Individual Reproductive Success Research Articles

Biology in the 21st Century: Natural Selection is Cognitive Selection

Natural selection has a formal definition as the natural process that results in the survival and reproductive success of individuals or groups best adjusted to their environment, leading to the perpetuation of those genetic qualities best suited to that organism's environmental niche. Within conventional Neo-Darwinism, the largest source of those variations that can be selected is presumed to be secondary to random genetic mutations. As these arise, natural selection sustains adaptive traits in the context of a 'struggle for existence'. Consequently, in the 20th century, natural selection was generally portrayed as the primary evolutionary driver.. The 21st century offers a comprehensive alternative to Neo-Darwinian dogma within Cognition-Based Evolution. The substantial differences between these respective evolutionary frameworks have been most recently articulated in a revision of Crick's Central Dogma, a former centerpiece of Neo-Darwinism. The argument is now advanced that the concept of natural selection should also be comprehensively reappraised. Cognitive selection is presented as a more precise term better suited to 21st century biology. Since cognition began with life's origin, natural selection represents cognitive selection.

Correlation of personality with individual reproductive success in shrub-nesting birds depends on their life history style

Two questions in the research of animal personality—whether there is a correlation between a personality trait and individual reproductive success, and what is the genetic basis underlying a personality trait—remain unresolved. We addressed these two questions in three shrub-nesting birds, the Azure-winged Magpie (Cyanopica cyanus, AM), White-collared Blackbird (Turdus albocinctus, WB), and Brown-cheeked Laughingthrush (Trochalopteron henrici, BL). The personality type of an individual was first identified according to its response to a territorial intruder. Then, we compared the fleeing distance, breeding parameters, and differential expressed genes (DEGs) in the brain transcriptome between bold and shy breeders. In the three species, bold breeders exhibited more aggressiveness towards an intruder of their territory than did shy breeders. The reproductive success of bold breeders was significantly higher than that of shy breeders in AM but not in WB and BL. The three species shared one DEG, crabp1, which was up-regulated in bold relative to in shy individuals. By regulating the expression of corticotropin-releasing hormone, higher crabp1 gene expression can decrease cellular response to retinoic acid. Therefore, bold individuals are insensitive to external stresses and able to exhibit more aggressiveness to intruders than their shier counterparts. Aggressiveness is beneficial to bold individuals in AM but not in WB and BL because the former could evoke neighbors to make the same response of defending against intruders but the latter could not. Although a personality trait may have the same genetic basis across species, its correlation with reproductive success depends largely on the life history style of a species.

Personality traits differentially affect components of reproductive success in a Neotropical poison frog.

Individual reproductive success has several components, including the acquisition of mating partners, offspring production, and offspring survival until adulthood. While the effects of certain personality traits-such as boldness or aggressiveness-on single components of reproductive success are well studied, we know little about the composite and multifaceted effects behavioural traits can have on all the aspects of reproductive success. Behavioural traits positively linked to one component of reproductive success might not be beneficial for other components, and these effects may differ between sexes. We investigated the influence of boldness, aggressiveness, and exploration on the number of mating partners, mating events, and offspring surviving until adulthood in males and females of the Neotropical poison frog Allobates femoralis. Behavioural traits had different-even opposite-effects on distinct components of reproductive success in both males and females. For example, males who displayed high levels of aggressiveness and exploration (or low levels of aggressiveness and exploration) managed to attract high number of mating partners, while males with low levels of boldness, low levels of aggressiveness, and high levels of exploration had the most offspring surviving until adulthood. Our results therefore suggest correlational selection favouring particular combinations of behavioural traits.

