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Death-related behaviors have been often observed in cetaceans, frequently with a female caregiver (presumed mother) supporting a dead calf by carrying, lifting, or sinking it, generally accompanied by other escorting individuals. However, how cetaceans experience death and if their behavior could be compared to human grieving is still largely unknown, as well as their acoustic production in such contexts. This report describes two cases of an adult common bottlenose dolphin (Tursiops truncatus) supporting a dead newborn and associated acoustic behavior in the Tiber River estuary area (Rome, Mediterranean Sea, Italy). In both cases, a main supporter (putative mother) was observed interacting with the carcass of a newborn by lifting (case 1) and sinking it (case 2), always escorted by another adult individual. Several vocalizations were recorded, including a signature whistle (a tonal sound with a repeated, stereotyped, individual-specific frequency modulation pattern) in the first case, likely emitted by the putative mother to seek aid from other individuals. This result appears reasonable since bottlenose dolphins are a highly vocal species, that communicates mainly through acoustic signals. These observations confirm the occurrence of death-related supportive behavior in bottlenose dolphins and report a preliminary description of associated vocalizations, providing additional information on this largely unknown topic.
Epimeletic behavior toward dead calves has been frequently reported in cetaceans, mostly with females (presumed mothers) showing supportive behaviors such as carrying, lifting, or sinking, often assisted by “escort” individuals. However, information on acoustic production in such contexts is scarce. This report describes two observations of epimeletic behavior toward dead newborns in bottlenose dolphins and associated acoustic production. Data were collected at the Tiber River estuary (Rome, Mediterranean Sea, Italy) with one hydrophone for passive acoustic monitoring and two digital cameras. In both cases, an individual (presumed mother) acting as the main supporter and directly interacting with the carcass by lifting it (case 1) or sinking it (case 2) was observed. Another adult individual (escort) was present in both encounters showing standing-by and excitement behaviors (case 1) and supportive behavior (case 2). In both encounters, whistles, pulsed sounds, and bray-call elements were recorded. The consistent vocal activity observed likely conveyed context-specific information. A signature whistle in the first encounter was also recorded, likely emitted by the putative mother as a distress call. This report confirms the occurrence of epimeletic behavior in bottlenose dolphins and reports a preliminary description of the acoustic production when a dead calf is involved, providing additional information on this largely unknown topic.
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Behavioral responses to dead conspecifics have long been thought to be a unique feature of the human species. However, observations of individuals showing supportive behavior toward dead or dying conspecific in the wild have been reported for several terrestrial and marine mammal species, including chimpanzees [1], giraffes [2], elephants [3], and cetaceans [4]. Death-related behaviors can be considered a subtype of epimeletic behavior, which occurs when a healthy individual (supporter) gives attention to or takes care of another distressed, injured, dying or, dead individual (receiver) [5]. Caldwell and Caldwell [6] distinguished two different categories of epimeletic behavior: nurturant (supportive behavior directed toward younger individuals) and succorant (support aimed at adults). In cetaceans, nurturant behavior is the most reported, frequently directed toward dead newborns [7]. When a calf is involved, the caregiver appears to be a female (presumably the mother) in most of the cases in which its sex is known [5]. The supporter may often be accompanied by other individuals, called “escorts”, who either directly assist the caregiver interacting with the receiver or remain close and swim by the pair [5, 8]. Post-mortem supportive behavior in cetaceans generally includes stereotyped behaviors that can be divided into three main groups: (1) standing-by, with individuals remaining close to the carcass without directly interacting with it, (2) excitement, with participants showing behaviors typical of arousal states such as erratic swimming, and (3) supportive behavior, with individual interacting directly with the carcass, typically lifting, sinking or carrying it with the melon, the rostrum, or the edge of the dorsal fin [9, 10].
