Coral reef fish density at a tourist destination responded rapidly to COVID-19 restrictions.

Throughout the world, anthropogenic pressure on natural ecosystems is intensifying, notably through urbanisation, economic development, and tourism. Coral reefs have become exposed to stressors related to tourism. To reveal the impact of human activities on fish communities, we used COVID-19-related social restrictions in 2021. In French Polynesia, from February to December 2021, there was a series of restrictions on local activities and international tourism. We assessed the response of fish populations in terms of changes in the species richness and density of fish in the lagoon of Bora-Bora (French Polynesia). We selected sites with varying human pressures-some dedicated to tourism activities, others affected by boat traffic, and control sites with little human presence. Underwater visual surveys demonstrated that fish density and richness differed spatially and temporally. They were lowest on sites affected by boat traffic regardless of pandemic-related restrictions, and when activities were authorised; they were highest during lockdowns. Adult fish density increased threefold on sites usually affected by boat traffic during lockdowns and increased 2.7-fold on eco-tourism sites during international travel bans. Human activities are major drivers of fish density and species richness spatially across the lagoon of Bora-Bora but also temporally across pandemic-related restrictions, with dynamic responses to different restrictions. These results highlight the opportunity provided by pauses in human activities to assess their impact on the environment and confirm the need for sustainable lagoon management in Bora-Bora and similar coral reef settings affected by tourism and boat traffic.

Long-term exposure to artificial light at night in the wild decreases survival and growth of a coral reef fish.

Artificial light at night (ALAN) is an increasing anthropogenic pollutant, closely associated with human population density, and now well recognized in both terrestrial and aquatic environments. However, we have a relatively poor understanding of the effects of ALAN in the marine realm. Here, we carried out a field experiment in the coral reef lagoon of Moorea, French Polynesia, to investigate the effects of long-term exposure (18-23 months) to chronic light pollution at night on the survival and growth of wild juvenile orange-fin anemonefish, Amphiprion chrysopterus. Long-term exposure to environmentally relevant underwater illuminance (mean: 4.3 lux), reduced survival (mean: 36%) and growth (mean: 44%) of juvenile anemonefish compared to that of juveniles exposed to natural moonlight underwater (mean: 0.03 lux). Our study carried out in an ecologically realistic situation in which the direct effects of artificial lighting on juvenile anemonefish are combined with the indirect consequences of artificial lighting on other species, such as their competitors, predators, and prey, revealed the negative impacts of ALAN on life-history traits. Not only are there immediate impacts of ALAN on mortality, but the decreased growth of surviving individuals may also have considerable fitness consequences later in life. Future studies examining the mechanisms behind these findings are vital to understand how organisms can cope and survive in nature under this globally increasing pollutant.

Anemone bleaching increases the metabolic demands of symbiont anemonefish.

Increased ocean temperatures are causing mass bleaching of anemones and corals in the tropics worldwide. While such heat-induced loss of algal symbionts (zooxanthellae) directly affects anemones and corals physiologically, this damage may also cascade on to other animal symbionts. Metabolic rate is an integrative physiological trait shown to relate to various aspects of organismal performance, behaviour and locomotor capacity, and also shows plasticity during exposure to acute and chronic stressors. As climate warming is expected to affect the physiology, behaviour and life history of animals, including ectotherms such as fish, we measured if residing in bleached versus unbleached sea anemones (Heteractis magnifica) affected the standard (i.e. baseline) metabolic rate and behaviour (activity) of juvenile orange-fin anemonefish (Amphiprion chrysopterus). Metabolic rate was estimated from rates of oxygen uptake [Formula: see text], and the standard metabolic rate [Formula: see text] of anemonefish from bleached anemones was significantly higher by 8.2% compared with that of fish residing in unbleached anemones, possibly due to increased stress levels. Activity levels did not differ between fish from bleached and unbleached anemones. As [Formula: see text] reflects the minimum cost of living, the increased metabolic demands may contribute to the negative impacts of bleaching on important anemonefish life history and fitness traits observed previously (e.g. reduced spawning frequency and lower fecundity).

Isolation and Synthesis of Laxaphycin B-Type Peptides: A Case Study and Clues to Their Biosynthesis.

