Demography of the sessile rotifers, <Emphasis Type="Italic">Limnias ceratophylli</Emphasis> and <Emphasis Type="Italic">Limnias melicerta</Emphasis> (Rotifera: Gnesiotrocha), in relation to food (<Emphasis Type="Italic">Chlorella vulgaris</Emphasis> Beijerinck, 1890) density

Circadian rhythms occur widely amongst living organisms, often in response to diel changes in environmental conditions. In aquatic animals, circadian activity is often synchronised with diel changes in the depths individuals occupy and may be related to predator–prey interactions, where the circadian rhythm is determined by ambient light levels, or have a thermoregulatory purpose, where the circadian rhythm is governed by temperature. Here, these two hypotheses are examined using animal-attached accelerometers in juvenile freshwater sawfish occupying a riverine environment displaying seasonal changes in thermal stratification. Across seasons, diel patterns of depth use (shallow at night and deep in the day) tended to occur only in the late dry seasons when the water was stratified, whereas individuals were primarily shallow in the early dry season which featured no thermal stratification. Activity was elevated during crepuscular and nocturnal periods compared to daytime, regardless of the thermal environment. Our observation of resting at cooler depths is consistent with behavioural thermoregulation to reduce energy expenditure, whereas activity appears linked to ambient light levels and predator–prey interactions. This suggests that circadian rhythms in activity and vertical migrations are decoupled in this species and respond to independent environmental drivers.     

Temperature as a cue for the periodicity in feeding of Brachycentrus occidentalis (Insecta: Trichoptera)

Filtering behaviour of two groups of Brachycentrus occidentalis larvae exposed to a diel temperature cycle and a LD12:12 photoperiod, was observed for 5 days at 4-hr intervals. Both groups were then subjected to 5 days of continuous light; however, a diel temperature pattern was maintained for only one group. Behaviour of larvae showed a diel periodicity under the initial conditions that was quickly lost in continuous light and constant temperature. Under continuous light and daily fluctuating temperatures, larvae could maintain a diel periodicity. However, the amplitude of the periodicity was moderated.     

Summer diel activity and movement of adult brown trout in high-elevation streams in Wyoming, U.S.A.

The variation in diel activity patterns was determined by implanting transmitters in adult brown trout of various sizes in two montane streams in two summers in the central Rocky Mountains, U.S.A. More brown trout tended to be active, and to move greater distances, each hour during twilight and night than during the day. Mean hourly light intensity was significantly negatively correlated with the proportion of fish active and with the distance moved. Maximum water temperature was not correlated with any measure of diel activity. Brown trout displayed fidelity to diurnal positions within a given diel cycle. Overall, fish were active an average of 11 h, had an average home range of 41 m, and moved an average of 121 m over the diel cycle. Fish length and distance moved were significantly positively correlated. Water temperature, predator avoidance, and foraging tactics may influence these patterns of diel behaviour in adult brown trout in U.S. streams.     

Diel drift of Chironomidae in a large lowland river (Central Poland)

The diel drift patterns of Chironomidae larvae were investigated in a seventh order section of the Warta River (Central Poland) over two diel cycles during May 1989. Three nets (mesh size 400 m) were installed in a cross section of the Warta River.The estimated drift density was low, but was comparable to that calculated for other large rivers. Spatio-temporal fluctuations in abundance and composition of macroinvertebrate drift, including Chironomidae, were observed with the highest density of drifting macrobenthos recorded near the depositional bank of this river. The ratio benthosdrift indicated differing propensities for of the older instars of a given chironomid taxon to drift. Orthocladiinae larvae were the most abundant subfamily of Chironomidae in drift but not in benthos, reaching up to 73% of the total drifting chironomid larvae. More taxa but fewer individuals (about 20% of the chironomid larvae collected) belonged to the tribe Chironomini, the dominant group in benthos.A major part of chironomid drift collection may represent behavioural drift because the net mesh size used in the Warta River was insufficient to catch the earliest instars (distributional drift). Both at the family and subfamily level chironomid larvae exhibited a distinct nocturnal drift periodicity. Nocturnal periodicity was documented for the dominant species, but due to the low density of many chironomid species, it was impossible to determine their diel drift pattern. Some Chironomidae appeared to be aperiodic.     

