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.     

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 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.     

Temporal variation and distribution of trace metals in freshwater and fish from Calabar River, S.E. Nigeria

An investigation on the abundance and distribution of trace metals (Fe, Cu, Zn, Mn, Cr, Cd and Pb) in water, and nine species of fish samples from Calabar river was carried out in 1992. The concentrations of iron (6000–7240gl^–1), zinc (4910–7230gl^–1), and cadmium (3–7gl^–1) showed moderate pollution while those of copper (420–630gl^–1), manganese (23–48gl^–1), chromium (<10–20gl^–1) and lead (<1–10gl^–1) in water were well below WHO permissible levels. Significant seasonal changes (0.001p0.25) were obtained for iron, copper, zinc, manganese and cadmium in water. Furthermore, iron, zinc and cadmium showed statistically significant spatial changes (0.005p0.10). Of the nine fish species studied, no statistically significant relationship between body weight and the concentrations of the metals was observed. The concentrations of the metals per mean total body weight apparently decreases in the order Fe>Zn>Cu>Mn>Pb>Cd=Cr and were within the limits that were safe for consumption.     

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.     

Lead and cadmium in urban allotment and garden soils and vegetables in the United Kingdom

In order to assess the intake of lead and cadmium by consumers of home grown vegetables in urban areas, replicated experimental plots of uniform size, comprising summer and winter crops, were established in 94 gardens and allotments in nine towns and cities in England.The geometric mean lead and cadmium concentrations for the soils (n = 94) were 217 g g^–1 (ranging from 27 to 1,676 g g^–1) and 0.53 g g^–1 (<0.2–5.9 g g^–1), respectively. Compared with agricultural soils, the garden and allotment soils contained elevated levels of lead but not cadmium.Lead concentrations in the vegetables ranged from <0.25 g g^–1 to 16.7 g g^–1 dry weight and cadmium concentrations ranged from <0.025 g g^–1 to 10.4 g g^–1 dry weight. Lead concentrations were higher than reported background levels, although <1% exceeded the statutory limit for saleable food in the UK (1 g g^–1 fresh weight). Cadmium concentrations were generally similar to background levels.     

Ecological constraints on the behaviour of mother long-tailed macaques (Macaca fascicularis)

Summary The ecological consequence of the effects of different social conditions on the behaviour of females with infants were examined in two different-sized groups of long-tailed macaques (Macaca fascicularis) that inhabited adjacent home ranges in Gunung Leuser National Park in Indonesia. This study aims to test predictions of the within-group competition/predation avoidance hypothesis of social organization. Females, particularly those carrying an infant, are considered to experience the most direct effects of environmental constraints on fitness prospects. Data on maintenance behaviour, height in canopy and spatial position were collected for five mothers in a small group and eight mothers in a large group. Comparisons were made on the basis of group size and dominance rank. Mothers from the large group spent less time feeding on clumped fruits and more time foraging on dispersed food items. Lower-ranking mothers, undergoing the highest rate of within-group competition, foraged most. For macaques in the study area, predation risk is considered to decrease with height in the canopy. Mothers in the small group, assumed to be more susceptible to predation, remained substantially higher in the canopy than mothers in the large group. In both groups, mothers adjusted their height to their spatial position in the group. Lower-ranking mothers were more often found without neighbours, presumably to avoid feeding competition. As a result they stay higher in the canopy than higher-ranking mothers. The results clearly demonstrate that individual females in the same reproductive state and living in the same area, but in different social environments, adapted their behaviour according to the hypothesis. Correspondence to: D.R. Vos     

Environmental Accumulation of Airborne Fluorides in Romania

The nature and extent of pollution from an aluminium smelter and a fertiliser factory in Romania were studied. These are large industrial complexes, and both types of industry are known to release fluorides into the atmosphere. In grass samples collected from around the aluminium smelter, the maximum fluoride levels were found to be 4023mgkg^–1 and 162mgkg^–1 in unwashed and washed grass samples respectively, and 89mgkg^–1 in soils. For the fertiliser factory, the maximum levels in washed grasses were found to be 207mgkg^–1, and 11mgkg^–1 in the soils. In both locations, these maximum values were obtained in samples collected from within 200m of the factory limits, and compare with regional background levels of less than 10mgkg^–1 for grasses and 2mgkg^–1 for soils. The high fluoride levels of fluoride in the grasses are sufficient to give cause for concern for the effects that these could have on the local population and on grazing animals.     

