Dive bout organization in the Chinstrap penguin at seal island, antarctica

Diving behavior of 2 breeding Chinstrap penguins (Pygoscelis antarctica) was studied focusing first and primarily on dive bouts rather than dives themselves. Analysis of dive bout organization revealed (1) though there are differences between solitary dives and dive bouts in dive duration and dive depth, the first dives of dive bouts do not differ from solitary dives in the dive parameters, (2) mean dive duration during bout correlates positively to both mean dive depth during bout and mean surface interval during bout, while number of dives during bout negatively correlates to both cost (consumed energy) and duration of a dive cycle during bout. These findings suggest the following possibilities on foraging behavior of penguins: (1) their decision to repeat diving depends on the result of the first dive at a site, and the first dives of bouts would tend to be searching or evaluating dives though they would be also successful foraging dives, (2) they repeat diving at a foraging patch until foraging efficiency decrease to a threshold of diminishing returns.     

Effect of age of calf on suckling behaviour and other behavioural activities of Zebu and crossbred calves during restricted suckling periods

The objective of this study was to investigate the effect of age of calf on the behaviour of Zebu and crossbred calves during restricted suckling (RS) periods. The behaviours of 20 Zebu and 16 crossbred calves were recorded during two 30-min sessions each day after milking when the calves and their dams were brought together in a group for suckling. This was made for a time period of 2 weeks/month for 6 months postpartum.The total suckling duration was significantly longer in Zebu calves (11.8+/-0.19 min) compared to the crossbred calves (9.4+/-0.19 min), but decreased significantly in both breeds with increasing age from 1 to 6 months. The number of suckling bouts decreased from a mean of 3.8 at 1 month of age to 1.1 at 6 month (P<0.05). The duration of each suckling bout decreased significantly from a mean of 3.5+/-0.15 min at 1 month of age to 1.6+/-0.01 min at 6 months (<0.05). The frequency of crossbred calves cross-suckling (3.7%) was significantly higher than that of the Zebu calves (1.9%; P<0.05). The frequency of calves cross-suckling decreased significantly from 4.2% at 1 month of age to 2.3% at 6 months. The duration and number of bouts of cross-suckling was significantly higher in the crossbred calves (duration 0.9+/-0.06 min; bouts 3, 7) than in the Zebu calves (duration 0.5+/-0.06 min; bouts 2, 7) and decreased with increasing age of calf. The duration and number of bouts of inter-sucking was significantly higher in the crossbred calves (duration 0.6+/-0.07 min; bouts 1, 6) than in the Zebu calves (duration 0.1+/-0.04 min; bouts 0, 5) and decreased with increasing age of calf. Exploration increased in duration as the calves increased in age from 1 to 6 months (P<0.05). The duration of play increased significantly with the increase in age of calf from 1 to 6 months, and occurred mainly after nursing.     

Diving patterns and performance in the Antarctic blue-eyed shag Phalacrocorax atriceps

The pattern and characteristics of diving of two male blue-eyed shags Phalacrocorax atriceps were studied, using continuous-recording time-depth recorders, for a total of 15 consecutive days during which the depth, duration, bottom time, ascent and descent rates and surface intervals of 674 dives were recorded. Deep dives (> 35 m, averages80–90 m, max. 116 m) were twice as common (64% versus 34%) as shallow dives (< 21 m and 90% < 10 m). Deep dives were long (averages 2.7-4.1 min, max. 5.2 min) with half the time spent near maximum depth and fast travel speeds (averages 1.0-2.4 m s⁻¹). Shallow dives were short (average 0.5 min, max. 1.3 min), without bottom time and with slow travel speeds (0.1–0.6 m s⁻¹). The time spent at depth and the diet (mainly benthic fish and octopus) is consistent with benthic foraging; the function of shallow dives is uncertain. Male shags forage mainly in the afternoon in3–5 distinct bouts of diving. Within bouts (and shorter homogeneous sequences of diving) surface intervals are consistently2–3 times the preceding dive duration; in other shags the reverse is the case. Blue-eyed shag diving depth, duration and pattern is extreme amongst shags; and the relationship between dives and surface intervals suggests that they may regularly exceed their aerobic dive limit.     