The laxaphyci's B family constitutes a group of five related cyclic lipopeptides isolated from diverse cyanobacteria from all around the world. This group shares a typical structure of 12 amino acids from the l and d series, some of them hydroxylated at the beta position, and all containing a rare beta-amino decanoic acid. Nevertheless, they can be differentiated due to slight variations in the composition of their amino acids, but the configuration of their alpha carbon remains conserved. Here, we provide the synthesis and characterization of new laxaphycin B-type peptides. In doing so we discuss how the synthesis of laxaphycin B and analogues was developed. We also isolate minor acyclic laxaphycins B, which are considered clues to their biosynthesis.

A new versatile primer set targeting a short fragment of the mitochondrial COI region for metabarcoding metazoan diversity: application for characterizing coral reef fish gut contents.

INTRODUCTION: The PCR-based analysis of homologous genes has become one of the most powerful approaches for species detection and identification, particularly with the recent availability of Next Generation Sequencing platforms (NGS) making it possible to identify species composition from a broad range of environmental samples. Identifying species from these samples relies on the ability to match sequences with reference barcodes for taxonomic identification. Unfortunately, most studies of environmental samples have targeted ribosomal markers, despite the fact that the mitochondrial Cytochrome c Oxidase subunit I gene (COI) is by far the most widely available sequence region in public reference libraries. This is largely because the available versatile ("universal") COI primers target the 658 barcoding region, whose size is considered too large for many NGS applications. Moreover, traditional barcoding primers are known to be poorly conserved across some taxonomic groups. RESULTS: We first design a new PCR primer within the highly variable mitochondrial COI region, the "mlCOIintF" primer. We then show that this newly designed forward primer combined with the "jgHCO2198" reverse primer to target a 313 bp fragment performs well across metazoan diversity, with higher success rates than versatile primer sets traditionally used for DNA barcoding (i.e. LCO1490/HCO2198). Finally, we demonstrate how the shorter COI fragment coupled with an efficient bioinformatics pipeline can be used to characterize species diversity from environmental samples by pyrosequencing. We examine the gut contents of three species of planktivorous and benthivorous coral reef fish (family: Apogonidae and Holocentridae). After the removal of dubious COI sequences, we obtained a total of 334 prey Operational Taxonomic Units (OTUs) belonging to 14 phyla from 16 fish guts. Of these, 52.5% matched a reference barcode (>98% sequence similarity) and an additional 32% could be assigned to a higher taxonomic level using Bayesian assignment. CONCLUSIONS: The molecular analysis of gut contents targeting the 313 COI fragment using the newly designed mlCOIintF primer in combination with the jgHCO2198 primer offers enormous promise for metazoan metabarcoding studies. We believe that this primer set will be a valuable asset for a range of applications from large-scale biodiversity assessments to food web studies.

Roadside sales activities in a South Pacific Island (Bora-Bora) reveal sustainable strategies for local food supply during a pandemic.

During the COVID-19 pandemic, the reduced exports and imports as well as the lack of activity due to the interruption in the international tourism economy seriously impacted food security in many Pacific Islands. People often returned to natural resources to provide for themselves, their families, or to generate income. On Bora-Bora Island, the major tourist destination in French Polynesia, roadside sales are widespread. Our study analyses the impact of the COVID-19 pandemic on roadside sales activities through a census of roadside stalls on the five Bora-Bora districts conducted before (January and February 2020), during (from March 2020 to October 2021) and after (from November to December 2021) health-related activity and travel restrictions. Our results showed that the marketing system for local products (fruits, vegetables, cooked meals, and fish) increased in the form of roadside sales during the COVID-19 in two of the five districts of Bora-Bora. Roadside selling would be an alternative system for providing food to the population at Bora-Bora during a global crisis and that could reveal itself sustainable after this pandemic.

Intralocus sexual conflict for fitness: sexually antagonistic alleles for testosterone.

Intralocus sexual conflict occurs when a trait encoded by the same genetic locus in the two sexes has different optima in males and females. Such conflict is widespread across taxa, however, the shared phenotypic traits that mediate the conflict are largely unknown. We examined whether the sex hormone, testosterone (T), that controls sexual differentiation, contributes to sexually antagonistic fitness variation in the bank vole, Myodes glareolus. We compared (opposite-sex) sibling reproductive fitness in the bank vole after creating divergent selection lines for T. This study shows that selection for T was differentially associated with son versus daughter reproductive success, causing a negative correlation in fitness between full siblings. Our results demonstrate the presence of intralocus sexual conflict for fitness in this small mammal and that sexually antagonistic selection is acting on T. We also found a negative correlation in fitness between parents and their opposite-sex progeny (e.g. father-daughter), highlighting a dilemma for females, as the indirect genetic benefits of selecting reproductively successful males (high T) are lost with daughters. We discuss mechanisms that may mitigate this disparity between progeny quality.