Mother-pup separation and adoption in northern elephant seals

Summary The evolution of fostering behavior, parental care directed toward another's young, has been the focus of much recent interest. During a five-year study of northern elephant seals (Mirounga angustirostris) at Año Nuevo, California, we recorded the frequency of mother-pup separation, reunion, and adoption of orphaned pups in crowded and low-density breeding areas. While most females nursed their own pup exclusively until it was weaned, many females, especially young mothers (age 3–5 years), were unable to raise a pup successfully. In the crowded main breeding harem on Año Nuevo Island, 24 to 47 percent of the pups born each year were separated from their mothers from 1977 to 1980. Mother-pup separation and pup mortality were associated with the following inter-related factors: 1) female density; 2) weather and tidal conditions; 3) topographical features of the breeding areas (i.e., degree of exposure to high tides and surf); and 4) the proportion of young, maternally inexperienced females pupping in a particular area. Most motherpup separations were caused directly by 1) adult males moving through the harem; 2) pups wandering from their mothers; 3) female aggression; and 4) inclement weather. Most of the separations, as well as adoptions, occurred when pups were quite young. Mother-pup recognition appeared to be based on a combination of acoustic, visual, and olfactory cues, and most mother-pup reunions were effected by the female rather than her pup. On the main island breeding area, 572 orphans were marked. Of these, five percent relocated their mother, 27 percent were adopted or frequently cared for by foster mothers, and 68 percent were not adopted, or rarely fostered. The survival of an orphan was clearly contingent on the amount of care it received; most orphans which were not nursed or protected by females died before reaching 6 weeks of age. Frequently, an adopted orphan's foster mother was in the stage of lactation which corresponded closely to that of its own mother. The most common fostering event involved females that had lost their own pup and adopted a single orphan. Other pupless females attempted to steal a pup, cared for a pup while it was still with its mother, adopted a weaned pup, adopted two pups, or indiscriminately nursed any orphaned pup that approached. Some females kept their own pup in addition to fostering in alien pup. Most foster mothers were young and had little or no previous maternal experience. The formation of large, high-density breeding rookeries, due to a scarcity of suitable breeding sites, results in frequent mother-pup separations, especially during inclement weather and tidal conditions. Many opportunities for adoptive behavior are therefore presented, because of the great number of orphans and pupless females. Increased maternal experience appears to be a benefit associated with adoption. Some instances of fostering behavior may also be based on reproductive errors on the part of the foster mother.     

Tolerance and ultrastructural responses of branchial chloride cells to salinity changes in the euryhaline teleost Oreochromis mossambicus

The changes of intercellular organization and junctional structures in branchial chloride cells reflect respective functions in different salinities. Under TEM, leaky junctions and intercellular digitations occurred between branchial chloride cells of Oreochromis mossambicus Peters adapted to seawater, but not in those adapted to freshwater. The fish transferred directly to 30 S seawater from freshwater died within 6 h, and their chloride cells developed neither leaky junctions nor interdigitations. The fishes acclimated to 20 S seawater for 12 h did not develop the characteristics of seawater-adapted chloride cells and died after transfer to 30 S seawater. The fish acclimated to 20 S seawater for 24 h started to develop seawater-adapted chloride cells, and were able to survive when transferred to 30 S seawater. Thus, the development of leaky junctions and interdigitations in branchial chloride cells appears to correlate to seawater adaptation in O. mossambicus. These changes of seawater-adapted chloride cells seem to be associated with the increase of ion permeability in the gill of teleosts adapted to seawater rather than those adapted to freshwater.     

Do Tidal-Channel Turbulence Measurements Support k−5/3?

We seriously challenge the belief that tidal-channel data of Grant et al. support the theory for the classical energy-spectrum behavior, k ^–5/3 in the inertial subrange [where k is wave number and (k) is the one-dimensional energy-spectrum function]. The behavior revealed by our re-examination is explained in terms of a more recent theory.     