Odor cues rather than personality affect tadpole deposition in a neotropical poison frog

Abstract Animals constantly need to evaluate available external and internal information to make appropriate decisions. Identifying, assessing, and acting on relevant cues in contexts such as mate choice, intra-sexual competition, and parental care is particularly important for optimizing individual reproductive success. Several factors can influence decision-making, such as external environmental cues and the animal’s own internal state, yet, we have limited knowledge on how animals integrate available information. Here, we used an entire island population (57 males, 53 females, and 1,109 tadpoles) of the neotropical brilliant-thighed poison frog Allobates femoralis to investigate how 2 factors (olfactory cues and personality traits) influence the ability of males to find and use new resources for tadpole deposition. We experimentally manipulated the location of tadpole deposition sites and their associated olfactory cues, and repeatedly measured exploration and boldness in adult males. We further reconstructed tadpole deposition choices via inferred parent–offspring relationships of adult frogs and tadpoles deposited in our experimental pools using molecular parentage analysis. We found that the discovery and use of new rearing sites were heavily influenced by olfactory cues; however, we did not find an effect of the measured behavioral traits on resource discovery and use. We conclude that in highly dynamic environments such as tropical rainforests, reliable external cues likely take priority over personality traits, helping individuals to discover and make use of reproductive resources.

Are yellow dung flies domesticated cow dung specialists?

The theory of niche differentiation implies some extent of specialization of species with regard to key resources, notably food. Coprophagous (dung-eating) insect larvae play a critical role in the decomposition of livestock dung in modern and traditional agricultural grasslands. The yellow dung fly (Scathophaga stercoraria L.; Diptera: Scathophagidae) is one of the largest, most common and abundant dung decomposers on pastures in cold-temperate regions across the entire northern hemisphere. As this fly is often associated with domesticated cattle or dairy cows, which are commonly kept for human nutrition worldwide (beef, milk, cheese, etc.), it is sometimes suspected to be a cow dung specialist. However, yellow dung flies are regularly active on and around other dung types, and must have reproduced on dung of wild vertebrates before the domestication of cattle. We therefore experimentally studied the performance of yellow dung fly larvae on dung of various large domestic vs. wild mammals (cow, horse, wild boar, red deer) in the laboratory in Switzerland. Larval performance in terms of juvenile survival, egg-to-adult development time, growth rate, and final adult body size, the major life history indicators of individual reproductive success, did not vary greatly among the various dung types tested. Thus, yellow dung flies can successfully reproduce on multiple types of mammal (vertebrate) dung, wild and domestic, and are therefore dung generalists rather than specialists. We conclude that yellow dung flies are common in European low- and highlands because they could plastically shift to dung of common herbivorous livestock after their domestication without losing the ability to reproduce on dung of common wild mammals.

Neural dynamics in the limbic system during male social behaviors

Sexual and aggressive behaviors are vital for species survival and individual reproductive success. Although many limbic regions have been found relevant to these behaviors, how social cues are represented across regions and how the network activity generates each behavior remains elusive. To answer these questions, we utilize multi-fiber photometry (MFP) to simultaneously record Ca2+ signals of estrogen receptor alpha (Esr1)-expressing cells from 13 limbic regions in male mice during mating and fighting. We find that conspecific sensory information and social action signals are widely distributed in the limbic system and can be decoded from the network activity. Cross-region correlation analysis reveals striking increases in the network functional connectivity during the social action initiation phase, whereas late copulation is accompanied by a "dissociated" network state. Based on the response patterns, we propose a mating-biased network (MBN) and an aggression-biased network (ABN) for mediating male sexual and aggressive behaviors, respectively.

Robustness of Hill's overlapping-generation method for calculating Ne to extreme patterns of reproductive success.