Sexual antagonism for testosterone maintains multiple mating behaviour.

1. The persistence of multiple mating remains one of the fundamental questions in evolutionary biology. In theory, multiple mating is predicted to improve female fitness cumulatively through direct and/or genetic benefits. However, intra-locus sexual conflicts may potentially constrain or even eliminate these benefits owing to the gender load imposed by sexually antagonistic selection. 2. Here, we tested whether sexually antagonistic selection can maintain the variance in multiple mating behaviour of bank voles (Myodes glareolus) by manipulating the hormone testosterone through artificial selection in the laboratory. Among mammals, testosterone is a sexually dimorphic fitness-related trait under selection and is known to affect mating behaviour. We conducted mating trials in which females derived from family-based selection of testosterone were sequentially paired with four males of different testosterone profiles. 3. We show that artificial selection for high testosterone increased the mating rate of males, but clearly decreased the number of partners that females mated with (and vice versa). Because multiple mating was beneficial for the reproductive success of both sexes, as evidenced by the positive Bateman gradients, the divergent evolutionary interests of testosterone between the sexes can maintain this polygynandrous mating system. 4. Our results highlight how mating rate is concordantly selected in both sexes; however, it is largely influenced by testosterone, which is under sexually antagonistic selection. 5. This study is the first one to emphasise the direct and indirect effects of the endocrine system not only on reproductive physiology and behaviour but also for the evolution of genetic mating strategies in mammals.

Harbours as unique environmental sites of multiple anthropogenic stressors on fish hormonal systems.

Fish development and acclimation to environmental conditions are strongly mediated by the hormonal endocrine system. In environments contaminated by anthropogenic stressors, hormonal pathway alterations can be detrimental for growth, survival, fitness, and at a larger scale for population maintenance. In the context of increasingly contaminated marine environments worldwide, numerous laboratory studies have confirmed the effect of one or a combination of pollutants on fish hormonal systems. However, this has not been confirmed in situ. In this review, we explore the body of knowledge related to the influence of anthropogenic stressors disrupting fish endocrine systems, recent advances (focusing on thyroid hormones and stress hormones such as cortisol), and potential research perspectives. Through this review, we highlight how harbours can be used as "in situ laboratories" given the variety of anthropogenic stressors (such as plastic, chemical, sound, light pollution, and invasive species) that can be simultaneously investigated in harbours over long periods of time.

d-Peptidase Activity in a Marine Mollusk Detoxifies a Nonribosomal Cyclic Lipopeptide: An Ecological Model to Study Antibiotic Resistance.

In the marine environment, sessile cyanobacteria have developed chemical strategies for protection against grazers. In turn, herbivores have to circumvent these defenses and in certain cases even take advantage of them as shelter from their own predators. This is the case of Stylocheilus striatus, a sea hare that feeds on Anabaena torulosa, a cyanobacterium that produces toxic cyclic lipopeptides of the laxaphycin B family. S. striatus consumes the cyanobacterium without being affected by the toxicity of its compounds and also uses it as an invisibility cloak against predators. In this article, using different substrates analogous to laxaphycin B, we demonstrate the presence of an enzyme in the digestive gland of the mollusk that is able to biotransform laxaphycin B derivatives. The enzyme belongs to the poorly known family of d-peptidases that are suspected to be involved in antibiotic resistance.

Intra- and intersexual trade-offs between testosterone and immune system: Implications for sexual and sexually antagonistic selection.