Adaptation and equivalence in open-ocean phytoplankton

Two marine phytoplankton species of the North Atlantic Ocean have inherently differt growth responses to changing nutrient conditions. Yet they co-occur. Thus, they can be compared under the same conditions via equivalence, provided that their adaptedness is linked to their responsiveness as follows. If Species x is adapted to nutrient changes, then it is responsive to these conditions and, certainly, if it is responsive to these conditions, then it is adapted to them. There is another expression that is equivalent to this one (that says the same thing): either Species x is adapted and responsive to these conditions or it is neither adapted nor responsive to them. The option just mentioned is resolved so that Emiliania huxleyi as Species x is adapted and responsive and Cyclococcolithus leptoporus as Species x is not adapted and not responsive. All the frequently observed species in the winter half of the year, November to April, from 1961 to 1972 are like E. huxleyi in being adapted and responsive or like C. leptoporus in being unadapted and unresponsive.Contribution No. 5806 from the Woods Hole Oceanographic Institution     

Temporal trends in urban and rural blood lead concentrations

A long term programme to assess the impact of reduced lead emissions in the UK has been established in central London and rural Suffolk. Blood lead concentrations of 5 and 6 year old children and their mothers both show an urban-rural difference of about 1.5 g/dl, which is related to the urban-rural differences in air and dust lead concentrations between areas.This paper is also being published in Proceedings of the Conference on Lead in the Home Environment and is one of several selected from SEGH sponsored conferences for simultaneous publication in this journal.     

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.     

The methane mussel: roles of symbiont and host in the metabolic utilization of methane

Methane mussels (Bathymodiolus sp., undescribed; personal communication by R. Turner to CRF) were collected in September 1989 and April 1990 from offshore Louisiana in the Gulf of Mexico. These mussels contain endosymbiotic methane-oxidizing bacteria and are capable of utilizing environmental methane as a source of energy and carbon. Oxygen consumption, methane consumption, and carbon dioxide production were measured in mussels with intact symbionts, functionally aposymbiotic mussels, and separated symbiont preparations under controlled oxygen and methane conditions, in order to study the roles of the symbionts and the hosts in methane utilization. The association was found to be very efficient in fixing methane carbon (only 30% of CH₄ consumed is released as CO₂), and to be capable of maximal rates of net carbon uptake of nearly 5 mol g^-1 h^-1. Rates of oxygen and methane consumption were dependent upon oxygen and methane concentrations. Maximal consumption rates were measured at 250 to 300 M O₂ and 200 to 300 M CH₄, under which conditions, oxygen consumption by the gill tissues (containing symbionts) had increased more than 50-fold over rates measured in the absence of methane. A model is proposed for the functioning of the intact association in situ, which shows the symbiosis to be capable of achieving growth rates (net carbon assimilation) in the range of 0.003 to 0.50% per day depending upon oxygen and methane concentrations. Under the conditions measured in the seep environment (200 M O₂, 60 M CH₄), a mussel consuming methane at rates found to be typical (4 to 5 mol g^-1 h^-1) should have a net carbon assimilation rate of about 0.1% per day. We suggest that the effectiveness of this symbiosis arises through integration of the morphological and physiological characteristics inherent to each of the symbiotic partners, rather than from extensive specialization exhibited by other deep-sea chemotrophic associations.     

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     

Suspension-feeding and assimilation efficiency in Lanice conchilega (Polychaeta)

In laboratory experiments, measurements of ingested ration, assimilated ration and assimilation efficiency in the polychaete Lanice conchilega (Pallas) were carried out at 12°C and at a food concentration of 40x10⁶ cells of Dunaliella marina/l. Over periods of 20 to 46 days, the ingested ration was determined photometrically by the continous automatic recording apparatus described by Winter (1973). The assimilated ration was calculated from the calorific content of the food ingested minus the calorific content of the faeces produced. Within the range of body size investigated (1.3 to 35.0 mg dry-tissue weight), the food intake (=filtration rate=F) was found to be a linear function of body size (W) according to the equation F=0.88 W ^0.32. The daily food intake-expressed as percentage of dry-tissue weight-decreased from 35.7 to 3.9% with increasing body size. The assimilation efficiency was found to vary between 70.6 and 77.2% and showed no significant correlation with body size. The absolute amounts of food retained from suspension and the assimilation efficiencies calculated for L. conchilega are within the range typical for obligatory suspension-feeding organisms. This implies that L. conchilega is capable of completely replacing deposit-feeding by suspension-feeding.This work was made possible through a research grant from the Deutsche Forschungsgemeinschaft in connection with the program Litoralforschung-Abwässer in Küstennähe.     