Among cetaceans, the common bottlenose dolphin (Tursiops truncatus) is one of the species most frequently reported to exhibit epimeletic behavior [4, 5, 11]. Consistently with other cetacean species, death-related behaviors are mainly directed from an adult female toward a younger individual (presumably its offspring) and they include carrying, mouthing, lifting, and diving [4]. Bottlenose dolphins are a highly vocal species with a complex vocal repertoire [12]. They show strong vocal plasticity [13] and are known to adapt the emission rates and the acoustic features of their vocalizations depending on the contexts and emotional state [14, 15, 16]. However, despite the consistent number of reports on epimeletic behavior for this species, still little is known about their acoustic production in such contexts, and little evidence has been reported. The only available description of acoustic production during epimeletic behavior in bottlenose dolphins is reported by Kuzcaj et al. [17] that characterized the whistles emitted during a case of succorant behavior directed toward a distressed conspecific. They found out that signature whistles were emitted as distress calls, with higher emission rates and higher intensity to seek aid from other individuals [17]. Few other reports provide a general description of the acoustic production in epimeletic-related contexts. The study from Perrtree et al. [18] reported higher emission rates of the signature whistle emitted by a mother during an infanticide attempt in bottlenose dolphins, possibly expressing context-related information. The one from Cheng et al. [9] characterized the whistles produced by an adult Indo-pacific humpback dolphin (Sousa chinensis) during nurturant behavior directed toward a dead calf. They reported a longer duration and higher number of inflection points in whistles emitted during nurturant behavior than in whistles produced in other contexts and hypothesized that these variations may convey information on the emotional state of the emitter [9]. Finally, [19] reported consistent whistles emission during a case of succorant behavior in bottlenose dolphins, with possible distress function.
Here, two episodes of nurturant behavior toward dead calves observed in 2021 in the population of bottlenose dolphins inhabiting the Tiber River estuary area (Rome, Italy, Mediterranean Sea), and associated acoustic production, are reported. This is the first detailed evidence of epimeletic behavior involving a dead calf in this population, although another case was observed by a sea-user in 2015 [20]. The final aim of this report is to provide additional information on death-related behaviors in such a species, moving toward a deeper comprehension of this still largely unknown behavioral context.
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And is located in the central Mediterranean Sea (Tyrrhenian Sea, Rome, Italy) (Figure 1). It includes the Tiber River estuary, which flows into the sea through the two mouths of Fiumara Grande (natural mouth) and Fiumicino (artificial mouth).
The area is rich in habitats and environmental conditions, and is characterized by a flourishing marine community that developed partly because of the large amount of organic material transported by the river [21]. However, crossing the city of Rome, the Tiber also contributes to enhancing pollution levels, bringing to the sea big quantities of pollutants, heavy metal and waste [21]. The study site also includes the “Secche di Tor Paterno” marine protected area and two single point moorings (SPMs, called R1 and R2) for the reception of crude oil and the supply of petroleum products (structures known to attract bottlenose dolphins [22]).
The area is well recognized for the presence of bottlenose dolphins [23], with the regular observation of mother-calf pairs and foraging behavior, thus appearing a suitable calving and feeding ground for the species [20, 21]. To date, 347 unique individuals have been photo identified in the area, showing different patterns of site fidelity [21].
Underwater World (2018 2021) By Mdm
And is located in the central Mediterranean Sea (Tyrrhenian Sea, Rome, Italy) (Figure 1). It includes the Tiber River estuary, which flows into the sea through the two mouths of Fiumara Grande (natural mouth) and Fiumicino (artificial mouth).
The area is rich in habitats and environmental conditions, and is characterized by a flourishing marine community that developed partly because of the large amount of organic material transported by the river [21]. However, crossing the city of Rome, the Tiber also contributes to enhancing pollution levels, bringing to the sea big quantities of pollutants, heavy metal and waste [21]. The study site also includes the “Secche di Tor Paterno” marine protected area and two single point moorings (SPMs, called R1 and R2) for the reception of crude oil and the supply of petroleum products (structures known to attract bottlenose dolphins [22]).
The area is well recognized for the presence of bottlenose dolphins [23], with the regular observation of mother-calf pairs and foraging behavior, thus appearing a suitable calving and feeding ground for the species [20, 21]. To date, 347 unique individuals have been photo identified in the area, showing different patterns of site fidelity [21].
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