Temporal plasticity in thermal‐habitat selection of burbot Lota lota a diel‐migrating winter‐specialist

In this study, animal‐borne telemetry with temperature sensors was coupled with extensive habitat temperature monitoring in a dimictic reservoir, to test the following hypotheses: behavioural thermoregulation occurs throughout the year and temperature selection varies on a diel and seasonal basis, in a winter‐specialist diel‐migrating fish. Burbot Lota lota demonstrated nightly behavioural thermoregulation throughout the year, with a large seasonal shift between selection for very cold temperatures (<2° C) optimal for reproduction during the spawning period and selection for warmer temperatures (12–14° C) optimal for hunting and feeding during non‐reproductive periods. During daylight hours, while L. lota avoided habitats warmer than optimal for reproduction and feeding during the spawning and non‐reproductive periods, respectively, active selection was limited to selection for 4–6° C habitat during the prespawning period. Although behavioural thermoregulation explained the night‐time migration, behavioural thermoregulation only partially explained daytime behaviour, indicating that diel migration is best explained by a combination of factors. Thus, thermal‐habitat selection was a good predictor of night‐time habitat occupancy in a diel‐migrating species. Together, these results show that thermal‐habitat selection by fishes may be important throughout the year and a more seasonally plastic behaviour than previously recognized.     

A cold-adapted reptile becomes a more effective thermoregulator in a thermally challenging environment

Thermoregulation is of great importance for the survival and fitness of ectotherms as physiological functions are optimized within a narrow range of body temperature (T _b). The precision with which reptiles thermoregulate has been proposed to be related to the thermal quality of their environments. Although a number of studies have looked at the effect of thermal constraints imposed by diel, seasonal and altitudinal variation on thermoregulatory strategies, few have addressed this question in a laboratory setting. We conducted a laboratory experiment to test whether tuatara, Sphenodon punctatus (order Rhynchocephalia), a cold-adapted reptile endemic to New Zealand, modify their thermoregulatory behaviour in response to different thermal environments. We provided tuatara with three thermal treatments: high-quality habitat [preferred T _b (T _sel) could be reached for 8 h/day], medium-quality habitat (T _sel available for 5 h/day) and low-quality habitat (T _sel available for 3 h/day). All groups maintained body mass, but tuatara in the low-quality habitat thermoregulated more accurately and tended to maintain higher T _bs than tuatara in the high-quality habitat. This study thus provides experimental evidence that reptiles are capable of adjusting their thermoregulatory behaviour in response to different thermal constraints. This result also has implications for the conservation of tuatara. A proposed translocation from their current habitat to a higher latitudinal range within New Zealand (similar to the shift from our 8 h/day to our 5 h/day regime) is unlikely to induce thermoconformity; rather, tuatara will probably engage in more effective thermoregulatory behaviour.     

The physiological ups and downs of thermal variability in temperate freshwater ecosystems

Freshwater fish face a variety of spatiotemporal thermal challenges throughout their life. On a broad scale, temperature is an important driver of physiological, behavioural and ecological patterns and ultimately affects populations and overall distribution. These broad patterns are partly underpinned by the small-scale local effects of temperature on individuals within the population. Climate change is increasing the range of daily thermal variation in most freshwater ecosystems, altering behaviour and performance of resident fishes. The aim of this review is understanding how daily thermal variation in temperate rivers affects individual fish physiology, behaviour and overall performance. The following are highlighted in this study: (a) the physical characteristics of rivers that can either buffer or exacerbate thermal variability, (b) the effects of thermal variability on growth and metabolism, (c) the approaches for quantifying thermal variation and thermal stress and (d) how fish may acclimatize or adapt to our changing climate.     

Temporal Links in Daily Activity Patterns between Coral Reef Predators and Their Prey

Few studies have documented the activity patterns of both predators and their common prey over 24 h diel cycles. This study documents the temporal periodicity of two common resident predators of juvenile reef fishes, Cephalopholis cyanostigma (rockcod) and Pseudochromis fuscus (dottyback) and compares these to the activity and foraging pattern of a common prey species, juvenile Pomacentrus moluccensis (lemon damselfish). Detailed observations of activity in the field and using 24 h infrared video in the laboratory revealed that the two predators had very different activity patterns. C. cyanostigma was active over the whole 24 h period, with a peak in feeding strikes at dusk and increased activity at both dawn and dusk, while P. fuscus was not active at night and had its highest strike rates at midday. The activity and foraging pattern of P. moluccensis directly opposes that of C. cyanostigma with individuals reducing strike rate and intraspecific aggression at both dawn and dusk, and reducing distance from shelter and boldness at dusk only. Juveniles examined were just outside the size-selection window of P. fuscus. We suggest that the relatively predictable diel behaviour of coral reef predators results from physiological factors such as visual sensory abilities, circadian rhythmicity, variation in hunting profitability, and predation risk at different times of the day. Our study suggests that the diel periodicity of P. moluccensis behaviour may represent a response to increased predation risk at times when both the ability to efficiently capture food and visually detect predators is reduced.     

