Does body contact contribute towards repairing relationships?: The association between flipper-rubbing and aggressive behavior in captive bottlenose dolphins

Bottlenose dolphins exhibit complex social affiliations that may be shaped by interactions among individuals. Affiliative body contact among dolphins may repair deteriorated relationships or reduce tension within the group following aggressive interactions. We investigated the time-series association between one type of contact behavior (flipper-rubbing) and aggression by continuous observation of three captive bottlenose dolphins. For all three dolphin pairs, the elapsed time to aggressive events was significantly greater following flipper-rubbing. In two dolphin pairs comprised of a young male and an adult female, one-zero score of inter-opponent flipper-rubbing was higher for 10 min following aggressive bouts (post-AG periods) than for the same length of control (Ctrl) periods. For all three focal pairs, one-zero score of third-party rubbing was higher for post-AG than Ctrl periods. Neither the direction of rubbing nor the identity of the partner that approached prior to rubbing showed any significant tendencies. Flipper-rubbing may contribute to restore friendly relationships between former opponents or reduce conflicts in at least juvenile-adult female associations. Our results also give preliminary suggestions of the functions of third-party flipper-rubbing among bottlenose dolphins, including tension easing by the third party, or displacement as a result of aggressive interactions.     

Contexts of rubbing behavior in Alouatta guariba clamitans: a scent‐marking role?

Rubbing behaviors are well known in several primate species and are usually seen as scent-marking behaviors, with several functions proposed but still widely debated. The genus Alouatta is highly sexually dimorphic and a suitable subject for the study of sexual and hierarchical divergences associated with rubbing behavior: males should mark more than females, and dominant individuals more than subordinate ones. Three wild groups of Southern brown howler monkeys, Alouatta guariba clamitans, were studied at Morro Geisler, Indaial, Brazil, from September 2004 to February 2005. One hundred and twenty-three rubbing episodes were registered; data on performers and associated contexts showed that anogenital, dorsum and hyoid regions were the most often rubbed. Adult males rubbed significantly above expected levels, whereas subordinated females and juveniles tended to rub below the expected levels. Females were the main performers of anogenital rubbing, often preceded by defecation. The predominance of rubbing in males probably serves an important function in intrasexual communication and social interactions. Intrasexual competition can also lead to a relationship between rubbing and social status in females. Hyoid and sternum rubbing by males are probably agonistic signals associated with extragroup conflict. The possible cleaning function of anogenital rubbing does not preclude a communicative function. Whether rubbing behavior in howlers is solely for the function of scent marking or can also be a visual signal (e.g. as a display or to color the substrate with pigment) requires further study.     

BEHAVIOR AND ECOLOGY OF THE ATLANTIC WHITE-SIDED DOLPHIN (LAGENORHYNCHUS ACUTUS) IN COASTAL NEW ENGLAND WATERS

Atlantic white-sided dolphins ( Lagenorhynchus acutus ) are among the most abundant, and least studied, cetaceans in coastal New England. Between April and October 1984 through 1997 we sighted 1,231 groups of Atlantic white-sided dolphins, primarily on Stellwagen Bank and Jeffreys Ledge (two shallow glacial deposits along the coasts of Massachusetts, New Hampshire, and Maine). Mean group size was 52 (≥90.9), and was significantly larger from August through October (71.9 ± 111.4) than April through June (35.0 ± 45.4). Calf sightings were uncommon until June and July, after which they were present in over 50% of groups. Combined with observations of apparent newborn calves, this confirms that early summer is an important calving period. The presence of calves did not, however, solely account for the increase in group size. Boat interaction (bow- and stern-wake riding) was the most commonly recorded behavior (47.4% of sightings), followed by traveling (31.4%), interactions with other cetacean species (27.6%), social interaction (15.5%), and feeding (9.5%). While feeding was uncommon, one observation of apparently coordinated "ball" feeding was seen with sand lance ( Ammodytes spp.) as the visible prey. Aerial behavior showed a positive correlation with group size, although it was often impossible to tell whether the same dolphins were leaping repeatedly. Eighty-eight dolphins were photo-identified using either unusual body pigment or a distinctive dorsal fin. While several individuals were reidentified between years and between areas, no reidentifications were made within a year in the same area. Unusually pigmented individuals were much more likely to be reidentified than those with distinctive dorsal fins, most likely due to higher visibility. We suggest that Atlantic white-sided dolphins are generally using the study area as transients in what appears to be a large home range.     

