Hierarchical analysis of avian re-nesting behavior: mean, across-individual, and intra-individual responses

© 2015, Springer-Verlag Berlin Heidelberg. Anti-predator behavior is a key aspect of life history evolution, usually studied at the population (mean), or across-individual levels. However individuals can also differ in their intra-individual (residual) variation, but to our knowledge, this has only been studied once before in free-living animals. Here we studied the distances moved and changes in nest height and concealment between successive nesting attempts of marked pairs of grey fantails (Rhipidura albiscapa) in relation to nest fate, across the breeding season. We predicted that females (gender that decides where the nest is placed) should on average show adaptive behavioral responses to the experience of prior predation risk such that after an unsuccessful nesting attempt, replacement nests should be further away, higher from the ground, and more concealed compared with replacement nests after successful nesting attempts. We found that, on average, females moved greater distances to re-nest after unsuccessful nesting attempts (abandoned or depredated) in contrast to after a successful attempt, suggesting that re-nesting decisions are sensitive to risk. We found no consistent across-individual differences in distances moved, heights, or concealment. However, females differed by 53-fold (or more) in their intra-individual variability (i.e., predictability) with respect to distances moved and changes in nest height between nesting attempts, indicating that either some systematic variation went unexplained and/or females have inherently different predictability. Ignoring these individual differences in residual variance in our models obscured the effect of nest fate on re-nesting decisions that were evident at the mean level.

Species-specific characteristics and individual variation of the bark call produced by male Australian fur seals (Arctocephalus pusillus doriferus)

Australian fur seals Arctocephalus pusillus doriferus are colonial breeding animals forming dense social groups during the breeding season. During this time, males establish and defend territories through physical conflicts, stereotyped posturing and vocalisations. While vocalisations are suggested to play an important role in male recognition systems, it has received little attention. Recordings of nine adult male Australian fur seals were made during the 2000 and 2001 breeding seasons at Kanowna Island (39° 10’S, 146° 18' E), Bass Strait, Australia. The in-air bark vocalisations of territory-holding males were used to characterise the Bark Call and to determine whether males produce individually distinct calls, which could be used as a basis for vocal recognition. Seventeen frequency and temporal variables were measured from a total of 162 barks from nine individual males. The Bark Series was more reliably classified (83%) to the correct caller compared to the Bark Unit. This was assigned with less certainty (68%), although the classification was still relatively high. Findings from this study indicate that there is sufficient stereotypy within individual calls, and sufficient variation between them, to enable vocal recognition in male Australian fur seals.

Individual variation in the pup attraction call produced by female Australian fur seals during early lactation

Otariid seals (fur seals and sea lions) are colonial breeders with large numbers of females giving birth on land during a synchronous breeding period. Once pups are born, females alternate between feeding their young ashore and foraging at sea. Upon return, both mother and pup must relocate each other and it is thought to be primarily facilitated by vocal recognition. Vocalizations of thirteen female Australian fur seals (Arctocephalus pusillus doriferus) were recorded during the breeding seasons of December 2000 and 2001, when pups are aged from newborns to one month. The pup attraction call was examined to determine whether females produce individually distinct calls which could be used by pups as a basis for vocal recognition. Potential for individual coding, discriminant function analysis (DFA), and classification and regression tree analysis were used to determine which call features were important in separating individuals. Using the results from all three analyses: F0, MIN F and DUR were considered important in separating individuals. In 76% of cases, the PAC was classified to the correct caller, using DFA, suggesting that there is sufficient stereotypy within individual calls, and sufficient variation between them, to enable vocal recognition by pups of this species.

Host intra-clutch variation, cuckoo egg matching and egg rejection by great reed warblers

Prevailing theory predicts that lower levels of intra-clutch variation in host eggs facilitate the detection of brood parasitism. We assessed egg matching using both human vision and UV-VIS spectrophotometry and then followed the nest fate of great reed warblers naturally parasitised by European cuckoos. Rejection was predicted by the following three variables: matching between cuckoo and host eggs on the main chromatic variable defined by principal components analysis of the egg spectra (which has a strong loading in the UV); the number of host eggs in the nest; and human estimates of intra-clutch variation. The first variable is not correlated to human estimates of matching, which do not predict rejection. In line with another recent study, rejection rates were predicted by higher levels of intra-clutch variation in the host eggs, suggesting that higher rather than lower levels of intra-clutch variation can facilitate the discrimination of cuckoo eggs by hosts. We suggest that the importance of intra-clutch variation is context dependent, with intra-clutch variation being important when there is good matching between the host and the cuckoo eggs. Our results also suggest that both spectrometric and human visual assessments of egg matching and intra-clutch variation are prudent: the former provide the best method of estimating reflectance variation, whereas the latter include some assessment of patterns of maculation.

