First-Year Common Nightingales (Luscinia megarhynchos) Have Smaller Song-Type Repertoire Sizes Than Older Males

Based on the assumptions that birdsong indicates male quality and that quality is related to age, one might expect older birds to signal their age. That is, in addition to actual body condition, at least some song features should vary with age, presumably towards more complexity. We investigated this issue by comparing repertoire sizes of free‐ranging common nightingale males in their first breeding season with those of older males. Nightingales are a good model species as they are open‐ended learners, where song acquisition is not confined to an early sensitive period of learning. Moreover, nightingales develop an extraordinarily large song‐type repertoire (approx. 180 different song types per male), and differences in repertoire size among males are pronounced. We analysed repertoire characteristics of the nocturnal song of nine nightingales in their first breeding season and compared them with the songs of nine older males. The repertoire size of older males was on average 53% larger than that of yearlings. When analysing two song categories of nightingales, whistle and non‐whistle songs separately, we found similar results. Our findings show marked differences in repertoire size between age categories, suggesting that this song feature may reflect a male's age. We discuss those mechanisms that may constrain the development of larger repertoires in first‐year males. Whether repertoire sizes are crucial for female mate choice or in vocal interactions among conspecific males remains open to further investigations.     

Evolving communicative complexity: insights from rodents and beyond

Social living goes hand in hand with communication, but the details of this relationship are rarely simple. Complex communication may be described by attributes as diverse as a species' entire repertoire, signallers' individualistic signatures, or complex acoustic phenomena within single calls. Similarly, attributes of social complexity are diverse and may include group size, social role diversity, or networks of interactions and relationships. How these different attributes of social and communicative complexity co-evolve is an active question in behavioural ecology. Sciurid rodents (ground squirrels, prairie dogs and marmots) provide an excellent model system for studying these questions. Sciurid studies have found that demographic role complexity predicts alarm call repertoire size, while social group size predicts alarm call individuality. Along with other taxa, sciurids reveal an important insight: different attributes of sociality are linked to different attributes of communication. By breaking social and communicative complexity down to different attributes, focused studies can better untangle the underlying evolutionary relationships and move us closer to a comprehensive theory of how sociality and communication evolve.     

Dispersal Ability Determines the Role of Environmental,Spatial and Temporal Drivers of Metacommunity Structure

Recently, community ecologists are focusing on the relative importance of local environmental factors and proxies to dispersal limitation to explain spatial variation in community structure. Albeit less explored, temporal processes may also be important in explaining species composition variation in metacommunities occupying dynamic systems. We aimed to evaluate the relative role of environmental, spatial and temporal variables on the metacommunity structure of different organism groups in the Upper Paraná River floodplain (Brazil). We used data on macrophytes, fish, benthic macroinvertebrates, zooplankton, periphyton, and phytoplankton collected in up to 36 habitats during a total of eight sampling campaigns over two years. According to variation partitioning results, the importance of predictors varied among biological groups. Spatial predictors were particularly important for organisms with comparatively lower dispersal ability, such as aquatic macrophytes and fish. On the other hand, environmental predictors were particularly important for organisms with high dispersal ability, such as microalgae, indicating the importance of species sorting processes in shaping the community structure of these organisms. The importance of watercourse distances increased when spatial variables were the main predictors of metacommunity structure. The contribution of temporal predictors was low. Our results emphasize the strength of a trait-based analysis and of better defining spatial variables. More importantly, they supported the view that “all-or- nothing” interpretations on the mechanisms structuring metacommunities are rather the exception than the rule.     

Does sociality drive the evolution of communicative complexity? A comparative test with ground-dwelling sciurid alarm calls

While sociality has been hypothesized to drive the evolution of communicative complexity, the relationship remains to be formally tested. We derive a continuous measure of social complexity from demographic data and use this variable to explain variation in alarm repertoire size in ground-dwelling sciurid rodents (marmots, Marmota spp.; prairie dogs, Cynomys spp.; and ground squirrels, Spermophilus spp.). About 40% of the variation in alarm call repertoire size was explained by social complexity in the raw data set. To determine the degree to which this relationship may have been influenced by historical relationships between species, we used five different phylogenetic hypotheses to calculate phylogenetically independent contrasts. Less variation was significantly explained in contrast-based analyses, but a general positive relationship remained. Social complexity explained more variation in alarm call repertoire size in marmots, while sociality explained no variation in repertoire size in prairie dogs and no variation in phylogenetically based analyses of squirrels. In most cases, substantial variation remained unexplained by social complexity. We acknowledge that factors other than social complexity, per se, may contribute to the evolution of alarm call repertoire size in sciurid rodents, and we discuss alternative hypotheses. Our measure of social complexity could be used by other researchers to test explicit evolutionary hypotheses that involve social complexity.     