Individual differences in plasticity and sampling when playing behavioural games

When engaged in behavioural games, animals can adjust their use of alternative tactics until groups reach stable equilibria. Recent theory on behavioural plasticity in games predicts that individuals should differ in their plasticity or responsiveness and hence in their degree of behavioural adjustment. Moreover, individuals are predicted to be consistent in their plasticity within and across biological contexts. These predictions have yet to be tested empirically and so we examine the behavioural adjustment of individual nutmeg mannikins (Lonchura punctulata), gregarious ground-feeding passerines, when playing two different social foraging games: producer-scrounger (PS) and patch-choice (PC) games. We found: (i) significant individual differences in plasticity and sampling behaviour in each of the two games, (ii) individual differences in sampling behaviour were consistent over different test conditions within a game (PC) and over a six month period (PS), (iii) but neither individual plasticity nor sampling behaviour was correlated from one social foraging game to another. The rate at which birds sampled alternative tactics was positively associated with seed intake in PS trials but negatively associated in PC trials. These results suggest that games with frequency dependence of pay-offs can maintain differences in behavioural plasticity but that an important component of this plasticity is group- and/or context-specific.     

Thermal biology and mate acquisition in ectotherms

Many elements of the behavioural repertoire of ectothermic animals depend on body temperature. Under differing thermal conditions, behaviours in insects, reptiles and other terrestrial ectotherms may therefore vary widely, and in any given thermal regime there may be simple physical and physiological character differences between individuals that lead to a predictable variation of behaviour. Where mating behaviour patterns are involved, digferential fitness may result. Recent studies show that the interactions of physiology with behavioural ecology should be a fruitful area for future research.     

Diel and seasonal activity patterns of adult tautog, Tautoga onitis, in lower Chesapeake Bay, inferred from ultrasonic telemetry

Recent advances in ultrasonic telemetry have enabled researchers to simultaneously collect telemetry data on numerous tagged individuals without remaining in direct contact with them. We present a low cost, commercially available automated acoustic receiver system from which presence/absence data can be used to infer diel activity patterns. We monitored a large, temperate labrid, Tautoga onitis, but the methods and analyses presented here have direct application to other marine species. Data were analyzed using graphical analysis, harmonic analysis, and empirical eigenfunction analysis. Diel activity patterns were expressed in the context of photoperiod, water temperature, and tidal flow. Detection periodicity was strongly diel (24-h cycle), with detection predominantly during daylight hours only (diurnal). Diurnal detection persisted throughout the study. Daily detection was greatest in the morning or afternoon during slack tide. Daily detection varied seasonally with ambient water temperature and was greatest between 13–20°C, corresponding to the primary fishing seasons for Tautoga onitis in spring and fall. This study represented the first use of automated acoustic receivers to investigate seasonal changes in diel activity patterns of adult tautog and only the second application of ultrasonic telemetry to study this species. This study was also the first to address diel activity patterns of adult tautog in the southern range of this species' distribution.Senior author     

Seasonal variations in the behavioural thermoregulation of roosting Humboldt penguins ( Spheniscus humboldti) in north-central Chile

We examined the thermoregulatory behaviour (TRB) of roosting Humboldt penguins (Spheniscus humboldti) in north central Chile during summer and winter, when ambient temperatures (T_a) are most extreme. Each body posture was considered to represent a particular TRB, which was ranked in a sequence that reflected different degrees of thermal load and was assigned an arbitrary thermoregulatory score. During summer, birds exhibited eight different TRBs, mainly oriented to heat dissipation, and experienced a wide range of T_a (from 14 to 31°C), occasionally above their thermoneutral zone (TNZ, from 2 to 30°C), this being evident by observations of extreme thermoregulatory responses such as panting. In winter, birds exhibited only three TRBs, mainly oriented to heat retention, and experienced a smaller range of T_a (from 11 to 18°C), always within the TNZ, even at night. The components of behavioural responses increased directly with the heat load which explains the broader behavioural repertoire observed in summer. Since penguins are primarily adapted in morphology and physiology to cope with low water temperatures, our results suggest that behavioural thermoregulation may be important in the maintenance of the thermal balance in Humboldt penguins while on land.     

