

The effects of immigration on the behaviour of residents may have important implications for the local population characteristics. A manipulative laboratory experiment with yearlings of the common lizard (Lacerta vivipara) was performed to test whether the introduction of dispersing or philopatric individuals influences the short-term spacing behaviour of resident individuals. Staged encounters were carried out to induce interactions within dyads. The home cage of each responding individual was connected by a corridor to an unfamiliar “arrival cage” to measure the latency to leave their own home cage after each encounter. Our results showed that the time that pairs spent in close proximity was longer when a dispersing individual was introduced in the home cage. The latency to leave the home cage was longer after the introduction of a dispersing individual. These response variables were not influenced by the relative body sizes of contestants nor by the level of aggression towards each other. In contrast, the aggressive response was significantly influenced by the residency asymmetry established experimentally (“owner” of the home cage vs introduced individual). Our results suggest that the space use by resident individuals is influenced by the dispersal status of conspecifics. The potential ultimate causes driving this effect are discussed.     

Polygyny in the red-winged blackbird: do females prefer monogamy or polygamy?

Five hypotheses have been proposed to explain polygyny in the red-winged blackbird (Agelaius phoeniceus). We categorized the hypotheses into three groups based on female preference for unmated versus monogamously mated males: (1) the “polygyny threshold” model, “sexy son” hypothesis and the “asynchronous settlement” model, which assume that females prefer unmated males to mated males on breeding situations of homogeneous quality; (2) the “neutral mate choice” hypothesis, which assumes that females have no preference; and (3) the “cooperative female choice” model, which assumes that females prefer monogamously mated males to unmated males. We tested the direction of female preference in two field experiments. In both experiments, newly settling females were given a choice of two adjacent territories, one defended by an unmated male and the other by a monogamously mated male. Male mating status was randomized with respect to the variation in territory quality and male quality. Early in the breeding season, significantly more females settled with the unmated males than with the mated males. Although more females settled with the unmated males than with the mated males late in the breeding season, the difference was no longer significant. Female settlement late in the season appeared to be related to the tenure of the resident females: the new females avoided territories where the resident females were in early stages of their nesting, but settled on territories where the resident females were in late stages. The pattern of female settlement shows that females prefer unmated males to mated males. The preference is consistent with the polygyny threshold model, sexy son hypothesis and the asynchronous settlement model, and inconsistent with the neutral mate choice hypothesis and the cooperative female choice model. For this reason, the latter two hypotheses are unlikely to explain the occurrence of polygyny in our population of red-winged blackbirds. Received: 1 December 1994 / Accepted after revision: 28 October 1995     

Production and perception of situationally variable alarm calls in wild tufted capuchin monkeys (Cebus apella nigritus)

Many mammalian and avian species produce conspicuous vocalizations upon encountering a predator, but vary their calling based on risk urgency and/or predator type. Calls falling into the latter category are termed “functionally referential” if they also elicit predator-appropriate reactions in listeners. Functionally referential alarm calling has been well documented in a number of Old World monkeys and lemurs, but evidence among Neotropical primates is limited. This study investigates the alarm call system of tufted capuchin monkeys (Cebus apella nigritus) by examining responses to predator and snake decoys encountered at various distances (reflecting differences in risk urgency). Observations in natural situations were conducted to determine if predator-associated calls were given in additional contexts. Results indicate the use of three call types. “Barks” are elicited exclusively by aerial threats, but the call most commonly given to terrestrial threats (the “hiccup”) is given in nonpredatory contexts. The rate in which this latter call is produced reflects risk urgency. Playbacks of these two call types indicate that each elicits appropriate antipredator behaviors. The third call type, the “peep,” seems to be specific to terrestrial threats, but it is unknown if the call elicits predator-specific responses. “Barks” are thus functionally referential aerial predator calls, while “hiccups” are better seen as generalized disturbance calls which reflect risk urgency. Further evidence is needed to draw conclusions regarding the “peep.” These results add to the evidence that functionally referential aerial predator alarm calls are ubiquitous in primates, but that noncatarrhine primates use generalized disturbance calls in response to terrestrial threats.     

Culturally correct song dialects are correlated with male age and female song preferences in wild populations of brown-headed cowbirds

Variation in vocal behavior among local populations of songbirds may be significant to females in mate choice. In a study of dialect populations of brown-headed cowbirds, estradiol-implanted females from two dialects held the pre-copulatory lordosis posture longer in response to playback of the local flight whistle song than to foreign whistle types from adjacent and more distant dialects. Females were held in captivity for a relatively brief period prior to testing and received no tutoring so discrimination was based solely on experience in the wild. This is the first study to show evidence of discrimination by female cowbirds based on flight whistle type. Evidence is presented from one of the study dialects that the majority of yearling male cowbirds are vocally distinct from resident adults in having either foreign or incomplete local whistles. Although these yearlings are fully mature sexually, they have little or no mating success. The correlation between female whistle preference and male mating success suggests that the ability to give the correct local whistle type may be a characteristic used by females to assess age and quality of a potential mate. A learned female preference for the predominant local song type may be a factor in the stability of these dialects by making it adaptive for males to conform to the local dialect.     