Regulatory T-cell differentiation versus clonal deletion of autoreactive thymocytes

The concept of clonal deletion of immune cells that carry an autoreactive antigen receptor was a central prediction of Burnet's clonal selection theory. A series of classical experiments in the late 1980s revealed that certain immature thymocytes upon encounter of 'self' are indeed removed from the T-cell repertoire before their release into the blood circulation. A second essential cornerstone of immunological tolerance, not anticipated by Burnett, has more recently surfaced through the discovery of Foxp3(+) regulatory T cells (Treg). Intriguingly, it appears that the expression of an autoreactive T-cell receptor is a shared characteristic of T cells that are subject to clonal deletion as well as of those deviated into the Treg lineage. This is all the more striking as Treg differentiation for the most part branches off from mainstream CD4T cell development during thymocyte maturation in the thymus, that is, it may neither temporally nor spatially be separated from clonal deletion. This raises the question of how an apparently identical stimulus, namely the encounter of 'self' during thymocyte development, can elicit fundamentally different outcomes such as apoptotic cell death on the one hand or differentiation into a highly specialized T-cell lineage on the other hand. Here, we will review the T-cell intrinsic and extrinsic factors that have been implicated in intrathymic Treg differentiation and discuss how these parameters may determine whether an autoreactive major histocompatibility complex class II-restricted thymocyte is deviated into the Treg lineage or subject to clonal deletion.     

Early experience shapes the development of behavioral repertoires of hatchling snakes

Snakes are obligate predatory organisms that consume prey whole, and despite their precocious nature, snakes must develop effective feeding skills, especially when encountering large prey. I conducted two experiments that document the development of behavioral repertoires for naïve hatchling trinket snakes, Coleognathus helena. In the first experiment, I examined how experience with prey of different relative prey mass encountered at regular feeding intervals affects hatchling feeding response. I also examined whether hatchling feeding performance improved over time. Improvement was evaluated on the frequency of the most effective behavioral states such as complex prey restraint behaviors and anterior-first ingestion. In the second experiment I tested whether feeding experience with prey of a particular size influences the way hatchlings respond to a novel prey size. All hatchlings improved their predatory behavior when prey size and number of trials were controlled. Hatchlings feeding on larger prey, however, showed greater overall improvement in their feeding behavior and were quick to integrate complex prey restraint behaviors such as constriction into their feeding repertoire. Despite the fact that early experience with one prey size seemed to shape their restraint repertoire during their first four feeding events, hatchlings remained flexible and responded to prey of a novel size with size-specific prey restraint behaviors.     

Elevating perceived predation risk modifies the relationship between parental effort and song complexity in the song sparrow Melospiza melodia

Adult‐directed predation risk elevates costs of parental care, and may modify relationships between sexually selected ornaments and parental effort by accentuating the tradeoff between survival and parental investment. We assessed multiple hypotheses regarding the relationship between maternal effort, paternal effort, and the sexually selected trait of male song complexity in the song sparrow Melospiza melodia. Further, we explored whether experimentally elevating perceived adult‐directed predation risk near nests affected these relationships. We quantified two dimensions of song complexity: song repertoire size and residual syllable number (the relative number of syllables for a given song repertoire size). Under elevated perceived predation risk, but not in the absence of the predator stimuli, females mated to males with higher residual syllable number displayed higher nestling provisioning rates and performed a greater proportion of nestling provisioning trips. In other words, elevating perceived predation risk induced a pattern of maternal investment consistent with differential allocation. In contrast, under elevated perceived predation risk, only, females performed a lesser proportion of provisioning trips when mated to males with large song repertoire sizes. Further, consistent with the good parent hypothesis, males with large song repertoire sizes displayed lower latencies to return to the nest, independent of the predator stimuli. Results suggest that residual syllable number may reflect some aspect of male genetic quality, such that females are more willing to maintain maternal effort while facing heightened predation risk. On the other hand, females may gain paternal benefits when mated to males with large song repertoires. Our study supports the hypothesis that increased costs of parental care associated with predation risk may induce relationships between sexually selected traits and parental behavior, which may increase the strength of sexual selection. Additionally, results suggest that different aspects of song complexity may fulfill non‐equivalent signaling roles.     

