Horse racing is a multi-million pound industry, in which genetic information is increasingly used to optimize breeding programmes. To maximize the probability of producing a successful offspring, the owner of a mare should mate her with a high-quality stallion. However, stallions with big reputations command higher stud fees and paying these is only a sensible strategy if, (i) there is a genetic variation for success on the racecourse and (ii) stud fees are an honest signal of a stallion's genetic quality. Using data on thoroughbred racehorses, and lifetime earnings from prize money (LE) as a measure of success, we performed quantitative genetic analyses within an animal model framework to test these two conditions. Although LE is heritable (VA=0.299±0.108, Pr=0.002), there is no genetic variance for stud fee and the genetic correlation between traits is therefore zero. This result is supported by an absence of any relationship between stud fees for currently active stallions and the predicted LE for their (hypothetical) offspring. Thus, while there are good genes to be bought, a stallion's fees are not an honest signal of his genetic quality and are a poor predictor of a foal's prize winning potential.
thoroughbred; Equus caballus; heritability; genetic correlation
An individual's sex depends upon its genes (genotypic sex determination or GSD) in birds and mammals, but reptiles are more complex: some species have GSD whereas in others, nest temperatures determine offspring sex (temperature-dependent sex determination). Previous studies suggested that montane scincid lizards (Bassiana duperreyi, Scincidae) possess both of these systems simultaneously: offspring sex is determined by heteromorphic sex chromosomes (XX–XY system) in most natural nests, but sex ratio shifts suggest that temperatures override chromosomal sex in cool nests to generate phenotypically male offspring even from XX eggs. We now provide direct evidence that incubation temperatures can sex-reverse genotypically female offspring, using a DNA sex marker. Application of exogenous hormone to eggs also can sex-reverse offspring (oestradiol application produces XY as well as XX females). In conjunction with recent work on a distantly related lizard taxon, our study challenges the notion of a fundamental dichotomy between genetic and thermally determined sex determination, and hence the validity of current classification schemes for sex-determining systems in reptiles.
discordant sex; reptile; sex chromosomes; temperature-dependent sex determination
While we can usually understand the impacts of invasive species on recipient communities, invasion biology lacks methodologies that are potentially more predictive. Such tools should ideally be straightforward and widely applicable. Here, we explore an approach that compares the functional responses (FRs) of invader and native amphipod crustaceans. Dikerogammarus villosus is a Ponto-Caspian amphipod currently invading Europe and poised to invade North America. Compared with other amphipods that it actively replaces in freshwaters, D. villosus exhibited significantly greater predation, consuming significantly more prey with a higher type II FR. This corroborates the known dramatic field impacts of D. villosus on invaded communities. In another species, FRs were nearly identical in invasive and native ranges. We thus propose that if FRs of other taxa and trophic groups follow such general patterns, this methodology has potential in predicting future invasive species impacts.
amphipod; functional response; invasive species; predation; prediction
In most taxa, owners win fights when defending a territory against intruders. We calculated effect sizes for four factors that potentially contribute to an ‘owner advantage’. We studied male fiddler crabs Uca mjoebergi, where owners won 92% of natural fights. Owners were not more successful because they were inherently better fighters (r=0.02). There was a small effect (r=0.18) of the owner's knowledge of territory quality (food availability) and a medium effect (r=0.29) of his having established relations with neighbours (duration of active tenure), but neither was statistically significant. There was, however, a significant effect due to the mechanical advantage the owner gained through access to the burrow during fights (r=0.48, p<0.005).
fighting; ownership; territoriality
Changes in the relative abundance of sympatric carnivores can have far-reaching ecological consequences, including the precipitation of trophic cascades and species declines. While such observations are compelling, experimental evaluations of interactions among carnivores remain scarce and are both logistically and ethically challenging. Carnivores are nonetheless a particular focus of management practices owing to their roles as predators of livestock and as vectors and reservoirs of zoonotic diseases. Here, we provide evidence from a replicated and controlled experiment that culling Eurasian badgers Meles meles for disease control was associated with increases in red fox Vulpes vulpes densities of 1.6–2.3 foxes km−2. This unique experiment demonstrates the importance of intraguild relations in determining species abundance and of assessing the wider consequences of intervention in predator populations.
badger; culling; ecological processes; mesopredator release; predator removal; tuberculosis
Exceptional braincase preservation in some Devonian placoderm fishes permits interpretation of muscles and cranial nerves controlling eye movement. Placoderms are the only jawed vertebrates with anterior/posterior obliques as in the jawless lamprey, but with the same function as the superior/inferior obliques of other gnathostomes. Evidence of up to seven extraocular muscles suggests that this may be the primitive number for jawed vertebrates. Two muscles innervated by cranial nerve 6 suggest homologies with lampreys and tetrapods. If the extra muscle acquired by gnathostomes was the internal rectus, Devonian fossils show that it had a similar insertion above and behind the eyestalk in both placoderms and basal osteichthyans.
