Familiarity with adults,but not relatedness,affects the growth of juvenile red-backed salamanders (<Emphasis Type="Italic">Plethodon cinereus</Emphasis>)

The relative roles of kinship and familiarity in affecting an individual’s growth and fitness are not easy to disentangle. Not only is an individual more likely to have prior behavioral interactions with conspecifics in close proximity, it may also be related (in terms of kinship) to those nearby conspecifics. While some studies have inferred that kin discrimination affects fitness correlates, other studies found that familiarity alone can reduce aggressive interactions, thus increasing fitness. These studies have all focused on intra-age class pairs or groups. However, many animals interact with conspecifics from different cohorts. In many populations, adults of Plethodon cinereus territorially defend rocks and logs that retain moisture and food resources. We investigated whether juveniles of P. cinereus grew more in the presence of adults that were relatives or familiar. We collected pairs of juveniles and adults found under the same cover objects in the forest (familiar) and pairs of juveniles and adults found under different cover objects, approximately 10 m apart (unfamiliar). We determined parentage and relatedness of the adult–juvenile pairs and then placed these pairs in semi-natural mesocosms for 17 days. We found that juveniles housed with familiar adults had significantly greater increases in mass and snout–vent length than juveniles housed with unfamiliar adults in 2006 but not in 2007. Relatedness had no effect on growth. In addition, juveniles cohabitating with adults were not more likely to be their offspring. At least under certain environmental conditions, familiarity with adults, independent of relatedness or parentage, increased the growth of juvenile salamanders.     

Juvenile–adult relationship in the gregarious ophiuroid<Emphasis Type="Italic"> Ophiothrix fragilis</Emphasis> (Echinodermata): a behavioral and morphological study

Ophiothrix fragilis forms dense beds in the North Sea–English Channel region, where juveniles are exclusively found on adults. The aim of this study was to see how the behavior and morphology of juveniles could help elucidate the close juvenile–adult relationship found in this species. Juveniles are found on the disk, arms and in the bursae of adult conspecifics, the ones on the disks being significantly larger. No clear advantage seems to be gained by the juveniles being in the bursae, and their presence there is most likely due to juvenile movement on adults and to and from adults. Hooked spines serve as anchory organs during the early life of the juvenile, but growth of the arms enhances its anchory capabilities and the hooked spines become secondary in that respect. They regress as ophiuroids become older, to the advantage of the other spines used in suspension feeding. Juveniles are attracted to conspecifics, and true gregarious behavior has been observed. The tip of each arm, the terminal tentacle, plays a major role in distance chemoattraction, with juveniles needing at least one intact terminal tentacle to be able to initiate a response. Electron microscope observations of the terminal tentacle permitted us to recognize two different potential receptor structures designated stäbchens. The first possesses one long projecting cilia and is mostly present around the base of the terminal tentacle, while the other has one to five short projecting cilia and is mostly found on the tip. No receptors are found on the shaft. Receptors are not associated with secretory structures. Juveniles and adults are closely associated with one another, and both the morphology and behavior of juveniles play an important role in that relationship.Communicated by O. Kinne, Oldendorf/Luhe     

Environmental influence on the locomotor activity rhythm of Talitrus saltator (Crustacea: Amphipoda)

The persistence and precision of the endogenous, nocturnal, circadian locomotor rhythm of the sand-beach amphipod Talitrus saltator is characterised, and the influence of substrate availability, photoperiod, temperature and relative humidity as potential environmental synchronizers of the rhythm is assessed. Individual, cyclic light-dark and temperature regimes synchronize and modify the laboratory activity pattern, but substrate availability and relative humidity have no significant long-term effect. Under simulateneous experimental regimes simulating field conditions of photoperiod, temperature and relative humidity the rhythm is entrained solely by photoperiod. The implications are discussed in relation to field behaviour.     