Norwegian deep diving trials

In 1983 NUTEC, together with two diving companies, completed two dives with 12 divers (6 in each dive) to pressures equivalent to 350 m s.w., one dive lasted for 17 d, and the other, 24 d. The purpose of the dives was to demonstrate that the diving companies were prepared for diving to 300 m depth in the North Sea. No major medical or physiological problems arose during the dives, although all divers had minor symptoms of high pressure nervous syndrome during compressions. During decompression three decompression sickness incidents occurred, which involved pain only, and all were successfully treated. All divers went through comprehensive medical physiological examinations before and after the dives. No significant changes from values measured before diving have been found in the six divers who have so far been examined after diving, except that five of them were considerably more sensitive to CO2 after the dive than before. Several problems arose in connection with the divers' breathing equipment, thermal protection and communication, which need to be improved.     

Diving and haulout behavior of crabeater seals in the Weddell Sea,Antarctica, during March 1986

Summary Time-depth recorders were used to study the diving and haulout behavior of six crabeater seals in the marginal. ice edge zone of the Weddell Sea during March 1986. Haulout patterns revealed the seals' clear preference for diving during darkness and hauling out onto sea ice during daylight. Seals did not necessarily haul out every day; individual seals hauled out on 80–100% of days during the study period. Four general dive types were identified: 1) traveling dives, 2) foraging dives, 3) crepuscular foraging dives, and 4) exploratory dives. Nearly continual diving occurred for extended periods (about 16 h) nightly, with one individual diving up to 44 h without interruption. Foraging dives occurring during crepuscular periods were deeper than those made during the darkest hours. The authors suggest that the distinct diel pattern of dive timing and depth may be related to possible predator avoidance behavior by the seals' principal prey, Antarctic Krill.     

SPERM WHALE DIVES TRACKED BY RADIO TAG TELEMETRY

Dives of a 12-m sperm whale ( Physeter catodon Linnaeus, 1758) were tracked in the southeast Caribbean by long range, 30 MHz radio tag with dive-profile telemetry over 4.6 d, 26 April-1 May 1995. Over the 295-km track, average speed was 0.7 m/sec (2.6 km/h). Of 158 dives (defined as submergences longer than 3 min), 65 were shallow (<200 m). The 93 deep dives averaged 990 m (range 420–1,330 m) in depth, and 44.4 min in duration (range 18.2–65.3 min). Water depth was at least 200 m deeper than the whale dive depth. The whale was engaged in activities at or near the surface, shallow dives, and deep dives during 22.6%, 23.4%, and 54% of the time, respectively. Depth and duration of dives were correlated, but there was little relationship between the length or depth of dives with the duration of surfacings either before or after dives. Deep dives occurred day and night. In 44.4% of the deep dives, the vertical movement of descents and ascents was interrupted at intermediate depths, lengthening these dives by an average of 10.8 min. During dives without stops at intermediate depths, descents averaged 11 min at 1.52 m/sec, and ascents averaged 11.8 min at 1.4 m/sec.     

Flexible foraging techniques in breeding Cormorants Phalacrocorax carbo and Shags Phalacrocorax aristotelis: benthic or pelagic feeding?

A total of 8772 dive durations were recorded during 117 diving bouts in five Cormorants Phalacrocorax carbo and five Shags Phalacrocorax aristotelis breeding at the Chausey Islands, France. Diet of the birds was assessed by analysis of 526 pellets containing 13,016 otoliths. Radio-tracking data indicated that Cormorants fed exclusively on pelagic fish during social fishing (5% of the trips) and executed 11% pelagic and 60% benthic dives during the remaining 95% of the trips. In Shags, 44% of all trips were pelagic, and the remaining 56% included 9% pelagic and 67% benthic dives. The proportions of benthic to pelagic dives varied widely between dive sequences of single birds and between individuals and sexes in both species. The prey spectrum of the Cormorants contained both pelagic (29%) and benthic fish (67%) and confirmed considerable flexibility in foraging. In Shags, birds may adjust their diving patterns to accommodate the behaviour of their main prey, sandeels Ammodytidae (87% of all prey). We propose that the wetability of plumage may explain this flexibility.     