Observations of flipper rubbing in mother–calf pairs of captive bottlenose dolphins (Tursiops truncatus) suggest a body-surface care function

Journal of Ethology - Bottlenose dolphins perform flipper rubbing in which they rub their partners with their pectoral fins. Considered similar to social grooming in primates, which have social and...     

A comparison of pectoral fin contact behaviour for three distinct dolphin populations

Tactile exchanges involving the pectoral fin have been documented in a variety of dolphin species. Several functions (e.g., social, hygienic) have been offered as possible explanations for when and why dolphins exchange pectoral fin contacts. In this study, we compared pectoral fin contact between dolphin dyads from three distinct dolphin populations: two groups of wild dolphins; Atlantic spotted dolphins (Stenella frontalis) from The Bahamas and Indo-Pacific bottlenose dolphins (Tursiops aduncus) from around Mikura Island, Japan; and one group of captive bottlenose dolphins (Tursiops truncatus) residing at the Roatan Institute for Marine Sciences, Anthony's Key Resort. A number of similarities were observed between the captive and wild groups, including; rates of pectoral fin contact, which dolphin initiated contact, posture preference, and same-sex rubbing partner preference. Unlike their wild counterparts, however, dolphins in the captive study group engaged in petting and rubbing at equal rates, females were more likely to contact males, males assumed the various rubbing roles more frequently than females, and calves and juveniles were more likely to be involved in pectoral fin contact exchanges. These results suggest that some aspects of pectoral fin contact behaviour might be common to many dolphin species, whereas other aspects could be species specific, or could be the result of differing environmental and social conditions.     

Behavior of dolphins towards adults and children during swim-with-dolphin programs and towards children with disabilities during therapy sessions

Abstract

In recent years, dolphin-assisted therapy has become very popular and an increasing number of facilities worldwide offer therapy programs with dolphins. In contrast to other animal-assisted therapy programs, dolphins are not domestic animals; they are mostly caught in the wild and there are still no studies on their behavior during these therapies. However, there is speculation that the behavior of dolphins toward people with mental and physical disabilities may play an important role in the success of the therapy. We observed 83 sessions with five untrained dolphins (Tursiops truncatus) at Dolphins Plus, a fenced area with ocean water in the Florida Keys, USA. Our detailed observations of contact and distance behavior between dolphins and different groups of swimmers (adults, children, and children with mental and physical disabilities) show that, in general, dolphins prefer small humans to adults. One dolphin showed a clear preference toward children with mental and physical disabilities, and we conclude that she showed assisting behavior.     

A latitudinal gradient in intensity of applying ant-repellent substance to the nest petiole in paper wasps (Hymenoptera: Vespidae)

Summary Observations on pre-emergence, single-foundress colonies of Japanese paper wasps (Polistes) revealed that there was a latitudinal gradient in intensity of application of an ant-repellent substance (secreted by the metasomal sternum VI glands) to the nest petiole. Thus the lower the latitude, the more frequently a foundress rubbed ant repellent onto the nest petiole. Estimation of potential ant predation on wasp brood using bait traps showed that there was a positive correlation between the frequency of rubbing and potential predation pressure from ants which are guided predominantly by substrate cues for foraging. There was also a latitudinal gradient in the degree of temporal association of rubbing behavior with foraging: the lower the latitude, the more closely foundress departure from the nest was associated with rubbing. Enlargement of the nest petiole by applying oral secretion potentially obliterated previous coats of ant repellent; however, this behavior was not always followed by rubbing behavior. The ant-repellent chemical barrier around the nest petiole may have evolved in tropical regions of the world as a defense against ant predation on wasp brood. I argue that as ant predation pressure diminishes towards the cooler regions, so does selection maintaining the behavioral sequence where foundress departure from the nest is preceded by rubbing behavior.     