For species with overlapping generations, the most widely used method to calculate effective population size (Ne) is Hill's, the key parameter for which is lifetime variance in offspring number ([Formula: see text]). Hill's model assumes a stable age structure and constant abundance, and sensitivity to those assumptions has been evaluated previously. Here I evaluate the robustness of Hill's model to extreme patterns of reproductive success, whose effects have not been previously examined: (1) very strong reproductive skew; (2) strong temporal autocorrelations in individual reproductive success; and (3) strong covariance of individual reproduction and survival. Genetic drift (loss of heterozygosity and increase in allele frequency variance) was simulated in age-structured populations using methods that generated no autocorrelations or covariances (Model NoCor); or created strong positive (Model Positive) or strong negative (Model Negative) temporal autocorrelations in reproduction and covariances between reproduction and survival. Compared to Model NoCor, the other models led to greatly elevated or reduced [Formula: see text], and hence greatly reduced or elevated Ne, respectively. A new index is introduced (ρα,α+), which is the correlation between (1) the number of offspring produced by each individual at the age at maturity (α), and (2) the total number of offspring produced during the rest of their lifetimes. Mean ρα,α+ was ≈0 under Model NoCor, strongly positive under Model Positive, and strongly negative under Model Negative. Even under the most extreme reproductive scenarios in Models Positive and Negative, when [Formula: see text] was calculated from the realized population pedigree and used to calculate Ne in Hill's model, the result accurately predicted the rate of genetic drift in simulated populations. These results held for scenarios where age-specific reproductive skew was random (variance ≈ mean) and highly overdispersed (variance up to 20 times the mean). Collectively, these results are good news for researchers as they demonstrate the robustness of Hill's model even in extreme reproductive scenarios.

Unraveling female communication through scent marks in the Norway rat

Chemical communication by females remains poorly understood, with most attention focused on female advertisement of sexual receptivity to males or mother-offspring communication. However, in social species, scents are likely to be important for mediating competition and cooperation between females determining individual reproductive success. Here, we explore chemical signaling by female laboratory rats (Rattus norvegicus) to test i) whether females target their deployment of scent information differentially according to their sexual receptivity and the genetic identity of both female and male conspecifics signaling in the local environment and ii) whether females are attracted to gain the same or different information from female scents compared to males. Consistent with targeting of scent information to colony members of similar genetic background, female rats increased scent marking in response to scents from females of the same strain. Females also suppressed scent marking in response to male scent from a genetically foreign strain while sexually receptive. Proteomic analysis of female scent deposits revealed a complex protein profile, contributed from several sources but dominated by clitoral gland secretion. In particular, female scent marks contained a series of clitoral-derived hydrolases and proteolytically truncated major urinary proteins (MUPs). Manipulated blends of clitoral secretion and urine from estrus females were strongly attractive to both sexes, while voided urine alone stimulated no interest. Our study reveals that information about female receptive status is shared between females as well as with males, while clitoral secretions containing a complex set of truncated MUPs and other proteins play a key role in female communication.

Thelong-lasting legacy of reproduction: lifetime reproductive success shapes expected genetic contributions of humansafter 10 generations.

An individual's lifetime reproductive success (LRS) measures its realized genetic contributions to the next generation, but how well does it predict this over longer periods? Here we use human genealogical data to estimate expected individual genetic contributions (IGC) and quantify the degree to which LRS, relative to other fitness proxies, predicts IGC over longer periods. This allows an identification of the life-history stages that are most important in shaping variation in IGC. We use historical genealogical data from two non-isolated local populations in Switzerland to estimate the stabilized IGC for 2230 individuals approximately 10 generations after they were born. We find that LRS explains 30% less variation in IGC than the best predictor of IGC, the number of grandoffspring. However, albeit less precise than the number of grandoffspring, we show that LRS does provide an unbiased prediction of IGC. Furthermore, it predicts IGC better than lifespan, and accounting for offspring survival to adulthood does not improve the explanatory power. Overall, our findings demonstrate the value of human genealogical data to evolutionary biology and suggest that reproduction-more than lifespan or offspring survival-impacts the long-term genetic contributions of historic humans, even in a population with appreciable migration.