Individual variation of in-air female ‘pup contact’ calls in Weddell seals, Leptonychotes weddellii

Vocal recognition may function as a critical factor in maintaining the phocid mother-pup bond during lactation. For vocal recognition to function, the caller must produce individually distinct calls that are recognised by their intended recipient. Mother-pup vocal recognition has been studied extensively in colonial otariids and appears to be characteristic of this family. Although less numerous, empirical studies of phocid species have revealed a range of recognition abilities. This study investigated whether Weddell seal (Leptonychotes weddellii) females produce individually distinct 'pup contact' calls that function during natural pair reunions. Fifteen calls from each of nine females recorded in the Vestfold Hills, Antarctica were analysed. One temporal, nine fundamental frequency and five spectral characteristics were measured. Results of the cross-validated Discriminant Function Analysis revealed that mothers produce individually distinct calls with 56% of calls assigned to the correct individual. The probability of achieving this level of discrimination on novel data by chance alone is highly improbable. Analysis of eight mother-pup reunions recorded near McMurdo Sound, Antarctica further demonstrated that these 'pup contact' calls function during natural pair reunions. Behavioural analysis also revealed that pups were chiefly responsible for establishing and maintaining close contact throughout the reunion process. Our study therefore demonstrates that Weddell seal females produce calls with sufficient stereotypy to allow pups to identify them during pair reunions, providing evidence of a functioning mother-pup vocal recognition system.

Stable isotopes reveal inter-annual and inter-individual variation in the diet of female Australian fur seals

Understanding the temporal and spatial variation of foraging habits of apex predators is central to understanding their role in marine ecosystems and how their populations may respond to environmental variability. In the present study, stable isotope analysis (C and N) of blood was used to investigate inter-individual and inter-annual differences in the diet of adult female Australian fur seals Arctocephalus pusillus doriferus. Positive correlations were observed between red cell and plasma values for δ¹³C and δ¹⁵N (r² = 0.47 and r² = 0.66, respectively, p < 0.001 in both cases), suggesting relatively consistent individual prey choices over 3 or 4 foraging trips. Mean δ¹⁵N values (12.8 to 17.5%) confirm the species occupies the highest marine trophic niche in the region. A significant decrease in plasma δ¹⁵N values, corresponding to two-thirds of a trophic level (ca. 2%), was observed between the 1998 to 2000 and 2003 to 2005 sampling periods. This was associated with a significant decrease in adult female body condition and is consistent with a decline, previously documented by faecal analysis, of the proportion of red cod Pseudophysis bachus, barracouta Thyrsites atun and Gould's squid Nototodarus gouldi in the diet and an increase in redbait Emmelichthys nitidus. While substantial variation in δ¹⁵N was observed within each age cohort, a significant decrease was observed with age, suggesting individual specialisation for particular prey types is evident early in adulthood, but that its composition changes as females age. In addition, generalized linear models indicated body mass had a negative influence on δ¹⁵N, which may reflect larger total body oxygen stores, facilitating individuals hunting cryptic prey of lower trophic level (e.g. octopus) on the sea floor.

Host behaviour and physiology underpin individual variation in avian influenza virus infection in migratory Bewick's swans

Individual variation in infection modulates both the dynamics of pathogens and their impact on host populations. It is therefore crucial to identify differential patterns of infection and understand the mechanisms responsible. Yet our understanding of infection heterogeneity in wildlife is limited, even for important zoonotic host-pathogen systems, owing to the intractability of host status prior to infection. Using novel applications of stable isotope ecology and eco-immunology, we distinguish antecedent behavioural and physiological traits associated with avian influenza virus (AIV) infection in free-living Bewick's swans (Cygnus columbianus bewickii). Swans infected with AIV exhibited higher serum δ13C (-25.3 ± 0.4) than their non-infected counterparts (-26.3±0.2). Thus, individuals preferentially foraging in aquatic rather than terrestrial habitats experienced a higher risk of infection, suggesting that the abiotic requirements of AIV give rise to heterogeneity in pathogen exposure. Juveniles were more likely to be infected (30.8% compared with 11.3% for adults), shed approximately 15-fold higher quantity of virus and exhibited a lower specific immune response than adults. Together, these results demonstrate the potential for heterogeneity in infection to have a profound influence on the dynamics of pathogens, with concomitant impacts on host habitat selection and fitness.