The influence of kinship on foraging competition in Siberian jays

Foraging competition in Siberian jay groups was examined in relation to dominance and kinship to determine whether juvenile offspring, by associating with adults, gained in food acquisition relative to juvenile immigrants. Members of the adult pair were dominant over juvenile cohort members and males were dominant to females, although an inter-sexual hierarchy, with male juveniles occasionally overlapping adult females, was suggested. Few competitive asymmetries were found between adults and retained offspring or adults and immigrant juveniles when they were competing for food together, but in kin and non-kin foraging groups, respectively. Male offspring visited the bait site more frequently than adult males, and female immigrants spent less time at the bait site than adult females. Under these circumstances, hoarding activities may limit the ability of alpha members to control resources. In mixed groups containing both juvenile offspring and juvenile immigrants, no difference was found in the number of visits made to the bait site, although load sizes and foraging rates were lower for immigrant birds. Retained juveniles obtained greater load sizes and foraging rates when associating with adults. The social dominance of parents suggests that they control juvenile foraging. Although offspring benefit in the presence of adults, adults may incur a cost to their restraint by spending more time at the bait site when competing with immigrants. These results extend conclusions from previous work describing the role of selective tolerance by adults which relaxes competition with retained offspring in Siberian jay winter groups. The present findings suggest that offspring benefit in both immediate and future energy gains, which may have a direct influence on survival. Received: 18 September 1996 / Accepted after revision: 26 January 1997     

Differences in herbivore preferences, phlorotannin production, and nutritional quality between juvenile and adult tissues from marine brown algae

 Juvenile and adult marine organisms differ in their morphology, chemistry, physiology, behavior, and ecology. Because juvenile algae are thinner, smaller, and have more delicate tissues than adults, they are often assumed to be more susceptible to grazers. We examined within-species food preferences of four common generalist herbivores for juvenile and adult tissues of eight common brown algae in two-choice laboratory food-preference experiments. Our results showed that juvenile algae did not tend to be a preferred food of herbivores. Juvenile tissues were significantly preferred over adult tissues in only four of the 32 combinations of algae and herbivores tested. In 12 experiments, adult tissues were preferred over juvenile tissues, and no choice occurred in the remaining 16 experiments. When sea urchins exhibited a preference, it was always for adult tissues. The other three herbivores, an isopod and two snails, were more variable in their choices, sometimes preferring juveniles, sometimes adults, and sometimes having no preference. We measured nitrogen and phlorotannin concentrations in adult and juvenile seaweeds to see whether these parameters were correlated with herbivore food preferences. Nitrogen levels were similar in juveniles and adults of three algal species and were higher in juveniles of two. Phlorotannin levels were higher in juveniles of four species and lower in juveniles of one. The other three species showed no differences in phlorotannin levels. Phlorotannin concentrations decreased with increasing juvenile size in three species and increased with increasing size in one species. Neither nitrogen nor phlorotannin concentrations explained overall herbivore food preferences for algae of different stages. Our results suggest that preferences of certain grazers for juvenile algae are not as strong as previously assumed and are dependent on herbivore species. Preferences between juveniles and adults are likely to be determined by a combination of morphological and chemical features of the tissues and the unique responses of herbivore species to those features. Received: 10 April 2000 / Accepted: 19 November 2000     

Seasonal changes of locomotor activity patterns in ruin lizards Podarcis sicula

The daily pattern of locomotor activity of the ruin lizard Podarcis sicula in its natural environment changes from unimodal in spring (with only one activity peak per day) to bimodal in summer (with two well-separated activity peaks per day) and it becomes unimodal again in autumn. In order to establish whether such seasonal changes in pattern might be at least in part controlled by endogenous temporal programs, lizards were collected at different times of the year and immediately after capture their locomotor behavior was tested in the laboratory under constant temperature (29°C) and in darkness. For some individuals tested in the laboratory the locomotor pattern previously expressed in the field was known. Seasonal differences in pattern have been unequivocally found to have an endogenous component, as most lizards in constant conditions retained the locomotor pattern shown in the field during the same season. Besides, in the bimodal lizards the freerunning period of locomotor rhythms () was significantly shorter and circadian activity time (a) longer than in the unimodal ones. Altogether the data are compatible with the idea that both the interdependent changes of and a and the changes in locomotor pattern occurring seasonally in the circadian activity rhythms of P. sicula would depend on changes in the phase relationship between mutually coupled oscillators which drive these rhythms. Correspondence to: A. Foà     

The importance of life stage to population connectivity in whiting (Merlangius merlangus) from the northern European shelf