Survive the north: transplantation for conservation of mangrove forests requires consideration of influences of low temperature, mating system and their joint effects on effective size of the reforested populations

Mangrove forests, which occur in the intertidal regions of tropical and sub-tropical zones, have high ecological and economic values. They have rapidly reduced over the past decades due to various reasons. Reforestation is a common strategy for the conservation of mangroves, but information on the renewal capacity and viability of these artificial mangrove populations is still lacking. Here, we estimated the effective population size (NE) of an artificial population of Kandelia obovata (Rhizophoraceae) and explored the effects of intrinsic and extrinsic factors on changes in NE to evaluate population viability and develop appropriate management strategies for mangrove forests. This population consisted of three ecologically and genetically differentiated groups that had experienced a major low-temperature event during the winter of 2015–2016 and varied in cold resistance. We first detected population bottlenecks and estimated contemporary values of NE for different groups using microsatellite data. Next, we performed paternity analyses for seedlings and propagules to explore variations in the reproductive success of individuals within the three groups before and after the low-temperature event. Lastly, we simulated four scenarios to characterize the effects of low temperature, mating system, and variance in reproductive success on changes in NE in a hypothetical metapopulation based on empirical estimates. Our results show that groups with moderate or poor cold resistance experienced local bottlenecks, and shifts in effective breeders occurred following the low-temperature event, which indicates that low temperature has an effect on not only population size but also reproductive success. Furthermore, our simulations revealed that changes in NE are jointly affected by reproductive success, mating system, and environmental conditions. These findings enhance our understanding of the multiple factors that affect NE, and provide key information that will aid the reforestation and management of mangrove forests, especially when they are introduced to high-latitude areas.

Reproductive success of jack and full-size males in a wild coho salmon population.

Despite the wealth of research on Pacific salmon Oncorhynchus spp. life histories there is limited understanding of the lifetime reproductive success of males that spend less time at sea and mature at a smaller size (jacks) than full-size males. Over half of returning male spawners can be jacks in some populations, so it is crucial to understand their contribution to population productivity. We quantified adult-to-adult reproductive success (RS) of jacks and their relative reproductive success (RRS) compared to full-size males in a wild population of coho salmon in the Auke Creek watershed, Juneau, Alaska. We used genetic data from nearly all individuals (approx. 8000) returning to spawn over a decade (2009-2019) to conduct parentage analysis and calculate individual RS. The average adult-to-adult RS of jacks (mean = 0.7 and s.e. = 0.1) was less than that of full-size males (mean = 1.1 and s.e. = 0.1). Jack RRS was consistently below 1.0 but ranged widely (0.23 to 0.96). Despite their lower average success, jacks contributed substantially to the population by siring 23% of the total returning adult offspring (1033 of 4456) produced between 2009 and 2015. Our results imply that jacks can affect evolutionary and population dynamics, and are relevant to the conservation and management of Pacific salmon.

Is Fluctuating Asymmetry a Sufficient Indicator of Stress Level in Two Lizard Species (Zootoca vivipara and Lacerta agilis) from Alpine Habitats?

Alpine habitats are exposed to increasing anthropogenic pressure and climate change. The negative impacts can lead to chronic stress that can affect the survival and reproductive success of individuals and even lead to population extinction. In this study, we analyse different morphological and ecological traits and indices of abiotic and biotic stressors (such as head size and shape, fluctuating asymmetry, body condition index, tail autotomy, and population abundance) in alpine and subalpine populations of two lacertid species (Zootoca vivipara and Lacerta agilis) from Serbia and North Macedonia. These lizards live under different conditions: allotopy/syntopy, different anthropogenic pressure, and different levels of habitat protection. We found differences between syntopic and allotopic populations in pileus size, body condition index (in both species), pileus shape, fluctuating asymmetry (in L. agilis), and abundance (in Z. vivipara). Differences between populations under anthropogenic pressure and populations without it were observed in pileus shape, body condition index (in both species), pileus size, fluctuating asymmetry, tail autotomy and abundance (in L. agilis). On the basis of our results, it is necessary to include other stress indicators in addition to fluctuating asymmetry to quickly observe and quantify the negative effects of threat factors and apply protective measures.