"PUSHY" BEHAVIOR IN A WILD DOLPHIN FEEDING PROGRAM AT TANGALOOMA, AUSTRALIA

A program where wild bottlenose dolphins (Tursiops truncatus) are fed by tourists in shallow water adjacent to a wharf has been established at Tangalooma, Queensland, Australia. Up to nine dolphins attend the nightly feedings, and between 60 and 80 resort guests are permitted to hand feed these dolphins each night. Since this program began in 1992, the dolphins have increased in confidence and have started, at times, to make forceful contact with guests who enter the water to feed them. This paper categorizes such behavior as "pushy" and reports on a study which quantifies the "pushiness" of the dolphins which feed at Tangalooma. The study examines ecological variables which may determine how pushy the dolphins are at different feeding sessions. The number of dolphins attending a particular feeding significantly increases the pushiness. In addition, the presence of adult males at a feeding is likely to increase pushing. Tidal state also influences how pushy the dolphins are. At low tide, when the dolphins'mobility is restricted by the water depth, they are less likely to be pushy. Given the problems experienced in a number of other situations where wild animals are fed by humans, it is important to monitor carefully the escalation of pushy behavior in this dolphin feeding program, as it may be a precursor to more aggressive actions on the part of the dolphins.     

Seasonal patterns of heat loss in wild bottlenose dolphins (<Emphasis Type="Italic">Tursiops truncatus</Emphasis>)

This study investigated patterns of heat loss in bottlenose dolphins (Tursiops truncatus) resident to Sarasota Bay, FL, USA, where water temperatures vary seasonally from 11 to 33°C. Simultaneous measurements of heat flux (HF) and skin surface temperature were collected at the body wall and appendages of dolphins during health-monitoring events in summer (June 2002–2004) and winter (February 2003–2005). Integument thickness was measured and whole body conductance (W/m² °C) was estimated using HF and colonic temperature measurements. Across seasons, HF values were similar at the appendages, but their distribution differed significantly at the flipper and fluke. In summer, these appendages displayed uniformly high values, while in winter they most frequently displayed very low HF values with a few high HF values. In winter, blubber thickness was significantly greater and estimated conductance significantly lower, than in summer. These results suggest that dolphins attempt to conserve heat in winter. In winter, though, HF values across the body wall were similar to (flank) or greater than (caudal keel) summer values. It is likely that higher winter HF values are due to the steep temperature gradient between the body core and colder winter water, which may limit the dolphin’s ability to decrease heat loss across the body wall.     

Rubbing behavior and morphology of van der Vecht's gland inBelonogaster petiolata (Hymenoptera: Vespidae)

Females of the social wasp, Belonogaster petiolata,rub the secretion of van der Vecht's gland, located on their terminal gastral sternite, onto the nest pedicel. In bioassays, the secretion was repellent to two species of ants, while shortchain acids were effective releasers of rubbing behavior. Rubbing was associated with pedicel enlargement and departure from the nest in preemergence colonies. Its frequency was high where wasps were often exposed to ants and low where ants were rare or absent. Rubbing also decreased significantly from the pre-to the postemergence stage of the colony cycle. In both stages, subordinate foundresses rubbed more often than queens or workers. These observations support the hypothesis that rubbing behavior and the secretion of van der Vecht's gland function in chemical defense of the nest against ant predation. The general morphology of the gland in B. petiolataresembles that of the four other independent-founding polistine wasp genera.     

Fine-scale analysis of synchronous breathing in wild Indo-Pacific bottlenose dolphins (Tursiops aduncus)

We quantitatively analysed synchronous breathing for dyads in Indo-Pacific bottlenose dolphins at Mikura Island, Tokyo, Japan. For most cases, we observed dyads swimming in the same direction (97%), in close proximity (i.e., less than 1.5 m) and with their body axes parallel as they breathed synchronously. Moreover, the pairs engaged in identical behaviour before and after the synchronous breathing episodes. These results suggest that the dolphins synchronize their movements, and that synchronous breathing is a component of “pair-swimming”, an affiliative social behaviour. Same sex pairs of the same age class frequently engaged in synchronous breathing for adults and subadults, as well as mother-calf and escort-calf pairs. The distance between individuals during synchronous breathing for mother-calf pairs was less than for other pairs. The distance observed between individuals for female pairs was less than for male pairs. The time differences between each exhale for each of the two dolphins involved in synchronous breathing episodes for female pairs were smaller than for male pairs, and time differences for adult pairs were smaller than subadult pairs. These results suggest that age and sex class influenced the characteristics of this behaviour.     