Juvenile movements are rarely considered in studies of connectivity. This study investigates movements in juvenile and adult life stages of an ecologically important temperate piscivore, the whiting (Merlangius merlangus). Adult movement was assessed from tag-recapture data collected between 1958 and 1980 around the coast of Scotland. Displacement distances ranged from 53 to 123 km. Adults rarely dispersed offshore in the North Sea, and exchange with the west of Scotland was negligible. Connectivity between nursery and spawning grounds was investigated using otolith microchemistry. Five juvenile nursery areas were discriminated with an accuracy of 75–85%. Comparing isotopic signatures of adults and juveniles indicated that several nursery areas contributed to each spawning aggregation. The lack of evidence for return migrations suggested an opportunistic and non-philopatric recruitment strategy within a single population unit. Differences in the proportion contribution of juveniles to adult aggregations also indicated that the “value” of nursery areas differed and the Scottish west coast could be viewed as a net source of recruits to the North Sea. Given the extent of juvenile dispersal, this study highlights the fact that all stages of the life cycle should be considered in studies of connectivity.     

Effects of sudden changes in salinity on endogenous rhythms of the spotted sea bass Lateolabrax sp.

The endogenous rhythm of oxygen consumption in juvenile spotted sea bass (Lateolabrax sp.) was measured to test the effects of sudden changes in salinity on the metabolic activity. Mean oxygen consumption rates of this euryhaline fish decreased by 13.5 to 16.0% and 25.3 to 36.4% when they were transferred from 31.5 to 15‰ seawater and to fresh water (0‰), respectively. The maximum rate of oxygen consumption was observed between 18:00 and 19:00 hrs local time, 1 to 2 h before sunset, even though they were kept in constant darkness. The peaks of oxygen consumption occurred in 23.2- and 23.3-h intervals, which correspond with a circadian rhythm, as revealed by maximum entropy spectral analysis. A markedly weakened rhythm in oxygen consumption occurred from 8 to 10 d after onset of the experiments. This study indicates that spotted sea bass can withstand sudden drops in salinity from 31.5‰ to fresh water, and yet maintain a regular though somewhat dampened endogenous rhythm of oxygen consumption. Received: 16 June 1997 / Accepted: 3 February 1998     

Circadian rhythms in the respiration of eight species of cardinal fishes (Pisces: Apogonidae): comparative analysis and adaptive significance

A comparative study was made of the entrained (12:12 LD) and endogenous (DD) respiratory rhythms of 8 species of cardinal fishes. Volumetric respirometers, designed specifically for long-term measurements of oxygen uptake by small fishes under controlled conditions, were used during standard 5 day experiments. Despite some individual variation, certain species-specific and genus-specific patterns of respiration emerged. Each species showed a persistant (advancing) rhythm of oxygen consumption during 3 days of DD. Light appeared to be the Zeitgeber, with the light-dark transition phasing the nocturnal peaks and the dark-light transition possibly phasing the diurnal ones. Genus-specific and species-specific trends in field behavior, habitat selection, and distribution were studied. There were correlations between the field behavior and the entrained respiratory rhythms of these nocturnally active fishes. Entrained and endogenous rhythms of each species showed characteristic similarities and were interpreted as integral components of a level of organization evolved for nocturnal life in the shallow marine environment.     

Juvenile–adult associations in sea urchins ( Strongylocentrotus franciscanus and S. droebachiensis ): protection from predation and hydrodynamics in S. franciscanus

Juvenile red sea urchins, Strongylocentrotus franciscanus, aggregate under adult conspecifics, whereas sympatric juvenile green sea urchins, Strongylocentrotus droebachiensis, are typically more solitary and dispersed. Neither the potential advantage of juvenile sheltering nor the differences in post-settlement behavior between the two species has been demonstrated experimentally, but may be related to protection from predators and/or hydrodynamics. In predation experiments, juvenile vulnerability differed in the two species as the seastar Pycnopodia helianthoides consistently chose juvenile S. franciscanus over S. droebachiensis (100% vs. 0%). When associated with adults, juvenile mortality decreased dramatically in S. franciscanus (90% alone vs. 5% with adults), but very little in S. droebachiensis (85% vs. 75%). Not surprisingly, juvenile behavioral responses in the two species reflect this difference in vulnerability. Juvenile S. franciscanus sheltered under adults when predation risk was high, but not when risk was low (44% vs. 13%), whereas sheltering in S. droebachiensis was infrequent and not related to predation risk (7% for high risk versus 5% for low risk). From a hydrodynamic perspective, the presence of an adult led to the creation of a hydrodynamic refuge for juveniles, where average water velocities were reduced by > 60% around the adult urchin. Again, striking differences in sheltering rate were apparent in S. franciscanus (52% vs. 13% for high flow and low flow, respectively), but not S. droebachiensis (5% for high flow versus 4% for low flow). Sheltering behavior was also species-specific as juveniles did not shelter at high rates under adults of the opposite species (≤ 16%). A field survey confirmed these finding in that juvenile S. franciscanus abundance was associated with both adults and water motion (R ² = 0.80, P = 0.008, best-subsets regression). These results suggest that sheltering confers juvenile S. franciscanus with a degree of protection from predators and water motion, and that species-specific differences in this post-settlement behavior may be related to the differences in the protection afforded by adults.     