Syllable sharing and changes in syllable repertoire size and composition within and between years in the great reed warbler, Acrocephalus arundinaceus

The syllable repertoire size of the great reed warbler (GRW) is known to underlay female preferences. However, its stability over a season—a key factor of this quality-related feature—has never been tested. Similarly, syllable repertoire composition, syllable sharing and its function in GRW have received little attention so far. GRW syllables fit into two categories: whistles and rattles. They differ in structure but nothing is known about their function. In the present study we analyze the above aspects of GRW vocalization. We found that syllable repertoire size and composition were stable within a season but significantly changed between years. Longitudinal analyses revealed that the changes in repertoire size were influenced more by the season than the age of individuals. The repertoire size correlated with harem size. Changes in repertoire size were significantly smaller than changes in composition, indicating varying seasonal usage of syllables present in the repertoires of individual birds. Syllable sharing was high and indifferent between neighbors and distant males. This pattern suggests that syllable sharing in GRW acts as a dialect. High levels of within- and between-season repertoire similarity among all recorded birds supported this view. Whistles were more shared and repeatable within and between seasons than rattles. We suppose that whistles may play an important role in determining a local dialect due to their lower changeability both between individuals and years. Within-season similarity of whistle repertoire to other males correlated with harem size, but similarity of rattle repertoire did not. This correlation may result from a female preference for philopatric males using the local dialect.     

Are there age-related differences in the song repertoire size of Eurasian blackbirds?

Most oscine bird species possess a repertoire of different song patterns, and repertoire size is thought to signal aspects of male quality. As age is assumed to be related to male quality in terms of experience and/or viability, repertoire size may be expected to reflect male age. Here, we investigated the relationship between repertoire size and age (yearlings or older) in Eurasian blackbirds, Turdus merula, a species with a large repertoire delivered in a highly variable manner. We found that older males tended to have larger repertoires than yearlings though the two age groups overlapped considerably. Thus, compared to other species with large repertoires, age-related differences in repertoire size seem rather small in male Eurasian blackbirds. We also compared repertoires of three individuals in two successive years (as yearlings and in the year following) and found a large element turnover. Our investigation revealed that this turnover was almost complete in the quiet terminating twitter part of the song. Such turnover may allow a young bird to adjust his repertoire to his neighbours?? repertoires, which could be useful for song matching interactions.     

Antibody-antigen complex formation with immobilized immunoglobulins.

We have investigated the complex formation between an immobilized monoclonal antibody and antigens that differ in size about 50-fold. As a model system, we used an iodinated progesterone derivative and a progesterone-horseradish peroxidase conjugate as tracer and a monoclonal antibody as binding protein. The antibody was immobilized by four different methods: physical adsorption, chemical binding, and binding via protein G in the absence or presence of a protective protein (gelatin). These investigations have shown that the performance of competitive immunoassays is determined by a combination of factors: (a) the relative size of the analyte and the tracer, (b) the antibody density on the solid matrix, (c) the method of immobilization of the antibody, and (d) the binding constants between antibody-analyte and antibody-tracer. All of these interactions have to be considered in designing an optimal immunoassay. The smaller antigen can form a 3- to 35-fold higher maximal complex density than the larger antigen. Dose-response curves are less affected by the size of the tracer than by the binding constant with the antibody. A large enzyme tracer with a relatively low binding constant can, therefore, provide a more sensitive assay. On the other hand, the increase in complex density achieved with a smaller tracer yields a higher signal that in turn can provide a better signal-to-noise ratio in highly sensitive competitive solid-phase immunoassays. We have suggested a model for antibody immobilization that accounts for the interdependence of tracer size, complex formation, and antibody density. The methods described can be used to design and optimize immunoassays of predefined performance characteristics. The results are particularly useful for converting radioimmunoassays to enzyme immunoassays.     