eye muscles; jaw evolution; gnathostome phylogeny
Regular scale patterning, restricted to the caudalmost tail and organized into two opposing rows on each side of the tail, is observed in few chondrichthyans. These evenly spaced scales, in dorsal and ventral rows, develop in an iterative sequence from the caudal tip, either side of the notochord. They are subsequently lost as a scattered pattern of placoid scales develops on the body and fins. An identical organized pattern is observed in tail scales of Scyliorhinus canicula (catshark), where the expression of sonic hedgehog signal is restricted to the epithelium of developing scales and remains localized to the scale pocket. Regulation of iterative scale position by sonic hedgehog is deeply conserved in vertebrate phylogeny.
These scales also reveal an archaic histological structure of a dentine type found in the oldest known shark scales from the Ordovician and Silurian. This combination of regulated pattern and ancient dentine occurs only in the tail, representing the primary scalation. Scattered body scales in elasmobranchs such as S. canicula originate secondarily from differently regulated development, one with typical orthodentine around a central pulp cavity. These observations emphasize the modular nature of chondrichthyan scale development and illustrate previously undetected variation as an atavism in extant chondrichthyan dentine.
Scyliorhinus; chondrichthyan evolution; scale development; dentine structure
Among anuran amphibians, only two species, Odorrana tormota and Huia cavitympanum, are known to possess recessed tympanic membranes. Odorrana tormota is the first non-mammalian vertebrate demonstrated to communicate with ultrasonic frequencies (above 20 kHz), and the frogs' sunken tympana are hypothesized to play a key role in their high-frequency hearing sensitivity. Here we present the first data on the vocalizations of H. cavitympanum. We found that this species emits extraordinarily high-frequency calls, a portion of which are comprised entirely of ultrasound. This represents the first documentation of an anuran species producing purely ultrasonic signals. In addition, the vocal repertoire of H. cavitympanum is highly variable in frequency modulation pattern and spectral composition. The frogs' use of vocal signals with a wide range of dominant frequencies may be a strategy to maximize acoustic energy transmission to both nearby and distant receivers. The convergence of these species' call characteristics should stimulate additional, phylogenetically based studies of other lower vertebrates to provide new insight into the mechanistic and evolutionary foundations of high-frequency hearing in all vertebrate forms.
Odorrana tormota; Huia cavitympanum; ultrasonic communication; ultrasound; convergence
In cooperatively breeding species, helpers can alleviate reproductive constraints by assuming the role of primary carers to first-born young, liberating breeders to invest in subsequent broods. However, evidence on how first-born young are transferred to helpers is currently lacking. We propose that breeder–offspring aggression might facilitate inter-brood division and test this idea using data from a wild population of cooperatively breeding pied babblers (Turdoides bicolor). After second-brood young hatch, breeders become increasingly aggressive to first-brood fledglings and attack them when they beg for food. After an attack, fledglings reduce begging. Helpers are much less aggressive to begging fledglings and fledglings subsequently tend to target helpers, rather than breeders, when begging for food. In this way, first-born dependent young are transferred to helpers, resulting in a partitioning of tasks among breeders and helpers. Task partitioning in eusocial insects is thought to be determined by the morphological or physiological characteristics of individuals. This complementary study suggests that flexible behavioural strategies may also result in specialized roles in cooperatively breeding vertebrates.
parental aggression; task partitioning; brood division; pied babbler; cooperative; division of labour
The theory of reciprocal altruism offers an explanation for the evolution of altruistic behaviours among unrelated animals. Among primates, grooming is one of the most common altruistic behaviours. Primates have been suggested to exchange grooming both for itself and for rank-related benefits. While previous meta-analyses have shown that they direct their grooming up the hierarchy and exchange it for agonistic support, no comprehensive evaluation of grooming reciprocation has been made. Here we report on a meta-analysis of grooming reciprocation among female primates based on 48 social groups belonging to 22 different species and 12 genera. The results of this meta-analysis showed that female primates groom preferentially those group mates that groom them most. To the extent allowed by the availability of kinship data, this result holds true when controlling for maternal kinship. These results, together with previous findings, suggest that primates are indeed able to exchange grooming both for itself and for different rank-related benefits.
reciprocation; altruism; grooming; primates
Pitohui birds from New Guinea have been found to contain a toxin otherwise only found in neotropical poison arrow frogs. Pitohuis have been considered to be monophyletic and thus toxicity is thought to have evolved once in birds. Here, we show that Pitohuis, rather than being a tight-knit group, are polyphyletic and represent several lineages among the corvoid families of passerine birds. This finding demonstrates that the ability to be toxic is widespread among corvoid birds and suggests that additional members of this radiation, comprising more than 700 species, could prove to be toxic. It is postulated that toxic birds ingest the toxin through their insect diet and excrete it through the uropygial gland, from where it is applied to the skin and feathers. Thus, the ability to become toxic is most likely an ancestral condition but variation in diet determines the extent to which toxicity is expressed among corvoid birds. Variability in toxicity levels further suggests that the main function of the toxin is that of a deterrent against ectoparasites and bacterial infection rather than being a defence against predators as initially proposed.