Cardiac responses to first ever submergence in double-crested cormorant chicks (Phalacrocorax auritus)

Heart rates were recorded from double-crested cormorant chicks during their first ever and subsequent voluntary head submergences and dives, as well as during longer dives made after the chicks were accustomed to diving. Despite variation between chicks, the cardiac response to first ever and subsequent voluntary submergence (head submergences and dives) was similar to the response observed in adult cormorants. Upon submersion the heart rate fell rapidly when pre-submersion heart rate was high (325-350 beats min-1). The heart rate established within the first second of voluntary submergence was between 230 and 285 beats min-1, well above resting heart rate (143 beats min-1). The same initial cardiac response occurred during longer dives performed after the chicks were accustomed to diving. In these dives the heart rate remained at the level established on submersion, unlike the response observed in shallow diving adult cormorants in which the heart rate declined throughout the dive. The heart rate was also monitored in a separate group of chicks in which the first exposure to water was during whole body forced submergence. Again, the observed response was similar to the adult response, although the cardiac response of chicks to forced submergence was more extreme than to voluntary submergence. Our results do not support the hypothesis that learning (by conditioning or habituation) is involved in the cardiac adjustments to voluntary submergence. It is suggested that the initial cardiac adjustments are reflex in nature and this reflex is fully developed by the first submergence event. Although the nature of this reflex pathway is obscure, cessation of breathing before submersion and the close linkage between breathing and heart rate might provide a plausible mechanism.     

Diving pattern and performance in relation to foraging ecology in the gentoo penguin, Pygoscelis papua

We present data on diving pattern and performance (dive depth, duration, frequency and organization during the foraging trip) in gentoo penguins Pygoscelis papua , obtained using time-depth recorders ( n = 9 birds, 99 foraging trips). These data are used to estimate various parameters of foraging activity, e.g. foraging range, prey capture rates, and are compared in relation to breeding chronology. Foraging trip duration was 6 h and 10 h, and trip frequency 1.0/day and 0.96/day, during the brooding and creche periods, respectively. Birds spent on average 52%of each foraging trip diving. Dive depth and duration were highly bimodal: shallow dives (< 21 m) averaged 4 m and 0.23 min, and deep dives (> 30 m) 80 m and 2.5 min, respectively. Birds spent on average 71%and 25%of total diving time in deep and shallow dives, respectively. For deep dives, dive duration exceeded the subsequent surface interval, but shallow dives were followed by surface intervals 2–3 times dive duration. We suggest that most shallow dives are searching/exploratory dives and most deep dives are feeding dives. Deep dives showed clear diel patterns averaging 40 m at dawn and dusk and 80–90 m at midday. Estimated foraging ranges were 2.3 km and 4.1 km during the brood and creche period, respectively. Foraging trip duration increased by 4 h between the brood and creche periods but total time spent in deep dives (i.e. time spent feeding) was the same (3 h). Of 99 foraging trips, 56%consisted of only one dive bout and 44%of 2–4 bouts delimited by extended surface intervals > 10 min. We suggest that this pattern of diving activity reflects variation in spatial distribution of prey rather than the effect of physiological constraints on diving ability.     

Behaviour of Zebu and crossbed cows in restricted suckling groups

The aim of this study was to investigate whether Zebu (Bos indicus) and crossbred (Bos indicusxBos taurus) cows differ in maternal related behaviour during restricted group suckling. The behaviours of 20 Zebu and 16 crossbred cows were recorded during two 30min sessions each day after milking when their calves were present for suckling, for a time period of 2 weeks per month for 6 months postpartum. The main activity in both breeds during the 0.5h observation sessions was suckling. However, the two breeds differed significantly in several behavioural aspects. The Zebu cows had longer total duration of suckling than the crossbred cows, per 30min session, 11.8 and 9.4min, respectively. The separate suckling bouts were longer in the Zebu cows (2.8min in Zebu and 2.3min in crossbred) and this breed also had more suckling bouts than the crossbred per session, 3.8 and 3.2, respectively. Zebu cows stayed in close contact with their calves for longer time and directed more agonistic actions against alien calves and cows than the crossbred cows.     