Pups crying bass: vocal adaptation for avoidance of age-dependent predation risk in ground squirrels?

In most mammals, larger adult body size correlates with lower fundamental frequency and more closely spaced formants in vocalizations relative to juveniles. In alarm whistles of two free-living rodents, the speckled ground squirrel Spermophilus suslicus and yellow ground squirrel S. fulvus, these cues to body size were absent despite prominent differences in body weight and skull and larynx sizes between juveniles and adults. No significant correlations were found between the individual maximum fundamental frequency and body weight, both within age classes and for pooled samples of all animals within species. Furthermore, the mean alarm whistle maximum fundamental frequencies did not differ significantly between age classes (juvenile versus adult) in the speckled squirrel and were even significantly lower in juvenile yellow squirrels. We discuss the hypothesis that the obfuscation of vocal differences between juvenile and adult squirrels may represent a special adaptation of pup vocal behaviour—a form of “vocal mimicry,” resulting in imitation of adult vocal pattern to avoid infanticide and age-dependent predation risk.     

Circadian rhythm of swimming activity in juvenile pink salmon (Oncorhynchus gorbuscha)

The circadian rhythm of swimming activity and the role of the daily illumination cycle in the synchronization of this rhythm were studied in individual juvenile pink salmon. Sixty eight percent of all fish examined (n=38) were day-active when exposed to a 12 h L:12 h D cycle; the remaining fish were nocturnally active. One half of the fish tested under laboratory conditions of continuous, constant light intensity (LL) and constant temperature showed unambiguously endogenous activity rhythms with circadian periods for up to 10 d. The remaining fish were arrhythmic. Mean period length of the free-running activity rhythms for diurnal fish in LL shortened with constant light intensity increasing from 6 to 600 lx, as predicted by the circadian rule. In contrast, mean free-running period for nocturnal fish did not vary significantly with similarly increasing constant light intensity. Mean swimming speed (activity level) of both diurnal and nocturnal fish increased significantly with increasing light intensity. This is suggestive of a positive photokinetic response. When subjected to a phase-delayed LD cycle, the fish resynchronized their daily rhythms of activity with this new LD cycle after only one transient cycle in most instances. Hence, the timing of the daily activity rhythms appeared to occur through the direct masking action of the illumination cycle on activity, rather than through entrainment of an endogenous circadian system.     

Patterns of co-occurrence and interactions between age classes of the common triplefin,Forsterygion lapillum

Many marine organisms have pelagic larval stages that settle into benthic habitats occupied by older individuals; however, a mechanistic understanding of intercohort interactions remains elusive for most species. Patterns of spatial covariation in the densities of juvenile and adult age classes of a small temperate reef fish, the common triplefin (Forsterygion lapillum), were evaluated during the recruitment season (Feb–Mar, 2011) in Wellington, New Zealand (41°17′S, 174°46′E). The relationship between juvenile and adult density among sites was best approximated by a dome-shaped curve, with a negative correlation between densities of juveniles and adults at higher adult densities. The curve shape was temporally variable, but was unaffected by settlement habitat type (algal species). A laboratory experiment using a “multiple-predator effects” design tested the hypothesis that increased settler mortality in the presence of adults (via enhanced predation risk or cannibalism) contributed to the observed negative relationship between juveniles and adults. Settler mortality did not differ between controls and treatments that contained either one (p = 0.08) or two (p = 0.09) adults. However, post hoc analyses revealed a significant positive correlation between the mean length of juveniles used in experimental trials and survival of juveniles in these treatments, suggesting that smaller juveniles may be vulnerable to cannibalism. There was no evidence for risk enhancement or predator interference when adults were present alongside a heterospecific predator (F. varium). These results highlight the complex nature of intercohort relationships in shaping recruitment patterns and add to the growing body of literature recognizing the importance of age class interactions.     