Song, sexual selection, and a song control nucleus (HVc) in the brains of European sedge warblers

Female sedge warblers select males that have more complex songs as mates. This study tests two predictions concerning HVc, a telencephalic nucleus that is essential for song learning and production: first, that males with more complex songs will have a larger HVc, and second that males who pair successfully will have a larger HVc than unpaired males. Data on song composition and pairing status were collected from wild sedge warblers breeding in Hungary. We found significant positive correlations between three song attributes (repertoire size, song complexity, and song length) and the size of HVc. Males that paired successfully also had more complex songs (repertoire size and song complexity, though not song length) than males that did not. However, we find no direct evidence that males who paired successfully had a larger HVc than unpaired males. These findings are discussed in relation to the possible functions of HVc and also to current views on sexual selection and the evolution of the song control pathway.     

中国瓢虫体型的地理分异及其与环境因子的关系

生物体的外在形态(个体大小及异速生长)不仅是分类学的重要依据, 也是影响生物与环境相互作用的重要功能性状。昆虫体型的地理分异及其机制是昆虫生物地理学的重要基础。本文基于中国省级行政区的瓢虫体型、分布和环境因子数据, 运用相关和回归分析, 探索了中国瓢虫体型(体长、体宽与长宽比)的纬度地带性及其与环境因子的关系。为了确定营养级对瓢虫体型特征的影响, 我们对全部瓢虫、植食性瓢虫和捕食性瓢虫分别进行分析。研究结果表明: (1)瓢虫的体型大小具有显著的纬度地带性, 即纬度越高, 瓢虫的体长或体宽越大, 符合贝格曼法则。与温度相关的环境因子是导致瓢虫体型大小地理分异的主要环境因素, 因为瓢虫以成虫越冬, 体型越大则能量储存越多, 冬眠期间的耐饥力越强; (2)由于食性和食物空间分布的差异, 在各个纬度上, 植食性瓢虫体型总是大于捕食性瓢虫; 捕食性瓢虫的长宽比与纬度呈显著正相关, 即纬度越高, 瓢虫体型越狭长; 而植食性瓢虫的长宽比与纬度没有显著的相关性; 捕食性瓢虫的长宽比与年降水量呈显著负相关, 因为年降水量降低导致植被的斑块化增加, 引起捕食对象分布更为分散, 从而要求瓢虫具有更强的飞行能力, 即更大的长宽比。综上所述, 在省级尺度上, 温度和降水分别主要影响瓢虫的大小和长宽比, 其效应随营养级(食性)的变化而变化。     

A twelve-step program for evolving multicellularity and a division of labor

The volvocine algae provide an unrivalled opportunity to explore details of an evolutionary pathway leading from a unicellular ancestor to multicellular organisms with a division of labor between different cell types. Members of this monophyletic group of green flagellates range in complexity from unicellular Chlamydomonas through a series of extant organisms of intermediate size and complexity to Volvox, a genus of spherical organisms that have thousands of cells and a germ-soma division of labor. It is estimated that these organisms all shared a common ancestor about 50 +/- 20 MYA. Here we outline twelve important ways in which the developmental repertoire of an ancestral unicell similar to modern C. reinhardtii was modified to produce first a small colonial organism like Gonium that was capable of swimming directionally, then a sequence of larger organisms (such as Pandorina, Eudorina and Pleodorina) in which there was an increasing tendency to differentiate two cell types, and eventually Volvox carteri with its complete germ-soma division of labor.     

Ageing of the B-cell repertoire

Older people are more susceptible to infection, less responsive to vaccination and have a more inflammatory immune environment. Using spectratype analysis, we have previously shown that the B-cell repertoire of older people shows evidence of inappropriate clonal expansions in the absence of challenge, and that this loss of B-cell diversity correlates with poor health. Studies on response to vaccination, using both spectratyping and high-throughput sequencing of the repertoire, indicate that older responses to challenge are lacking in magnitude and/or delayed significantly. Also that some of the biologically significant differences may be in different classes of antibody. We have also previously shown that normal young B-cell repertoires can vary between different phenotypic subsets of B cells. In this paper, we present an analysis of immunoglobulin repertoire in different subclasses of antibody in five different populations of B cell, and show how the repertoire in these different groups changes with age. Although some age-related repertoire differences occur in naive cells, before exogenous antigen exposure, we see indications that there is a general dysregulation of the selective forces that shape memory B-cell populations in older people.     