Brood defense and age of young: a test of the vulnerability hypothesis

Summary In many altricial species including the great tit (Parus major) the intensity of brood defense against predators has been shown to increase with the age of the offspring. This effect has been ascribed amongst others to the young becoming more vulnerable as they age (vulnerability hypothesis). In a great tit population suffering heavy losses from brood depredation by the great spotted woodpecker (Dendrocopus major), we rendered first and second broods more vulnerable by artificially enlarging the entrance of the nest hole. Contrary to the vulnerability hypothesis, 16 experimental pairs defended their brood against a dummy great spotted woodpecker less vigorously than did 16 control pairs. Nest concealment behavior potentially compromising active defense was minimized by simultaneous playback of nestling distress calls, thus simulating the act of nest predation. This leaves the brood value hypothesis as an alternative functional explanation of the defense level — age effect. It predicts that parents should defend their brood in proportion to the reproductive value (or some more suitable cohortal equivalent measure) of their offspring. At present, this explanation pertains to one predator species. In first broods, but not in second broods, males defended them more vigorously than did their females. While this parallels previous experiments on brood defense against predators posing a much greater risk to the parents, two functional explanations previously put forward can hardly apply.     

Ammonium enhancement of dark carbon fixation and nitrogen limitation in zooxanthellae symbiotic with the reef coralsMadracis mirabilis andMontastrea annularis

The nutrient status (limitation vs sufficiency) of dinoflagellates (zooxanthellae) symbiotic with reef corals in Bermuda was assessed in 1989 and 1990 by measuring the enhancement of dark carbon fixation with 20 M ammonium by isolated symbionts. A colony ofMadracis mirabilis was kept in the laboratory and fed daily or starved for one month. Symbionts from fed portions of the colony had ammonium-enhancement ratios (NH _4dark ⁺ ; SW_dark;SW=seawater without added ammonium) similar to those of the original field population (1.2 to 1.3). Ammonium-enhancement ratios increased with starvation of the host (x1.7) as did values forV _D:V _L [(ammonium dark rate-seawater dark rate): light rate in seawater]. Both parameters indicated decreasing nitrogen sufficiency of the algae when the host was not fed, but starvation appeared to affect these algae less than symbionts of sea anemones. Field samples of zooxanthellae fromM. mirabilis (Three Hill Shoals and Bailey's Bay Flats) yielded results similar to those for fed corals, but those taken from Bailey's Bay Flats in May 1990 yielded exceptionally high values for enhancement (>3) andV _D:V _L indicating pronounced nitrogen limitation at the time of sampling. We sampled zooxanthellae from populations ofMontastrea annularis at 8 m (Three Hill Shoals) and 24 m (Soldier's Point) depths. Enhancement andV _D:V _L values for zooxanthellae from the 8 m corals were density-dependent: symbionts from corals with normal symbiont densities displayed the most nitrogen limitation (enhancement values=1.4 to 2.0), while those from bleached corals with lower density exhibited enhancement andV _D:V _L values typical of nitrogen-sufficient algae. Symbionts isolated from the 25 m corals yielded the highest values, and appeared to exhibit the least nitrogen-sufficiency for this species.     

Daily ration and specific daily ration of the chaetognath Sagitta enflata

Digestion times, obtained directly from laboratory feeding, were used in conjunction with a day/night series of field samples to estimate the daily ration and specific daily ration of Sagitta enflata Grassi, the most abundant chaetognath in the Gulf Stream near Miami, Florida (USA), during winter and early spring. Feeding was independent of time of day and increased with increasing chaetognath size from 1.7 prey day^-1 for the 11.5 mm size class to 2.9 prey day^-1 for the 21.5 mm class. The diet of s. enflata consisted of 94.8% copepods having a mean weight of 21.8 g, and 5.2% prey chaetognaths estimated to be two-thirds the length of the predators. The daily ration of S. enflata from 12.5 to 20.5 mm in length increased from 48.3 to 143.3 g of prey dry weight and from 18.5 to 48.1 g of prey carbon. The specific daily ration declined from 0.12 to 0.08 dry weight basis, and from 0.26 to 0.14, carbon basis, over the same size range. Prey chaetognaths contributed as much as 51% of the ration on a carbon basis.