Individual variation in foraging movements in a lake population of young-of-the-year brook charr (Salvelinus fontinalis)

We examined the variation in foraging movements in a population of young-of-the-year (YOY) brook charr living in the near-shore littoral zone of a lake. By repeating the methodology of an earlier stream study, we made direct comparisons between data from lake and stream populations. In general, the pattern of variation in foraging movements was similar between the two sites with greater variability and activity observed in the lake population. The dichotomous nature of the proportion of time spent moving in the stream was also observed in the lake population but in a reversal of the stream pattern. Charr that moved constantly while foraging represented the largest movement category in the lake. In general, variation in foraging movements were more strongly related to the rate of prey rejection, whereas environmental factors, such as distance from shore, submerged objects, and the amount of overhead riparian cover, were more strongly related to prey ingestion. This last finding directly contrasts with that found in the stream literature for YOY charr in still water where ingestion rate, as estimated using feeding attempt rate, increases with the mobility of YOY charr.

Individual variation in thermal performance curves: swimming burst speed and jumping endurance in wild-caught tropical clawed frogs

The importance of studying individual variation in locomotor performance has long been recognized as it may determine the ability of an organism to escape from predators, catch prey or disperse. In ectotherms, locomotor performance is highly influenced by ambient temperature (Ta), yet several studies have showed that individual differences are usually retained across a Ta gradient. Less is known, however, about individual differences in thermal sensitivity of performance, despite the fact that it could represent adaptive sources of phenotypic variation and/or additional substrate for selection to act upon. We quantified swimming and jumping performance in 18 wild-caught tropical clawed frogs (Xenopus tropicalis) across a Ta gradient. Maximum swimming velocity and acceleration were not repeatable and individuals did not differ in how their swimming performance varied across Ta. By contrast, time and distance jumped until exhaustion were repeatable across the Ta gradient, indicating that individuals that perform best at a given Ta also perform best at another Ta. Moreover, thermal sensitivity of jumping endurance significantly differed among individuals, with individuals of high performance at low Ta displaying the highest sensitivity to Ta. Individual differences in terrestrial performance increased with decreasing Ta, which is opposite to results obtained in lizards at the inter-specific and among-individual levels. To verify the generality of these patterns, we need more studies on individual variation in thermal reaction norms for locomotor performance in lizards and frogs.

Variation in energy intake and basal metabolic rate of a bird migrating in a wind tunnel

1. We studied the changes in body mass, metabolizable energy intake rate (ME) and basal metabolic rate (BMR) of a Thrush Nightingale, Luscinia luscinia, following repeated 12-h migratory flights in a wind tunnel. In total the bird flew for 176 h corresponding to 6300 km. This is the first study where the fuelling phase has been investigated in a bird migrating in captivity.

2. ME was very high, supporting earlier findings that migrating birds have among the highest intake rates known among homeotherms. ME was significantly higher the second day of fuelling, indicating a build-up of the capacity of the digestive tract during the first day of fuelling.

3. Further indications of an increase in size or activity level of metabolically active structures during fuelling come from the short-term variation in BMR, which increased over the 2-day fuelling period with more than 20%, and in almost direct proportion to body mass. However, mass-specific BMR decreased over the season.

4. The patterns of mass change, ME and BMR of our focal bird following two occasions of 12-h fasts were the same as after flights, indicating that fast and flight may involve similar physiological processes.

5. The relatively low ME the first day following a flight may be a contributing factor to the well-known pattern that migrating birds during stopover normally lose mass the first day of fuelling.

Sexual niche segregation and gender-specific individual specialisation in a highly dimorphic marine mammal