Darwin’s “tug-of-war” vs. starlings’ “horse-racing”: how adaptations for sequential encounters drive simultaneous choice

Charles Darwin aided his private decision making by an explicit deliberation, famously deciding whether or not to marry by creating a list of points in a table with two columns: “Marry” and “Not Marry”. One hundred seventy-two years after Darwin’s wedding, we reconsider whether this process of choice, under which individuals assign values to their options and compare their relative merits at the time of choosing (the tug-of-war model), applies to our experimental animal, the European Starling, Sturnus vulgaris. We contrast this with the sequential choice model that postulates that decision-makers make no comparison between options at the time of choice. According to the latter, behaviour in simultaneous choices reflects adaptations to contexts with sequential encounters, in which the choice is whether to take an opportunity or let it pass. We postulate that, in sequential encounters, the decision-maker assigns (by learning) a subjective value to each option, reflecting its payoff relative to background opportunities. This value is expressed as latency and/or probability to accept each opportunity as opposed to keep searching. In simultaneous encounters, choice occurs through each option being processed independently, by a race between the mechanisms that generate option-specific latencies. We describe these alternative models and review data supporting the predictions of the sequential choice model.     

Predator stimuli and calling behavior of Carolina chickadees (Poecile carolinensis), tufted titmice (Baeolophus bicolor), and white-breasted nuthatches (Sitta carolinensis)

Evidence from different chickadee species (Poecile genus) indicates that birds can modify the note composition of their “chick-a-dee” calls in the presence of predator stimuli. Here, we tested the effects of predator models and the distance of those models on calls of three species foraging together at feeding stations: Carolina chickadees (Poecile carolinensis) and tufted titmice (Baeolophus bicolor), both members of the Paridae family, and white-breasted nuthatches (Sitta carolinensis), a member of the Sittidae family. Model and distance affected seed-taking rates in all three species. “Chick-a-dee” calling rates were higher in the predator context for both chickadees and titmice, but we detected no predator context effects on “quank” call rates for nuthatches. Predator and distance contexts affected acoustic parameters of notes of the “chick-a-dee” calls of chickadees and titmice; no such effects were detected for nuthatch “quank” calls. These results suggest species differences in encoding of information in the primary social calls of these three species that commonly occur in multi-species flocks. Chickadees and titmice are “nuclear” species and nuthatches are “satellite” species, and these different roles might be related to the differences in vocal signaling that we detected.     

Sex difference in signature whistle production of free-ranging bottlenose dolphins,Tursiops truncates

Signature whistles of 42 free-ranging bottle-nose dophin calves were compared to those of their mothers. Humans judged their similarity by inspection of spectrograms. There was a sex difference in the tendency of calves to produce whistles similar to or different from those of their mothers; most female calves produced whistles that were different from those of their mothers, whereas male calves were more likely to produce whistles that were similar to those of their mothers. Because matrilineally related females associate together and use signature whistles to establish and/or maintain contact with their calves, there may be a selective pressure for females to produce whistles that are distinct from those of their mothers. There may be fewer constraints governing whistle development in males, with the result that some males produce whistles similar to those of their mothers and others do not.     

Mating system and individual reproductive success of sympatric anadromous and resident brook charr, <Emphasis Type="Italic">Salvelinus fontinalis</Emphasis>, under natural conditions

Salmonids are known for the occurrence in sympatry of two life-history forms, one that undergoes migration to sea before returning to freshwater to reproduce (anadromous) and one that inhabits freshwater without a migration phase (resident). Whereas one breeding population is often suggested by population genetic studies, mating patterns have rarely been directly assessed, especially when both sexes are found within each life-history form. By using highly polymorphic microsatellite loci and parentage analysis in a natural population of sympatric anadromous and resident brook charr (Salvelinus fontinalis), we found that gene flow occurred between the two forms and was mediated by resident males mating with both resident and anadromous females. Determinants of reproductive success, estimated by the number of surviving juveniles (ages 1 and 2 years), differed between the sexes. No strong evidence of the influence of size on individual reproductive success was found for males, whereas larger females (and hence most likely to be anadromous) were more successful. The higher individual reproductive success of anadromous fish compared to residents was mainly explained by this higher reproductive success of anadromous females. We suggest that resident males adopt a “sneaking” reproductive tactic as a way of increasing their reproductive success by mating with females of all sizes in all habitats. The persistence of the resident tactic among females may be linked to their advantage in accessing spatially constrained spawning areas in small tributary streams unavailable to larger females.     

Alarm calls of Belding's ground squirrels to aerial predators: nepotism or self-preservation?

Summary Belding's ground squirrels (Spermophilus beldingi) give acoustically distinct alarm calls to aerial and terrestrial predators. The animals typically give multiple-note trills to predatory mammals, and single-note whistles to flying hawks. During a 9-year study of free-living S. beldingi at Tioga Pass, California, the adaptive significance of the whistle call was investigated. Data were gathered on 664 ground squirrel-hawk interactions, most of which were induced by flying trained raptors over individually marked study animals of known sex and age. The sight of a flying hawk and the sound of whistles stimulated widespread calling and running to shelter by the ground squirrels (Fig. 1). Wild raptors were rarely successful at capturing the rodents once a whistle had been given, and fewer callers than noncallers were killed (Table 1). Individuals of both sexes and all ages whistled equally often (Fig. 4), and females' tendencies to whistle were not affected by the presence of relatives, including offspring (Fig. 5). The most frequent callers were animals in exposed positions: far from cover and close to the predatory bird (Table 2). Taken together the data suggest that unlike trills, which increase vulnerability to terrestrial predators (Table 1) and function to warn relatives, whistle directly benefit callers by increasing their chances of escaping from hawks.