Thermoregulatory behaviour explains countergradient variation in the upper thermal limit of a rainforest skink

The vulnerability of a terrestrial ectotherm to high environmental temperatures depends on the animal's thermal physiology and thermoregulatory behaviour. These variables – environment, physiology, and behaviour – interact with each other, complicating assessment of species vulnerability to global warming. We previously uncovered a counterintuitive pattern in rainforest sunskinks Lampropholis coggeri: a negative relationship between their critical thermal maximum (CT_max) and the temperature of their environment. Could this result be explained by a three‐way interaction between environment, physiology, and behaviour? Here we find that sunskink thermal preference is correlated positively with CT_max, but, importantly, skinks from hotter environments prefer lower temperatures than conspecifics from cooler environments. In an acclimation experiment, we find that CT_max is plastic and shifts in alignment with acclimation temperature. We also found heritable variation in this trait in a common garden study, but this variation was small relative to the plastic shifts observed in CT_max. Thus, our previous observation of a negative correlation between field CT_max and temperature is explained, at least in part, by the lizard's thermoregulatory behaviour: lizards from hot environments preferentially choose cool microenvironments, and their physiology acclimates to these cooler experienced temperatures. Our results suggest that behavioural adjustments to the environment can produce countergradient variation in physiological traits. More broadly, our work underscores the importance of interactions between environment, behaviour, and physiology in ectotherms. Understanding these interactions will be crucial in assessing vulnerability to climate change.     

Relaxed circadian rhythm in zooplankton along a latitudinal gradient

To test whether aquatic invertebrates are able to adjust their diel migratory cycle to different day length and presence of predators, we performed standardized enclosure experiments in shallow lakes at four different latitudes from southern Spain, with strong night–day cycles, to Finland where daylight is almost continuous during summer. We show here that nearly continuous daylight at high latitudes causes a relaxation in diel migratory behaviour in zooplankton irrespective of predation risk. At lower latitudes, however, similar conditions lead to pronounced diel rhythms in migration. Hence, zooplankton may show local behavioural adaptations in their circadian rhythm. They are also able to make risk assessments as to whether diel migration is beneficial or not, manifested in a lack of diel migration at near constant daylight, irrespective of predator presence. Our results provide an additional explanation to previous knowledge regarding diel migrations among aquatic invertebrates by showing that both physical (light) and biological (predation) factors may affect the migratory behaviour.     

Seasonal acclimation of preferred body temperatures improves the opportunity for thermoregulation in newts

Seasonal acclimation and thermoregulation represent major components of complex thermal strategies by which ectotherms cope with the heterogeneity of their thermal environment. Some ectotherms possess the acclimatory capacity to shift seasonally their thermoregulatory behavior, but the frequent use of constant acclimation temperatures during experiments and the lack of information about thermal heterogeneity in the field obscures the ecological relevance of this plastic response. We examined the experimentally induced seasonal acclimation of preferred body temperatures (T(p)) in alpine newts Ichthyosaura (formerly Triturus) alpestris subjected to a gradual increase in acclimation temperature from 5°C during the winter to a constant 15°C or diel fluctuations between 10° and 20°C during the spring/summer. Both the mean and range of T(p) followed the increase in mean acclimation temperature without the influence of diel temperature fluctuations. The direction and magnitude of this acclimatory capacity has the potential to increase the time window available for thermoregulation. Although thermoregulation and thermal acclimation are often considered as separate but coadapted adjustments to thermal heterogeneity, their combined response is employed by newts to tackle seasonal variation in a thermoregulatory-challenging aquatic environment.     

Shade as a thermoregulatory resource for captive chimpanzees

In nature, the thermoregulatory strategies of species have evolved in response to the environmental conditions in which they live. Primates display extensive behavioural flexibility but few have examined the role of this behavioural flexibility with regard to thermoregulation. Chimpanzees, under free-living conditions, utilise natural vertical light/thermal gradients in rainforest canopies as a means of thermoregulation, moving along the gradient to maintain homoeostasis. In contrast, captive chimpanzees are often housed at latitudes outside their natural range under conditions that do not allow for this natural behavioural thermoregulation. We investigated the use of shade by captive chimpanzees as a behavioural thermoregulatory strategy at the Johannesburg Zoo, South Africa. We performed behavioural observations over the austral winter/spring period on a group of captive chimpanzees recording their behaviour in relation to shade use. Despite experiencing temperatures in Johannesburg below or just within their thermoneutral zone, chimpanzees consistently spent more time in shade than in direct sun. This pattern of shade use was most pronounced for the hotter midday period than for other times, was not dependent on the overall availability of shade within the enclosure and was not predicted by mean daily or hourly temperatures nor thermal maxima or minima. Instead, ultraviolet radiation and humidity levels appear to predict the observed patterns of shade utilisation and these findings suggest that chimpanzees in captivity adopt a sun-avoidance strategy, possibly as a result of the rapid heat gain associated with their dark skin and pelage. The findings suggest that chimpanzees are flexible when responding to the thermal challenges associated with housing outside of their natural environment.     