Developmental stress, social rank and song complexity in the European starling (Sturnus vulgaris)

Bird song is a sexually selected trait and females have been shown to prefer males that sing more complex songs. However, for repertoire size to be an honest signal of male quality it must be associated with some form of cost. This experiment investigates the effects of food restriction and social status during development on song complexity in the European starling (Sturnus vulgaris). Birds that experienced an unpredictable food supply early in life produced a significantly smaller repertoire of song phrases than those with a constant food supply. Social status during development was also significantly correlated with repertoire size, with dominant birds producing more phrase types. This study therefore provides novel evidence that social as well as nutritional history may be important in shaping the song signal in this species.     

Phylogenetic comparative methods and the geographic range size – body size relationship in new world terrestrial carnivora

Most recent papers avoid describing macroecological relationships and interpreting then without a previous control of non-independence in data caused by phylogenetic patterns in data. In this paper, we analyzed the geographic range size – body size relationship for 70 species of New World terrestrial Carnivora (fissipeds) using various phylogenetic comparative methods and simulation procedures to assess their statistical performance. Autocorrelation analyses suggested a strong phylogenetic pattern for body size, but not for geographic range size. The correlation between the two traits was estimated using standard Pearson correlation across species (TIPS) and four different comparative methods: Felsenstein's independent contrasts (PIC), autoregressive method (ARM), phylogenetic eigenvector regression (PVR) and phylogenetic generalized least-squares (PGLS). The correlation between the two variables was significant for all methods, except PIC, in such a way that ecological mechanisms (i.e., minimum viable population or environmental heterogeneity- physiological homeostasis), could be valid explanations for the relationship. Simulations using different O-U processes for each trait were run in order to estimate true Type I errors of each method. Type I errors at 5% were similar for all phylogenetic methods (always lower than 8%), but equal to 13.1% for TIPS. PIC usually performs better than all other methods under Brownian motion evolution, but not in this case using a more complex combination of evolutionary models. So, recent claims that using independent contrasts in ecological research can be too conservative are correct but, on the other hand, using simple across-species correlation is too liberal even under the more complex evolutionary models exhibited by the traits analyzed here.     

Element repertoire: change and development with age in Whitethroat Sylvia communis song

Song repertoires are often important determining factors in sexual selection. In several species, older males have larger repertoires than 1-year-old males. The development of large song repertoires by an individual is, however, poorly understood. We studied song element repertoire changes in five individual male Whitethroats Sylvia communis sampled as 1- and 2-year olds. These males increased the size of their element repertoire between their first and second year, but song length and number of different elements per song did not change. On average, 44.3% of the song elements in the first-year repertoire were also found in the second-year repertoire. Elements shared between years were found earlier in the songs and tended to occur in sequences. Sequences of shared elements also seemed to be conserved between years. The study suggests that the song element repertoire of the second year is partly based on the first-year repertoire, which may explain why large song repertoires are mainly expressed by males at least 2 years of age. It would appear, therefore, that song element repertoire size could be a reliable signal of male age.     

Difficulties when assessing birdsong learning programmes under field conditions: a re-evaluation of song repertoire flexibility in the great tit

There is a remarkable diversity of song-learning strategies in songbirds. Establishing whether a species is closed- or open-ended is important to be able to interpret functional and evolutionary consequences of variation in repertoire size. Most of our knowledge regarding the timing of vocal learning is based on laboratory studies, despite the fact that these may not always replicate the complex ecological and social interactions experienced by birds in the wild. Given that field studies cannot provide the experimental control of laboratory studies, it may not be surprising that species such as the great tit that were initially assumed to be closed-ended learners have later been suggested to be open-ended learners. By using an established colour-ringed population, by following a standardized recording protocol, and by taking into account the species' song ecology (using only recordings obtained during peak of singing at dawn), we replicated two previous studies to assess song repertoire learning and flexibility in adult wild great tits elicited by social interactions. First, we performed a playback experiment to test repertoire plasticity elicited by novel versus own songs. Additionally, in a longitudinal study, we followed 30 males in two consecutive years and analysed whether new neighbours influenced any change in the repertoire. Contrary to the previous studies, song repertoire size and composition were found to be highly repeatable both between years and after confrontation with a novel song. Our results suggest that great tits are closed-ended learners and that their song repertoire probably does not change during adulthood. Methodological differences that may have led to an underestimation of the repertoires or population differences may explain the discrepancy in results with previous studies. We argue that a rigorous and standardized assessment of the repertoire is essential when studying age- or playback-induced changes in repertoire size and composition under field conditions.