Parasites indirectly affect life-history evolution of most species. Combating parasites requires costly immune defenses that are assumed to trade off with other life-history traits. In vertebrate males, immune defense is thought to trade off with reproductive success, as androgens enhancing sexual signaling can suppress immunity. The phenotypic relationship between male androgen levels and immune function has been addressed in many experimental studies. However, these do not provide information on either intra- or intersex genetic correlations, necessary for understanding sexual and sexually antagonistic selection theories. We measured male and female humoral antibody responses to a novel antigen (bovine gamma globulin), total immunoglobulin G, and the male testosterone level of a laboratory population of the bank vole (Myodes glareolus). Although we studied five traits, factor-analytic modeling of the additive genetic (co)variance matrix within a restricted maximum likelihood-animal model supported genetic variation in three dimensions. Sixty-five percent of the genetic variation contrasted testosterone with both immune measures in both sexes; consequently, selection for the male trait (testosterone) will have correlated effects on the immune system not only in males but also in females. Thus, our study revealed an intra- and intersexual genetic trade-off between immunocompetence and male reproductive effort, of which only indirect evidence has existed so far.

Crime and punishment in a roaming cleanerfish.

Cheating is common in cooperative interactions, but its occurrence can be controlled by various means ranging from rewarding cooperators to active punishment of cheaters. Punishment occurs in the mutualism involving the cleanerfish Labroides dimidiatus and its reef fish clients. When L. dimidiatus cheats, by taking scales and mucus rather than ectoparasites, wronged clients either chase or withhold further visits to the dishonest cleaner, which leads to more cooperative future interactions. Punishment of cheating L. dimidiatus may be effective largely because these cleaners are strictly site-attached, increasing the potential for repeated interactions between individual cleaners and clients. Here, we contrast the patterns of cheating and punishment in L. dimidiatus with its close relative, the less site-attached Labroides bicolor. Overall, L. bicolor had larger home ranges, cheated more often and, contrary to our prediction, were punished by cheated clients as frequently as, and not less often than, L. dimidiatus. However, adult L. bicolor, which had the largest home ranges, did not cheat more than younger conspecifics, suggesting that roaming, and hence the frequency of repeated interactions, has little influence on cheating and retaliation in cleaner-client relationships. We suggest that roaming cleaners offer the only option available to many site-attached reef fish seeking a cleaning service. This asymmetry in scope for partner choice encourages dishonesty by the partner with more options (i.e. L. bicolor), but to be cleaned by a cleaner that sometimes cheats may be a better option than not to be cleaned at all.

Hormonal and behavioural effects of motorboat noise on wild coral reef fish.

Anthropogenic noise is an emergent ecological pollutant in both terrestrial and aquatic habitats. Human population growth, urbanisation, resource extraction, transport and motorised recreation lead to elevated noise that affects animal behaviour and physiology, impacting individual fitness. Currently, we have a poor mechanistic understanding of the effects of anthropogenic noise, but a likely candidate is the neuroendocrine system that integrates information about environmental stressors to produce regulatory hormones; glucocorticoids (GCs) and androgens enable rapid individual phenotypic adjustments that can increase survival. Here, we carried out two field-based experiments to investigate the effects of short-term (30 min) and longer-term (48 h) motorboat-noise playback on the behaviour, GCs (cortisol) and androgens of site-attached free-living orange-fin anemonefish (Amphiprion chrysopterus). In the short-term, anemonefish exposed to motorboat-noise playback showed both behavioural and hormonal responses: hiding and aggression increased, and distance moved out of the anemone decreased in both sexes; there were no effects on cortisol levels, but male androgen levels (11-ketotestosterone and testosterone) increased. Some behaviours showed carry-over effects from motorboat noise after it had ceased, and there was no evidence for a short-term change in response to subsequent motorboat-noise playback. Similarly, there was no evidence that longer-term exposure led to changes in response: motorboat noise had an equivalent effect on anemonefish behaviour and hormones after 48 h as on first exposure. Longer-term noise exposure led to higher levels of cortisol in both sexes and higher testosterone levels in males, and stress-responses to an additional environmental challenge in both sexes were impaired. Circulating androgen levels correlated with aggression, while cortisol levels correlated with hiding, demonstrating in a wild population that androgen/glucocorticoid pathways are plausible proximate mechanisms driving behavioural responses to anthropogenic noise. Combining functional and mechanistic studies are crucial for a full understanding of this global pollutant.

Near-future ocean warming and acidification alter foraging behaviour, locomotion, and metabolic rate in a keystone marine mollusc.