Do foraging ecology and contaminants interactively predict parenting hormone levels in common eider?

Global climate change is causing abiotic shifts such as higher air and ocean temperatures, and disappearing sea ice in Arctic ecosystems. These changes influence Arctic-breeding seabird foraging ecology by altering prey availability and selection, affecting individual body condition, reproductive success, and exposure to contaminants such as mercury (Hg). The cumulative effects of alterations to foraging ecology and Hg exposure may interactively alter the secretion of key reproductive hormones such as prolactin (PRL), important for parental attachment to eggs and offspring and overall reproductive success. However, more research is needed to investigate the relationships between these potential links. Using data collected from 106 incubating female common eiders (Somateria mollissima) at six Arctic and sub-Arctic colonies, we examined whether the relationship between individual foraging ecology (assessed using δ13C, δ15N) and total Hg (THg) exposure predicted PRL levels. We found a significant, complex interaction between δ13C, δ15N and THg on PRL, suggesting that individuals cumulatively foraging at lower trophic levels, in phytoplankton-dominant environments, and with the highest THg levels had the most constant significant relationship PRL levels. Cumulatively, these three interactive variables resulted in lowered PRL. Overall, results demonstrate the potential downstream and cumulative implications of environmentally induced changes in foraging ecology, in combination with THg exposure, on hormones known to influence reproductive success in seabirds. These findings are notable in the context of continuing environmental and food web changes in Arctic systems, which may make seabird populations more susceptible to ongoing stressors.

Do females of Whiskered Tern Chlidonias hybrida renest after offspring desertion?

In species with biparental care, one parent may relieve him-/herself of the costs of parental care by deserting and leaving the mate to care for the offspring. Offspring desertion by a single parent, usually the female, occurs in many bird species, especially in Charadriiformes. The reasons for desertion are still being debated, and compelling evidence for the benefits and costs to either sex is not yet handed. The benefits of desertion for the deserter may include renesting with a new mate shortly after desertion. Therefore, desertion may enhance individual lifetime reproductive success. In Whiskered Tern Chlidonias hybrida almost all females desert, about half of females desert during chick-rearing period, the other half during the post-fledging period. Based on intensive fieldwork over a 4-year study, we investigated whether Whiskered Tern females remated and renested after desertion. We assumed that favourable conditions, such as an extended breeding season and food abundance resulting in a high breeding success would prompt most females to renest after desertion. In contrast to this expectation, however, only 5% of females that deserted during chick-rearing period (2 out of 41) remated and renested in the study area. About 50 days elapsed between the initiations of the first and second clutch by these two females. Females also participated in courtship behaviour after desertion (without renesting). We thus estimate that 15% (6 out of 41) of females that deserted during chick-rearing period were subsequently involved in some kind of breeding activity. Therefore, renesting does not seem to be a major direct benefit for deserting Whiskered Tern females. Females of this species probably benefit from desertion in other ways, but these have yet to be discovered.

NMπ 2.0: Software update to minimize the risk of false positives among determinants of reproductive success.

In plant populations, parentage analysis helps understand factors shaping individual reproductive success. However, estimating reproductive success determinants based on parentage counts requires decoupling the effects of individual fecundity and propagule dispersal. The neighbourhood model implemented in the NMπ software provides a standard solution for this problem based on the fixed-effects regression-like approach. Nonetheless, it has been recently shown that the method is prone to false discoveries when important fecundity determinants are omitted. To account for the unexplained variance in fecundity, the Bayesian approach was developed based on the new model (the hierarchical neighbourhood model; HNM). Here, I present the NMπ software update that allows the HNM approach to be used in the framework of a friendly interface. More importantly, the HNM approach is now made available for both dispersed (seedlings) and nondispersed (seeds with known mothers) progeny data. The Bayesian approach, among others, selects significant fecundity determinants, estimates the proportion of variance in reproductive potential explained by selected determinants (R2 ), and provides individual female and male fecundity values. Although the software was designed to handle microsatellite marker data, a solution is proposed for large sets of single nucleotide polymorphisms. The program can be run on Windows (using either a terminal or a graphical interface) as well as (using a terminal) on Linux, or macOS platforms. In any case, NMπ can utilize multicore processors to speed up the analysis. The updated package containing the code, the executable file, the user manual, and example data is available at https://www.ukw.edu.pl/pracownicy/plik/igor_chybicki/3694/.