Diel and lunar variation in diving behavior of rough-toothed dolphins (Steno bredanensis) off Kauaʻi,Hawaiʻi

Observational studies describe rough-toothed dolphins (Steno bredanensis) actively foraging during the day on epipelagic species. Using data from depth-transmitting satellite tags deployed on nine individuals off Kauaʻi, we investigated diving behavior and the effects of lunar phase and solar light levels on vertical movements. Overall, tagged rough-toothed dolphins primarily used near-surface waters, spending between 83.6% and 93.7% of their time in the top 30 m of the water column. When diving, grand mean, median, and maximum dive depths were 76.9 m, 67.5 m, and 399.5 m, although individuals were in water with depths from approximately 700–1,450 m. Dive rates varied by time of day, being lowest during the day and at dawn and highest at dusk and night. Dives were deepest (M = 133.7 m, SD = 52.6 m, median = 106.5 m) and longest (M = 4.0 min, SD = 0.4 min, median = 4.0 min) at dusk, suggesting dolphins were taking advantage of prey rising to the surface in response to reduced light levels. Lunar phase indirectly affected diving, with deeper and longer dives occurring with increasing illumination. The variations in dive behavior across solar and lunar cycles indicate diving patterns shift based on the distribution of prey.     

Sociability of female bottlenose dolphins (Tursiops spp.) and chimpanzees (Pan troglodytes): Understanding evolutionary pathways toward social convergence

On the northern tip of New Zealand's South Island, bottlenose dolphins (Tursiops truncatus) occasionally pass through Admiralty Bay in large, fast-traveling groups of 100 or so individuals. Watching such a group race and splash through the water is reminiscent of a stampeding herd of ungulates, cetaceans' closest terrestrial ancestors. At other times, smaller social groups of bottlenose dolphins appear in the bay and provide a glimpse of the behavioral complexity that dolphins share with their distant relatives, the primates (Fig. 1). Despite being evolutionarily separated for 95 million years and evolving in vastly different environments, cetaceans and primates share striking similarities in behavior, socioecological problem-solving, life-history patterns, and cognitive capacity. By comparing attributes shared by primates and cetaceans, distraction from phylogenetic "noise" is minimized and our understanding of evolutionary pathways is enhanced. In particular, cetaceans provide a powerful outgroup for studying the evolution of primate social organization.     

SURFACE BEHAVIOR OF BOTTLENOSE DOLPHINS IS RELATED TO SPATIAL ARRANGEMENT OF PREY

We tested the hypothesis that spatial arrangement and movements of bottlenose dolphins ( Tursiops truncatus ) are related to the spatial arrangement of their prey. From 65 groups we: (1) classified feeding behavior of dolphins according to spatial arrangement and movements of individuals; (2) assessed spatial arrangement and location of prey from direct observations, numbers of associated seabirds, and echosounder recordings; and (3) related feeding behavior of dolphins to spatial arrangement and location of prey. Four feeding categories were defined from cluster and principal component analyses: (1) moving rapidly with no diving, (2) milling with no diving, (3) diving in several locations, and (4) diving in one location. These feeding categories were related to spatial arrangement and location of prey.     

QUANTITATIVE BEHAVIORAL STUDY OF BOTTLENOSE DOLPHINS IN SWIM-WITH-DOLPHIN PROGRAMS IN THE UNITED STATES

The behavior of dolphins in four Swim-With-Dolphin programs was compared by type of Swim encounter, defined by the presence ("Controlled") or absence ("Not-Controlled") of explicit trainer regulation of interactions between dolphins and human swimmers. Dolphin-swimmer interactions involving aggressive, submissive, or sexual behavior were designated as "high-risk" in the Swim context; sexual behavior was included as high-risk based on analyses that demonstrated co-occurrence of sexual and agonistic behaviors. High-risk activity comprised a substantial proportion of dolphin-swimmer social activity during Not-Controlled Swims. In contrast, high-risk activity rarely occurred during Controlled Swims, even though agonistic and sexual behaviors were normal components of the same dolphins' free-time social repertoire. These results indicated that direct trainer control of dolphin-swimmer interactions virtually eliminated high-risk activity from the Swim context, and thereby diminished the potential for dolphin distress, swimmer injury, and rejection of dolphins from Swim programs due to swimmer injury. This study illustrates effective use of quantitative behavioral sampling techniques for evaluation of captive management concerns and promotes broader use of these techniques for a better understanding of cetacean behavior.     