Home-range use by coyotes in Idaho

Radio-collared coyotes (Canis latrans) were relocated every 15 min during continuous 24-h sampling periods. The data were used to estimate patterns of home-range use by coyotes. Utilization of the the home range was found to vary spatially and behaviourally. Spatial use was determined by relative amounts of time coyotes spent and amounts of distance they travelled within each are of their home ranges. Behavioural use was based on identification of three types of movement patterns that werew postulated to represent three general kinds of behaviour: (1) resting behaviour, (2) hunting or investigative behavour, and (3) ranging or traveling behaviour. Spatial and behavioural uses of the home range area were found to be interrelated; core areas in which animals spent most of their time were also used primarily for resting or hunting. In areas in which animals spent little time, coyotes exhibited primarily ranging behaviour. Use patterns were postulated to be the result of coyotes' selection of areas due to unique vegetal, faunal, or physiogfaphic characteristics. Temporal variatrions in home-range use were found and were postulated to result from seasonal and diel changes in coyote behaviour due to the annual reproductive cycle, the seasonal and diel cycle of temperature, possible cycles in prey behaviour.     

Mitigating thermal effect of behaviour and microhabitat on the intertidal snail Littorina saxatilis (Olivi) over summer

High shore intertidal ectotherms must withstand temperatures which are already close, at or beyond their upper physiological thermal tolerance. Their behaviour can provide a relief under heat stress, and increase their survival through thermoregulation. Here, we used infrared imaging to reveal the thermoregulatory behavioural strategies used by the snail Littorina saxatilis (Olivi) on different microhabitats of a high shore boulder field in Finistère (western France) in summer. On our study site, substrate temperature is frequently greater than L. saxatilis upper physiological thermal limits, especially on sun exposed microhabitats. To maintain body temperatures within their thermal tolerance window, withdrawn snails adopted a flat posture, or elevated their shells and kept appended to the rock on the outer lip of their aperture with dried mucous (standing posture). These thermal regulatory behaviours lowered snail body temperatures on average by 1–2 °C. Aggregation behaviour had no thermoregulatory effect on L. saxatilis in the present study. The occupation of biogenic microhabitats (barnacles) was associated with a 1 °C decrease in body temperatures. Barnacles and microhabitats that experienced low sun exposure, low thermal fluctuations and low thermal maxima, could buffer the heat extremes encountered at high shore level especially on sun exposed microhabitats.     

Spatial and Temporal Variation of Macroinvertebrate Drift in Two Neotropical Streams1

The detection of spatial variation in macroinvertebrate drift depends on the spatial scale of investigation in streams of the La Selva Biological Station, Costa Rica. Drift samples were taken in a spatially nested design, with two streams, two reaches per stream, two riffles per reach, and four replicate samples per riffle. Drift showed little variation among streams, but varied significantly at the scales of reach and riffle, with variation among samples also high. In addition, sampling took place at two temporal scales: diel and at two different periods that differed in rainfall conditions. Drift diel periodicity was a clear pattern, while only density of individuals varied among sampling periods. This is the first study of macroinvertebrate drift at multiple spatial scales, despite the recognition that multi‐scale studies are essential for a more complete understanding of community patterns and processes.     

The principles of collective animal behaviour

In recent years, the concept of self-organization has been used to understand collective behaviour of animals. The central tenet of self-organization is that simple repeated interactions between individuals can produce complex adaptive patterns at the level of the group. Inspiration comes from patterns seen in physical systems, such as spiralling chemical waves, which arise without complexity at the level of the individual units of which the system is composed. The suggestion is that biological structures such as termite mounds, ant trail networks and even human crowds can be explained in terms of repeated interactions between the animals and their environment, without invoking individual complexity. Here, I review cases in which the self-organization approach has been successful in explaining collective behaviour of animal groups and societies. Ant pheromone trail networks, aggregation of cockroaches, the applause of opera audiences and the migration of fish schools have all been accurately described in terms of individuals following simple sets of rules. Unlike the simple units composing physical systems, however, animals are themselves complex entities, and other examples of collective behaviour, such as honey bee foraging with its myriad of dance signals and behavioural cues, cannot be fully understood in terms of simple individuals alone. I argue that the key to understanding collective behaviour lies in identifying the principles of the behavioural algorithms followed by individual animals and of how information flows between the animals. These principles, such as positive feedback, response thresholds and individual integrity, are repeatedly observed in very different animal societies. The future of collective behaviour research lies in classifying these principles, establishing the properties they produce at a group level and asking why they have evolved in so many different and distinct natural systems. Ultimately, this research could inform not only our understanding of animal societies, but also the principles by which we organize our own society.