Sweepstakes reproductive success via pervasive and recurrent selective sweeps.

Highly fecund natural populations characterized by high early mortality abound, yet our knowledge about their recruitment dynamics is somewhat rudimentary. This knowledge gap has implications for our understanding of genetic variation, population connectivity, local adaptation, and the resilience of highly fecund populations. The concept of sweepstakes reproductive success, which posits a considerable variance and skew in individual reproductive output, is key to understanding the distribution of individual reproductive success. However, it still needs to be determined whether highly fecund organisms reproduce through sweepstakes and, if they do, the relative roles of neutral and selective sweepstakes. Here, we use coalescent-based statistical analysis of population genomic data to show that selective sweepstakes likely explain recruitment dynamics in the highly fecund Atlantic cod. We show that the Kingman coalescent (modelling no sweepstakes) and the Xi-Beta coalescent (modelling random sweepstakes), including complex demography and background selection, do not provide an adequate fit for the data. The Durrett-Schweinsberg coalescent, in which selective sweepstakes result from recurrent and pervasive selective sweeps of new mutations, offers greater explanatory power. Our results show that models of sweepstakes reproduction and multiple-merger coalescents are relevant and necessary for understanding genetic diversity in highly fecund natural populations. These findings have fundamental implications for understanding the recruitment variation of fish stocks and general evolutionary genomics of high-fecundity organisms.

Effects of an IgE receptor polymorphism acting on immunity, susceptibility to infection, and reproduction in a wild rodent.

The genotype of an individual is an important predictor of their immune function, and subsequently, their ability to control or avoid infection and ultimately contribute offspring to the next generation. However, the same genotype, subjected to different intrinsic and/or extrinsic environments, can also result in different phenotypic outcomes, which can be missed in controlled laboratory studies. Natural wildlife populations, which capture both genotypic and environmental variability, provide an opportunity to more fully understand the phenotypic expression of genetic variation. We identified a synonymous polymorphism in the high-affinity Immunoglobulin E (IgE) receptor (GC and non-GC haplotypes) that has sex-dependent effects on immune gene expression, susceptibility to infection, and reproductive success of individuals in a natural population of field voles (Microtus agrestis). We found that the effect of the GC haplotype on the expression of immune genes differed between sexes. Regardless of sex, both pro-inflammatory and anti-inflammatory genes were more highly relatively expressed in individuals with the GC haplotype than individuals without the haplotype. However, males with the GC haplotype showed a stronger signal for pro-inflammatory genes, while females showed a stronger signal for anti-inflammatory genes. Furthermore, we found an effect of the GC haplotype on the probability of infection with a common microparasite, Babesia microti, in females - with females carrying the GC haplotype being more likely to be infected. Finally, we found an effect of the GC haplotype on reproductive success in males - with males carrying the GC haplotype having a lower reproductive success. This is a rare example of a polymorphism whose consequences we are able to follow across immunity, infection, and reproduction for both males and females in a natural population.

A body composition model with multiple storage compartments for polar bears (Ursus maritimus).