Deep-diving foraging behaviour of sperm whales (Physeter macrocephalus)

1. Digital tags were used to describe diving and vocal behaviour of sperm whales during 198 complete and partial foraging dives made by 37 individual sperm whales in the Atlantic Ocean, the Gulf of Mexico and the Ligurian Sea. 2. The maximum depth of dive averaged by individual differed across the three regions and was 985 m (SD = 124.3), 644 m (123.4) and 827 m (60.3), respectively. An average dive cycle consisted of a 45 min (6.3) dive with a 9 min (3.0) surface interval, with no significant differences among regions. On average, whales spent greater than 72% of their time in foraging dive cycles. 3. Whales produced regular clicks for 81% (4.1) of a dive and 64% (14.6) of the descent phase. The occurrence of buzz vocalizations (also called 'creaks') as an indicator of the foraging phase of a dive showed no difference in mean prey capture attempts per dive between regions [18 buzzes/dive (7.6)]. Sperm whales descended a mean of 392 m (144) from the start of regular clicking to the first buzz, which supports the hypothesis that regular clicks function as a long-range biosonar. 4. There were no significant differences in the duration of the foraging phase [28 min (6.0)] or percentage of the dive duration in the foraging phase [62% (7.3)] between the three regions, with an overall average proportion of time spent actively encountering prey during dive cycles of 0.53 (0.05). Whales maintained their time in the foraging phase by decreasing transit time for deeper foraging dives. 5. Similarity in foraging behaviour in the three regions and high diving efficiencies suggest that the success of sperm whales as mesopelagic predators is due in part to long-range echolocation of deep prey patches, efficient locomotion and a large aerobic capacity during diving.     

Sperm whale dive behavior characteristics derived from intermediate‐duration archival tag data

Here, we describe the diving behavior of sperm whales (Physeter macrocephalus) using the Advanced Dive Behavior (ADB) tag, which records depth data at 1‐Hz resolution and GPS‐quality locations for over 1 month, before releasing from the whale for recovery. A total of 27 ADB tags were deployed on sperm whales in the central Gulf of California, Mexico, during spring 2007 and 2008, of which 10 were recovered for data download. Tracking durations of all tags ranged from 0 to 34.5 days (median = 2.3 days), and 0.6 to 26.6 days (median = 5.0 days) for recovered tags. Recovered tags recorded a median of 50.8 GPS‐quality locations and 42.6 dives per day. Dive summary metrics were generated for archived dives and were subsequently classified into six categories using hierarchical cluster analysis. A mean of 77% of archived dives per individual were one of four dive categories with median Maximum Dive Depth >290 m (V‐shaped, Mid‐water, Benthic, or Variable), likely associated with foraging. Median Maximum Dive Depth was <30 m for the other two categories (Short‐ and Long‐duration shallow dives), likely representing socializing or resting behavior. Most tagged whales remained near the tagging area during the tracking period, but one moved north of Isla Tiburón, where it appeared to regularly dive to, and travel along the seafloor. Three whales were tagged on the same day in 2007 and subsequently traveled in close proximity (<1 km) for 2 days. During this period, the depth and timing of their dives were not coordinated, suggesting they were foraging on a vertically heterogeneous prey field. The multiweek dive records produced by ADB tags enabled us to generate a robust characterization of the diving behavior, activity budget, and individual variation for an important predator of the mesopelagos over temporal and spatial scales not previously possible.     

Comparing multiple sampling platforms for measuring the behavior of humpback whales (Megaptera novaeangliae)

Multiple platform approaches for measuring the behavior of marine mammals may strengthen data quality and quantity. However, if this approach is chosen, reconciliation of behavioral measures between each platform is required. This study compared typical measures of humpback whale (Megaptera novaeangliae) behavior collected from three different sampling platforms; land‐based, boat‐based, and digital tags (Dtags), to explore similarities and/or discrepancies in data. Visual observations from land‐based platforms significantly underestimated group blow rate when compared to boat‐based platforms, whereas broad‐scale spatial movements (speed and course traveled) were measured similarly by these two platforms. Dtags were used to define dive behavior based on a time and depth criteria, where bouts of short shallow respiration dives (≤80 s and ≤10 m) were separated from longer, deeper dives (>80 s and >10 m). At a group level, land and boat platforms agreed on the number of long dives but land platforms missed bouts of short dives. At an individual level, the number of short and long dives observed by boat‐based platforms agreed with Dtag recordings. This study highlights the importance of data exploration prior to analyses in multiple platform studies to identify potential discrepancies and appropriately account for any biases.     