A comparison of pectoral fin contact between two different wild dolphin populations

Contact behaviour involving the pectoral fin has been documented in a number of dolphin species, and various explanations about its function have been offered. Pectoral fin contact can take a variety of forms, and involves a number of body parts and movements, likely differing depending upon social or ecological context. For this study, we compare the pectoral fin contact behaviour of two species of wild dolphins: Indo-Pacific bottlenose dolphins (Tursiops aduncus) from around Mikura Island, Japan, and Atlantic spotted dolphins (Stenella frontalis) from The Bahamas. The two study populations exhibit surprising similarity in the ways in which pectoral fin contacts are used, despite differences in species and environmental conditions at the two sites. Differences in contact rates for calves between the two sites suggest that calf-focused aggression from adult dolphins is more prevalent at Mikura than in The Bahamas. Our results suggest that pectoral fin contact behaviour seems to be driven primarily by social pressures, and may be similar in function to allogrooming described in primates.     

Flipper Bands Modify the Short-Term Diving Behavior of Little Penguins

ABSTRACT Flipper banding has long been the primary method to identify individual penguins, despite studies indicating that it may be detrimental to breeding success and survival. Our objectives were to measure the effects that flipper bands may have on diving performance of little penguins to determine whether the bands may be detrimental. We studied short- and long-term direct effects of flipper banding on diving behavior of free-ranging little penguins (Eudyptula minor) by comparing diving behavior before and after banding and by comparing diving performance of unbanded birds to those that had carried flipper bands for several years, respectively. Recently banded birds displayed increases in multiple variables following banding. Long-term banded penguins did not exhibit differences to their unbanded counterparts in most variables examined. Our findings are useful to those considering or reviewing the use of bands in penguin study and management.     

Social facilitation of fur rubbing behavior in white-faced capuchins

In their natural environment, capuchins select certain plants, containing secondary compounds with bactericide, insecticide or fungicide properties, to rub their pelage energetically (i.e. fur rubbing). Fur rubbing can be performed in solitary, or collectively in subgroups of variable size and composition, and most of the time fur rubbing happens in synchrony with other group members. The aim of this study is to understand the underlying mechanisms of this phenomenon, and, more particularly, to determine the processes involved in its synchronization. For this purpose, we designed a set of experiments where white-faced capuchins (Cebus capucinus) were presented with onions (Allium cepa) that they use to fur rub. We conducted a detailed kinetic study of fur rubbing behavior to determine if its synchronization is the consequence of simultaneous responses of different individuals to the same stimulus or if, on the contrary, there is a real collective phenomenon where individuals respond to conspecific behavior. Our results reveal that fur rubbing is a collective behavior with a mimetic underlying mechanism. If fur rubbing with onions (a plant with antifungal and repellent properties) allows capuchins to treat their fur against parasites or pathogens, its synchronization would optimize the treatment by acting as a group barrier to ectoparasite propagation.     

SEXUAL DIMORPHISM IN BOTTLENOSE DOLPHINS FROM THE EAST COAST OF FLORIDA

Skulls of 69 bottlenose dolphins (genus Tursiops ) from the Indian/Banana River on the east coast of Florida were examined for evidence of sexual dimorphism. The only sexual dimorphism shown by t -tests on 28 morphological and four meristic skull characters was that males have, on average, more teeth than females in all four arcades. Results of covariance analysis, employed to account for variation in size, indicate minor dimorphism in parietal width of the skull. Twenty body measurements of 29 Tursiops originating in the same area were also analysed for differences between males and females. Statistical results indicate the possibility of sexual dimorphism in the length from the snout to the umbilicus and in flipper width. No evidence was found for differences in overall skull or body length between the sexes.