While sexual segregation is expected in highly dimorphic species, the local environment is a major factor driving the degree of resource partitioning within a population. Sexual and individual niche segregation was investigated in the Australian fur seal (Arctocephalus pusillus doriferus), which is a benthic foraging species restricted to the shallow continental shelf region of south-eastern Australia. Tracking data and the isotopic values of plasma, red blood cells and whiskers were combined to document spatial and dietary niche segregation throughout the year. Tracking data indicated that, in winter, males and females overlapped in their foraging habitat. All individuals stayed within central Bass Strait, relatively close (< 220 km) to the breeding colony. Accordingly, both genders exhibited similar plasma and red cell δ13C values. However, males exhibited greater δ13C intra-individual variation along the length of their whisker than females. This suggests that males exploited a greater diversity of foraging habitats throughout the year than their female counterparts, which are restricted in their foraging grounds by the need to regularly return to the breeding colony to suckle their pup. The degree of dietary sexual segregation was also surprisingly low, both sexes exhibiting a great overlap in their δ15N values. Yet, males displayed higher δ15N values than females, suggesting they fed upon a higher proportion of higher trophic level prey. Given that males and females exploit different resources (mainly foraging habitats), the degree of individual specialisation might differ between the sexes. Higher degrees of individual specialisation would be expected in males which exploit a greater range of resources. However, comparable levels of inter-individual variation in δ15N whisker values were found in the sampled males and females, and, surprisingly, all males exhibited similar seasonal and inter-annual variation in their δ13C whisker values, suggesting they all followed the same general dispersion pattern throughout the year.

HbA1c as a screening tool for detection of type 2 diabetes: A systematic review

Aim: To assess the validity of glycated haemoglobin A1c (HbA1c) as a screening tool for early detection of Type 2 diabetes.

Methods: Systematic review of primary cross-sectional studies of the accuracy of HbA1c for the detection of Type 2 diabetes using the oral glucose tolerance test as the reference standard and fasting plasma glucose as a comparison.

Results: Nine studies met the inclusion criteria. At certain cut-off points, HbA1c has slightly lower sensitivity than fasting plasma glucose (FPG) in detecting diabetes, but slightly higher specificity. For HbA1c at a Diabetes Control and Complications Trial and UK Prospective Diabetes Study comparable cut-off point of ≥ 6.1%, the sensitivity ranged from 78 to 81% and specificity 79 to 84%. For FPG at a cut-off point of ≥ 6.1 mmol/l, the sensitivity ranged from 48 to 64% and specificity from 94 to 98%. Both HbA1c and FPG have low sensitivity for the detection of impaired glucose tolerance (around 50%).

Conclusions: HbA1c and FPG are equally effective screening tools for the detection of Type 2 diabetes. The HbA1c cut-off point of > 6.1% was the recommended optimum cut-off point for HbA1c in most reviewed studies; however, there is an argument for population-specific cut-off points as optimum cut-offs vary by ethnic group, age, gender and population prevalence of diabetes. Previous studies have demonstrated that HbA1c has less intra-individual variation and better predicts both micro- and macrovascular complications. Although the current cost of HbA1c is higher than FPG, the additional benefits in predicting costly preventable clinical complications may make this a cost-effective choice.

Criterion validity of the activPAL™ and ActiGraph for assessing children's sitting and standing time in a school classroom setting

BACKGROUND: Few studies have investigated the accuracy of the ActiGraph (AG) GTX3 accelerometer for assessing children's sitting and standing time. The activPAL (aP) has an inclinometer function that enables it to distinguish between sitting/lying and standing; however, its accuracy for assessing sitting and standing in older children is unknown. This study validated the accuracy of these devices for estimating sitting and standing time in a school classroom against a criterion measure of direct observation (DO).

FINDINGS: Forty children in grades 5-7 wore both devices while being video recorded during two school lessons. AG and aP data were simultaneously collected in 15-s epochs. Individual participant DO and aP data were recorded as total time spent sitting/lying, standing and stepping. AG data were converted into time spent sitting and standing using previously established cut-points. Compared with DO, the aP underestimated sitting time (mean bias = -1.9 min, 95 % LoA = -8.9 to 5.2 min) and overestimated standing time (mean bias = 1.8 min, 95% LoA = -9.6 to 13.3 min). The best-performing AG cut-point across both sitting and standing (<75 counts/15 s) was more accurate than the aP, underestimating sitting time (mean bias = -0.8 min, 95 % LoA = -10.5 to 9.9 min) and standing time (mean bias = -0.4 min, 95% LoA = -9.8 to 9.1 min), but was less precise as evidenced by wider LoAs and poorer correlations with DO (sitting r = 0.86 aP vs 0.80 AG; standing r = 0.78 aP vs 0.60 AG).

CONCLUSIONS: The aP demonstrated good accuracy and precision for assessing free-living sitting and standing time in classroom settings. The AG was most accurate using a cut-point of < 75 counts/15 s. Further studies should validate the monitors in settings with greater inter- and intra-individual variation in movement patterns.