Diving in shallow water: the foraging ecology of darters (Aves: Anhingidae)

Diving birds have to overcome buoyancy, especially when diving in shallow water. Darters and anhingas (Anhingidae) are specialist shallow-water divers, with adaptations for reducing their buoyancy. Compared to closely-related cormorants (Phalacrocoracidae), darters have fully wettable plumage, smaller air sacs and denser bones. A previous study of darter diving behaviour reported no relationship between dive duration and water depth, contrary to optimal dive models. In this study I provide more extensive observations of African darters Anhinga melanogaster rufa diving in water<5 m deep at two sites. Dive duration increases with water depth at both sites, but the relationship is weak. Dives were longer than dives by cormorants in water of similar depth (max 108 s in water 2.5 m deep), with dives of up to 68 s observed in water<0.5 m deep. Initial dives in a bout were shorter than expected, possibly because their plumage was not fully saturated. Dive efficiency (dive:rest ratio) was 5–6, greater than cormorants (2.7±0.4 for 18 species) and other families of diving birds (average 0.2–4.3). Post-dive recovery periods increased with dive duration, but only slowly, resulting in a strong increase in efficiency with dive duration. All dives are likely to fall within the theoretical anaerobic dive limit. Foraging bouts were short (17.8±4.3 min) compared to cormorants, with birds spending 80±5% of time underwater. Darters take advantage of their low buoyancy to forage efficiently in shallow water, and their slow, stealthy dives are qualitatively different from those of other diving birds. However, they are forced to limit the duration of foraging bouts by increased thermoregulatory costs associated with wettable plumage.     

Diving pattern and performance in the macaroni penguin Eudptes chrysolophus

The pattern and characteristics of diving in two female macaroni penguins Eudyptes chrysolophus was studied, during the brooding period, using continuous-recording time-depth recorders, for a total of I8 days (15 consecutive days) during which the depth, duration and timing of 4876 dives were recorded. Diving in the first 11 days was exclusively diurnal, averaging 244 dives on trips lasting 12 hours. Near the end of the brooding period trips were longer and included diving at night. About half of all trips (except those involving continuous night-time diving) was spent in diving and dive rate averaged 14–25 dives per hour (42 per hour at night). The duration of day time dives varied between trips, and averaged 1.4–1.7 min, with a subsequent surface interval of 0.5–0.9 min. Dive duration was significantly directly related to depth, the latter accounting for 53% of the variation. The average depths of daytime dives were 20–35 m (maximum depth 11 5 m). Dives at night were shorter (average duration 0.9 min) and much shallower (maximum 11 m); depth accounted for only 6% of the variation in duration. Estimates of potential prey capture rates (3–5 krill per dive; one krill every 17–20 s) are made. Daily weight changes in chicks were directly related to number of dives, but not to foraging trip duration nor time spent diving. Of the other species at the same site which live by diving to catch krill, gentoo penguins forage exclusively diurnally, making longer. deeper dives; Antarctic fur seals, which dive to similar depths as macaroni penguins, do so mainly at night.     

Sperm whale behaviour indicates the use of echolocation click buzzes "creaks" in prey capture

During foraging dives, sperm whales (Physeter macrocephalus) produce long series of regular clicks at 0.5-2 s intervals interspersed with rapid-click buzzes called "creaks". Sound, depth and orientation recording Dtags were attached to 23 whales in the Ligurian Sea and Gulf of Mexico to test whether the behaviour of diving sperm whales supports the hypothesis that creaks are produced during prey capture. Sperm whales spent most of their bottom time within one or two depth bands, apparently feeding in vertically stratified prey layers. Creak rates were highest during the bottom phase: 99.8% of creaks were produced in the deepest 50% of dives, 57% in the deepest 15% of dives. Whales swam actively during the bottom phase, producing a mean of 12.5 depth inflections per dive. A mean of 32% of creaks produced during the bottom phase occurred within 10 s of an inflection (13x more than chance). Sperm whales actively altered their body orientation throughout the bottom phase with significantly increased rates of change during creaks, reflecting increased manoeuvring. Sperm whales increased their bottom foraging time when creak rates were higher. These results all strongly support the hypothesis that creaks are an echolocation signal adapted for foraging, analogous to terminal buzzes in taxonomically diverse echolocating species.     