Life history of Gonatus onyx (Cephalopoda: Teuthoidea): ontogenetic changes in habitat, behavior and physiology

In the present study 670 individuals of Gonatus onyx (Cephalopoda: Teuthoidea) were observed in Monterey Bay, California from a remotely operated vehicle. The vertical distribution of this species was bimodal, with peaks at 400 and 800 m depth during the day and 300 and 500 m during the night. The bimodal distribution reflects a life stage shift between younger, schooling juveniles living in shallower water and older, solitary adults which live deeper. Ontogenetic changes in behavior associated with this life stage shift are reflected in the physiology of the organisms as well. Both aerobic and anaerobic metabolism, as estimated using mitochondrial and glycolytic enzymes, decline with increasing body mass, suggesting reduced locomotory capacity in deeper-living adults. Oxygen consumption rates were also determined in relation to oxygen partial pressure. Oxygen consumption regulation was similar between juvenile and adult squids. The critical oxygen partial pressures (29 to 30 mmHg) correspond precisely to the oxygen concentrations found at the depth of maximal abundance for day and night populations of juveniles and adults, respectively. Behavioral and physiological changes with ontogeny of G. onyx are believed to result from reduced visual predator/prey interactions in the light-limited deep sea. Received: 7 February 1999 / Accepted: 26 January 2000     

Circannual variations of lizard circadian activity rhythms in constant darkness

The daily pattern of locomotor activity of Podarcis sicula in the field changes from unimodal in spring to bimodal in summer, becoming unimodal again in autumn. Short-term experiments in which P. sicula collected in different months were tested under constant conditions immediately after capture showed that the activity pattern typical of each season is retained in the lizard circadian locomotor rhythm. In constant conditions, the bimodal pattern is associated with a short free-running period (τ) of the circadian locomotor rhythm and a long circadian activity (α), while the unimodal pattern is associated with a long τ and short α. To test whether seasonal changes in circadian locomotor rhythms are driven by a circannual clock, we recorded locomotor activity of lizards over 12–15 months in constant temperature and darkness. The present results demonstrate, for the first time in a vertebrate, the existence of circannual changes in constant darkness of both τ and α. In most lizards, the longest τ along its circannual cycle is associated with a short α, and the shortest τ in the same cycle with a long α, so that the pattern of mutual association between τ and α is the same as in short-term experiments. Most lizards, however, stayed unimodal all the time. This shows that changes in activity pattern from unimodal to bimodal (and vice versa) are induced by seasonal changes in environmental factors, instead of being incorporated into a circannual rhythm. Circannual changes in τ and α of locomotor rhythms may adaptively predispose the circadian system of P. sicula to a change in activity pattern as soon as seasonal changes in the environment demand it. Received: 22 January 1999 / Received in revised form: 14 April 1999 / Accepted: 19 April 1999     

Growth costs of territorial overlap: experiments with juvenile lizards (Anolis aeneus)

Summary Several theoretical models have suggested that increases in territory overlap with conspecifics should result in lower rates of resource accumulation by territorial residents. This assumption was tested using juvenile Anolis aeneus lizards, which exhibit variable degrees of territory overlap in both the field and laboratory. In a series of laboratory trials under controlled density and resource conditions, juvenile growth rates were negatively related to the degree of territory overlap. This effect aplied to juveniles of all but the lowest status groups, although higher status juveniles had more exclusive territories than did lower status juveniles. Focal studies indicated that food depletion in the zones of overlap was the most likely reason for the negative relationship between territory overlap and growth. It is suggested that juveniles in the field may tolerate extensive territory overlap in spite of growth penalties because of a general shortage of predator-safe homesites in undisturbed habitats.     