Environmentally-induced changes in fitness are mediated by direct effects on physiology and behaviour, which are tightly linked. We investigated how predicted ocean warming (OW) and acidification (OA) affect key ecological behaviours (locomotion speed and foraging success) and metabolic rate of a keystone marine mollusc, the sea hare Stylocheilus striatus, a specialist grazer of the toxic cyanobacterium Lyngbya majuscula. We acclimated sea hares to OW and/or OA across three developmental stages (metamorphic, juvenile, and adult) or as adults only, and compare these to sea hares maintained under current-day conditions. Generally, locomotion speed and time to locate food were reduced ~1.5- to 2-fold when the stressors (OW or OA) were experienced in isolation, but reduced ~3-fold when combined. Decision-making was also severely altered, with correct foraging choice nearly 40% lower under combined stressors. Metabolic rate appeared to acclimate to the stressors in isolation, but was significantly elevated under combined stressors. Overall, sea hares that developed under OW and/or OA exhibited a less severe impact, indicating beneficial phenotypic plasticity. Reduced foraging success coupled with increased metabolic demands may impact fitness in this species and highlight potentially large ecological consequences under unabated OW and OA, namely in regulating toxic cyanobacteria blooms on coral reefs.

Testosterone-mediated effects on fitness-related phenotypic traits and fitness.

The physiological and behavioral mechanisms underlying life-history trade-offs are a continued source of debate. Testosterone (T) is one physiological factor proposed to mediate the trade-off between reproduction and survival. We use phenotypic engineering and multiple laboratory and field fitness-related phenotypic traits to test the effects of elevated T between two bank vole Myodes glareolus groups: dominant and subordinate males. Males with naturally high T levels showed higher social status (laboratory dominance) and mobility (distance between capture sites) than low-T males, and the effect of T on immune response was also T group specific, suggesting that behavioral strategies may exist in male bank voles due to the correlated responses of T. Exogenous T enhanced social status, mate searching (polygon of capture sites), mobility, and reproductive success (relative measure of pups sired). However, exogenous T also resulted in the reduction of immune function, but only in males from the high-T group. This result may be explained either by the immunosuppression costs of T or by differential sensitivity of different behavioral strategies to steroids. Circulating T levels were found to be heritable; therefore, female bank voles would derive indirect genetic benefits via good genes from mating with males signaling dominance.

Infanticide in the evolution of reproductive synchrony: effects on reproductive success.

Synchronous breeding in animals and plants has stimulated both a theoretical and empirical examination of the possible benefits of active synchronization. The selective pressures of predation and infanticide are the strongest candidates proposed to explain the evolution of reproductive synchrony. Alternatively, breeding asynchronously with conspecifics may ensure a greater availability of resources per breeder. However, the possible fitness benefits resulting from active asynchronization have not yet received attention in evolutionary ecology. Here we present a hypothesis, based on a graphical model, illustrating the costs and benefits of the two modes of reproduction. We tested the hypothesis empirically using a 2 x 2 full factorial study design, where reproductive synchrony and infanticide tactics were manipulated in enclosed populations of the bank vole. The results reveal a relationship between infanticide tactics and breeding synchrony as illustrated by our hypothesis. In general, female reproductive success (number and size of offspring surviving to weaning) was significantly lower in infanticidal populations. Moreover, an asynchronous breeding pattern proved to be advantageous in the noninfanticidal population but this advantage of asynchrony was lost as infanticide became common in the population. Our findings support the idea that synchronous reproduction could have evolved as a counterstrategy against infanticide.

Gonadotropin hormone modulation of testosterone, immune function, performance, and behavioral trade-offs among male morphs of the lizard Uta stansburiana.

Sexual selection predicts that trade-offs maintain trait variation in alternative reproductive strategies. Experiments often focus on testosterone (T), but the gonadotropins follicle-stimulating hormone and luteinizing hormone may provide a clearer understanding of the pleiotropic relationships among traits. We assess the activational role of gonadotropins on T and corticosterone regulation in traits expressed by polymorphic male side-blotched lizards Uta stansburiana. Gonadotropins are found to enhance and suppress multiple physiological, morphological, and behavioral traits independently, as well as indirectly via T, and we demonstrate selective trade-offs between reproduction and survival. The OBY locus, a genetic marker in our model vertebrate mating system, allows characterization of the interaction between genotype and hormone treatment on male traits. Our results suggest that oo, ob, and bb males are near their physiological and behavioral capacity for reproductive success, whereas yy and by males are maintained below their physiological maximum. Both by and yy morphs show trait plasticity, and we demonstrate that gonadotropins are likely proximate effectors that govern not only trait differences between alternative mating strategies but also morph plasticity. Gonadotropins clearly represent an important mechanism maintaining variation in physiological, morphological, and behavioral traits, as well as potentially maintaining the immunosuppression costs of male sexual signals.