KILLER WHALE PREDATION ON SPERM WHALES: OBSERVATIONS AND IMPLICATIONS

In October 1997 we observed a herd of approximately 35 killer whales ( Orcinus orca ) attack a pod of nine sperm whales ( Physeter macrocephalus ) 130 km off the coast of central California. During the four hours we watched, adult female killer whales, including some with calves, attacked in waves of four to five animals in what was apparently a "wound and withdraw" strategy. Adult male killer whales stood by until the very end when one charged in and quickly killed a seriously wounded sperm whale that had been separated from the group. The sperm whales appeared largely helpless: their main defensive behavior was the formation of a rosette ("marguerite"-heads together, tails out). When the killer whales were successful in pulling an individual out of the rosette, one or two sperm whales exposed themselves to increased attack by leaving the rosette, flanking the isolated individual, and leading it back into the formation. Despite these efforts, one sperm whale was killed and eaten and the rest were seriously, perhaps mortally, wounded. We also present details of two other encounters between sperm whales and killer whales that we observed. Although sperm whales, because of various behavioral and morphological adaptations, were previously thought to be immune to predation, our observations clearly establish their vulnerability to killer whales. We suggest that killer whale predation has potentially been an important, and underrated, selective factor in the evolution of sperm whale ecology, influencing perhaps the development of their complex social behavior and at-sea distribution patterns.     

DIVING BEHAVIOR OF THE SAIMAA RINGED SEAL (PHOCA HISPIDA SAIMENSIS NORDQ.)

The activity and diving patterns of four adult Saimaa ringed seals ( Phoca hispida saimensis , a landlocked subspecies living in Lake Saimaa, Finland) were examined during spring, summer, and autumn by the use of VHF-transmitters. Over 17,000 dives were registered. The duration of the dives and diving patterns differed among individuals. The mean duration of dives increased from spring to autumn; e.g. , in one individual the mean dive duration increased from 6 min in June to 10.5 min in October. The haul-out periods of one individual in May to early June made up 46.2% of its total activity budget, but in another individual in July to August the haul-out periods made up only 11% of the budget and the seal was submerged for 80% of the time. Periods of successive long duration dives (>10 min) were observed in three individuals in summer and autumn. The longest dive measured was 23 min. The duration of the periods containing long dives was often over three hours (maximum six hours) and the mean duration of the dives about 15 min. These long duration dives are assumed to be aerobic resting dives. Generally, the dives of the Saimaa ringed seal appear to be of longer duration than previously assumed.     

Distribution and diving behaviour of harp seals (<Emphasis Type="Italic">Pagophilus groenlandicus</Emphasis>) from the Greenland Sea stock

The distribution and diving behaviour of 16 adult harp seals (Pagophilus groenlandicus) from the Greenland Sea stock were studied in 1993 and 1999, using satellite-linked dive recorders (SDRs). The seals remained near the pack-ice edge in the Greenland Sea between breeding and moulting (April/May 1993; 6F) and during the first 7 weeks after moulting (June/July, 1999; 4F, 6M), there diving to depths of <100 m. In mid-July 1999, seven out of eight seals with active SDRs migrated into the Barents Sea, there diving to <400 m and sharing feeding grounds with the Barents Sea harp seal stock. Between September and December, six of these seals joined the eighth seal in the Denmark Strait until March 2000, there diving to depths of 100–400 m. Overall, dives were significantly deeper in the day and in winter than at night and in summer, with some regional differences. Harp seals are considered pack-ice-associated seals, but our tagged seals spent a considerable proportion of their time in open water, their distribution largely overlapping with that of capelin (Mallotus villosus).