Analysis of individual year-classes of a marine fish reveals little evidence of first-generation hybrids between cryptic species in sympatric regions

As settled juveniles and adults, blue rockfish (Sebastes mystinus) are nonmigratory inhabitants of kelp and rocky reef habitats along the California coast, USA, and prior to settlement, they possess a pelagic larval and juvenile stage lasting 3–5 months. A previous study of adults revealed two cryptic species within S. mystinus and evidence of reproductive isolation in a region where both cryptic adults co-occur. Given this pattern of reproductive isolation, we investigated the degree of hybridization or introgression in individual year-classes shortly after juvenile settlement in two different years (2001 and 2002). Using microsatellite markers, we found little indication of hybridization in new juvenile year-classes despite an adult population that comprised both cryptic species. However, we found an average of two percent of hybrid or introgressed individuals in regions with a low frequency of one of the two species. Therefore, while the lack of hybrids or introgression supports the hypothesis of reproductive isolation between the cryptic species within S. mystinus, the age-structured analysis also revealed a spatial pattern of low-frequency differences in the number of introgressed individuals. These results suggest that reproductive barriers may breakdown when one of the two species predominates the regional adult gene pool.     

Intraspecific mimicry and status signals in juvenile African penguins

Summary Observations of resting groups of African penguins Spheniscus demersus on land showed that adults were more aggressive towards juveniles than towards adults. Head coloration was important in triggering this aggression. Adults probably discriminate against juveniles to exclude inexperienced birds from co-operative feeding groups and thus maximize their own energetic returns. There was a disproportionately low frequency of penguin groups at sea containing both adult and juvenile birds. Almost a quarter of juvenile penguins moulted a variable amount of their heads into adult plumage at sea. This reduced the amount of aggression received in proportion to the degree of head moult. Head moult probably occurs in only the fittest juveniles. Birds which moult can join adult feeding groups where communal feeding enhances success. The costs of moulting at sea prevent the occurrence of head moulting in the entire population of juveniles.     

Interactions between sea urchins (Strongylocentrotus droebachiensis) and their predators in field and laboratory experiments

Field observations and manipulative experiments in a nearshore cobble bed (2 to 3 m below mean low water) at Eagle Head, Nova Scotia, Canada, between 1984 and 1986, showed that small juveniles ofStrongylocentrotus droebachiensis (3 to 6 mm diam) sheltering beneath cobbles had a refuge from predators such as rock crabs, small lobsters, and fish. Sea urchins gradually outgrew these refuges and small adults (25 to 30 mm) required larger rocks as shelter from predators, particularly large cancrid crabs. Small juveniles were usually solitary and well dispersed beneath cobbles, whereas small adults tended to aggregate on the undersides and in the interstices of boulders. These aggregations may develop passively as sea urchins accumulate in suitablysized refuges. Chemotaxis experiments indicate that juvenileS. droebachiensis are repelled by waterborne stimuli from conspecifics. In a factorial experiment, effects of the presence of potential predators (rock crabs and lobsters) and/or food (kelp) on the behaviour of large juvenile (10 to 15 mm) and small adult sea urchins were examined in flowing seawater tanks. Both size classes formed exposed feeding aggregations when kelp was provided as food, irrespective of the presence or absence of predators. In the absence of kelp, each size class responded differently to the presence of a predator: juveniles became more cryptic, whereas adults aggregated on the tank sides. Increased movement to the sides of a tank in the presence of a predator may reflect a flight response, since chemotaxis experiments indicated thatS. droebachiensis is repelled by waterborne chemical stimuli from predators. Observational and experimental data suggest that predation at the late juvenile and early adult stages may influence population structure, distribution and abundance ofS. droebachiensis.     

Ontogenetic change in the radula of Conus magus (Gastropoda)

Specimens of the Indo-Pacific piscivorous gastropod Conus magus Linnaeus, 1758 were either collected by SCUBA in the field in Palau (1983–1984) or raised in the laboratory from egg capsules. C. magus captures adult fish with a specially modified radula tooth. Radula morphology was analyzed in a size range of individuals from 4.1 to 43.7 mm shell length, which encompassed early postmetamorphic juveniles to adults. Post-metamorphic C. magus juveniles are too small to consume fish, and all individuals below 9 mm possess a juvenile radula tooth totally different from that of the adult and resembling that of some vermivorous Conus species. The only food remains found in the digestive tracts of juveniles were the setae of syllid polychaetes. All individuals above 10.5 mm possessed adult teeth and had only fish remains in the gut. Two specimens, 10.1 and 9.2 mm in shell length, had intermediate-type radula teeth.