Climate warming is rapidly altering Arctic ecosystems. Polar bears (Ursus maritimus) need sea ice as a platform from which to hunt seals, but increased sea-ice loss is lengthening periods when bears are without access to primary hunting habitat. During periods of food scarcity, survival depends on the energy that a bear has stored in body reserves, termed storage energy, making this a key metric in predictive models assessing climate change impacts on polar bears. Here, we developed a body composition model for polar bears that estimates storage energy while accounting for changes in storage tissue composition. We used data of dissected polar bears (n =31) to link routinely collected field measures of total body mass and straight-line body length to the body composition of individual bears, described in terms of structural mass and two storage compartments, adipose and muscle. We then estimated the masses of metabolizable proteins and lipids within these storage compartments, giving total storage energy. We tested this multi-storage model by using it to predict changes in the lipid stores from an independent dataset of wild polar bears (n = 36) that were recaptured 8-200days later. Using length and mass measurements, our model successfully predicted direct measurements of lipid changes via isotopic dilutions (root mean squared error of 14.5kg). Separating storage into two compartments, and allowing the molecular composition of storage to vary, provides new avenues for quantifying energy stores of individuals across their life cycle. The multi-storage body composition model thus provides a basis for further exploring energetic costs of physiological processes that contribute to individual survival and reproductive success. Given bioenergetic models are increasingly used as a tool to predict individual fitness and population dynamics, our approach for estimating individual energy stores could be applicable to a wide range of species.

Growth in marine mammals: a review of growth patterns, composition and energy investment.

Growth of structural mass and energy reserves influences individual survival, reproductive success, population and species life history. Metrics of structural growth and energy storage of individuals are often used to assess population health and reproductive potential, which can inform conservation. However, the energetic costs of tissue deposition for structural growth and energy stores and their prioritization within bioenergetic budgets are poorly documented. This is particularly true across marine mammal species as resources are accumulated at sea, limiting the ability to measure energy allocation and prioritization. We reviewed the literature on marine mammal growth to summarize growth patterns, explore their tissue compositions, assess the energetic costs of depositing these tissues and explore the tradeoffs associated with growth. Generally, marine mammals exhibit logarithmic growth. This means that the energetic costs related to growth and tissue deposition are high for early postnatal animals, but small compared to the total energy budget as animals get older. Growth patterns can also change in response to resource availability, habitat and other energy demands, such that they can serve as an indicator of individual and population health. Composition of tissues remained consistent with respect to protein and water content across species; however, there was a high degree of variability in the lipid content of both muscle (0.1-74.3%) and blubber (0.4-97.9%) due to the use of lipids as energy storage. We found that relatively few well-studied species dominate the literature, leaving data gaps for entire taxa, such as beaked whales. The purpose of this review was to identify such gaps, to inform future research priorities and to improve our understanding of how marine mammals grow and the associated energetic costs.

Response to drift kelp is modified by both substratum and season: green sea urchin foraging behaviour in subsidized habitats

The ability of organisms to detect, locate and navigate to resource patches is modified by the surrounding landscape. Green sea urchinsStrongylocentrotus droebachiensisin barren grounds exist in a food-limited state and are subject to intense competition. Rapid detection and consumption of resource patches, particularly pieces of macroalgae from adjacent algal beds, are key in determining individual growth, survival and reproductive success. Detection and movement to resource patches requires moving through a heterogeneous benthic seascape composed of rocky and sandy patches, presenting different degrees of resistance to movement. We used time-lapse photography to describe the foraging behaviour of urchins in relation to the presence of a key resource subsidy (drift kelp) and different benthic seascapes. We demonstrated that urchins could detect the presence of drift kelp in barren-ground habitats and alter their movement behaviour in response, but did not exhibit the ability to directionally navigate towards kelp in field conditions. Seascapes with increased proportions of rocky substrata facilitated increased movement in response to the presence of drift. Moreover, urchin foraging behaviour was temporally variable, with no response to the presence of drift in early spring (May). This indicates not only that interpretations of observations of urchin behaviour must take intrinsic and extrinsic seasonal dynamics into account, but that extrapolating results to explain larger-scale patterns and processes must include both spatially explicit subtidal seascapes and temporal dynamics. In many temperate and boreal regions, this indicates the need for increased subtidal benthic research in the fall and winter.