Ephemeral and Localized Outbreaks of the Coral Predator Acanthaster cf. solaris in the Southwestern Lagoon of New Caledonia.

Mehdi Adjeroud, Mohsen Kayal, Christophe Peignon, Matthieu Juncker, Suzanne C. Mills, Ricardo Beldade, and Pascal Dumas (2018) Outbreaks of the coral predator Acanthaster spp., the crown-of-thorns seastar (COTS), cause major coral declines across the Indo-Pacific. However, the processes surrounding the initiation and propagation of COTS outbreaks are still unclear. We observed COTS outbreak abundances on several mid-shelf and inner-barrier reefs in the southern section of the New Caledonian lagoon, a multi-location initiation event that we expected to precede a broader region-wide disturbance. However, reef monitoring over 3 years revealed the highly localized and ephemeral character of these outbreaks. Outbreaks that were observed at four reef locations at the beginning of the survey simply faded away, without any specific management actions such as culling efforts. We also found no distinct reef biotope on which COTS outbreaks originated, although mid-shelf and inner-shelf barrier reefs seem to be favoured. New Caledonia has been exempted from to widespread regional COTS outbreaks, which is surprising given its proximity to the Australian Great Barrier Reef - a reef complex particularly vulnerable to COTS - and Vanuatu - where large-scale outbreaks were recorded during the same period. The availability of coral prey is probably not a limiting factor for the propagation of COTS outbreaks on New Caledonian reefs, given the high abundance and diversity of coral assemblages. Our findings reveal that localized and ephemeral COTS outbreaks can be naturally contained and do not necessarily result in widespread disturbances.

Signal reliability compromised by genotype-by-environment interaction and potential mechanisms for its preservation.

Sexual selection based on signaling requires that signals used by females in mate choice are reliable indicators of a male's heritable total fitness. A signal and the preference for it are expected to be heritable, resulting in the maintenance of genetic covariance between these two traits. However, a recent article has proposed that signals may quickly become unreliable in the presence of both environmental variation and genotype-by-environment interaction (G x E) with crossing reaction norms, potentially compromising the mechanisms of sexual selection. Here we examine the heritability and plasticity of a male dominance advertisement in the bank vole, Clethrionomys glareolus, in stable and changing rearing environments from father to son. The bank vole is naturally exposed to considerable sources of spatial and temporal environmental variation and male reproductive success is determined by both intra- (male-male competition) and inter- (females prefer to mate with dominant males) sexual selection. Significant G x E for male dominance was found with crossing reaction norms. Plasma testosterone level (T), rather than condition, determined a male's dominance and T also showed a significant G x E. Dominance showed a considerable plasticity across environments, but was only heritable under stable conditions. We document a negative between-environments correlation of male dominance, suggesting that when the environment changes between father and son, the dominance signal is unreliable to females and sexual selection may be compromised. We discuss how G x E and environmental variation interacting with other mechanisms may preserve the reliability of signals and thus the mechanism of sexual selection itself.

Quantitative measure of sexual selection with respect to the operational sex ratio: a comparison of selection indices.

Despite numerous indices proposed to predict the evolution of mating systems, a unified measure of sexual selection has remained elusive. Three previous studies have compared indices of sexual selection under laboratory conditions. Here, we use a genetic study to compare the most widely used measures of sexual selection in natural populations. We explored the mating and reproductive successes of male and female bank voles, Clethrionomys glareolus, across manipulated operational sex ratios (OSRs) by genotyping all adult and pup bank voles on 13 islands using six microsatellite loci. We used Bateman's principles (Is and I and Bateman gradients) and selection coefficients (s' and beta') to evaluate, for the first time, the genetic mating system of bank voles and compared these measures with alternative indices of sexual selection (index of monopolization and Morisita's index) across the OSRs. We found that all the sexual selection indices show significant positive intercorrelations for both males and females, suggesting that Bateman's principles are an accurate and a valid measure of the mating system. The Bateman gradient, in particular, provides information over and above that of other sexual selection indices. Male bank voles show a greater potential for sexual selection than females, and Bateman gradients indicate a polygynandrous mating system. Selection coefficients reveal strong selection gradients on male bank vole plasma testosterone level rather than body size.