Pitohui; toxicity; systematics; batrachotoxin; phylogeny
The endangered mountain pygmy possum is the only Australian marsupial that hibernates under snow cover. Most of its alpine habitat was burnt by a rare fire in 2003, and habitat loss and disturbance have also occurred owing to ski resort development. Here we show that there has been a rapid loss of genetic variation following habitat loss associated with resort development, but no detectable loss of alleles or decrease in heterozygosity following the fire.
genetic variation; Burramys; endangered; inbreeding
The birth sex ratio of vertebrates with chromosomal sex determination has been shown to respond to environmental variability, such as temperature. However, in humans the few previous studies on environmental temperature and birth sex ratios have produced mixed results. We examined whether reconstructed annual mean temperatures were associated with annual offspring sex ratio at birth in the eighteenth to nineteenth century Sami from northern Finland. We found that warm years correlated with a male-biased sex ratio, whereas a warm previous year skewed sex ratio towards females. The net effect of one degree Celsius increase in mean temperature during these 2 years corresponded to approximately 1% more sons born annually. Although the physiological and ecological mechanisms mediating these effects and their evolutionary consequences on parental fitness remain unknown, our results show that environmental temperature may affect human birth sex ratio.
climatic reconstruction; parental age; time-series analysis
Lateralization is a well-described phenomenon in humans and other vertebrates and there are interesting parallels across a variety of different vertebrate species. However, there are only a few studies of lateralization in invertebrates. In a recent report, we showed lateralization of olfactory learning in the honeybee (Apis mellifera). Here, we investigate lateralization of another sensory modality, vision. By training honeybees on a modified version of a visual proboscis extension reflex task, we find that bees learn a colour stimulus better with their right eye.
lateralization; insects; honeybee vision
Disparate psychrophiles (e.g. glacier ice worms, bacteria, algae and fungi) elevate steady-state intracellular ATP levels as temperatures decline, which has been interpreted as a compensatory mechanism to offset reductions in molecular motion and Gibb's free energy of ATP hydrolysis. In this study, we sought to manipulate steady-state ATP levels in the mesophilic bacterium, Escherichia coli, to investigate the relationship between cold temperature survivability and elevated intracellular ATP. Based on known energetic pathways and feedback loops, we targeted the AMP nucleotidase (amn) gene, which is thought to encode the primary AMP degradative enzyme in prokaryotes. By knocking out amn in wild-type E. coli DY330 cells using recombineering methodology, we generated a mutant (AMNk) that elevated intracellular ATP levels by more than 30% across its viable temperature range. As temperature was lowered, the relative ATP disparity between AMNk and DY330 cells increased to approximately 66% at 10°C, and was approximately 100% after storage at 0°C for 5–7 days. AMNk cells stored at 0°C for 7 days displayed approximately fivefold higher cell viability than wild-type DY330 cells treated in the same manner.
psychrophile; energetics; adenylates; cold tolerance
Animal signals are constrained by the environment in which they are transmitted and the sensory systems of receivers. Detection of movement-based signals is particularly challenging against the background of wind-blown plants. The Australian lizard Amphibolurus muricatus has recently been shown to compensate for greater plant motion by prolonging the introductory tail-flicking component of its movement-based display. Here I demonstrate that such modifications to signal structure are useful because environmental motion lengthens the time lizard receivers take to detect tail flicks. The spatio-temporal properties of animal signals and environmental motion are thus sufficiently similar to make signal detection more difficult. Environmental motion, therefore, must have had an influence on the evolution of movement-based signals and motion detection mechanisms.
movement-based signal; motion vision; lizard; signal evolution
Despite evidence that organismal distributions are shifting in response to recent climatic warming, we have little information on direct links between species' physiology and vulnerability to climate change. We demonstrate a positive relationship between upper thermal tolerance and its acclimatory ability in a well-defined clade of closely related European diving beetles. We predict that species with the lowest tolerance to high temperatures will be most at risk from the adverse effects of future warming, since they have both low absolute thermal tolerance and poor acclimatory ability. Upper thermal tolerance is also positively related to species' geographical range size, meaning that species most at risk are already the most geographically restricted ones, being endemic to Mediterranean mountain systems. Our findings on the relationship between tolerance and acclimatory ability contrast with results from marine animals, suggesting that generalizations regarding thermal tolerance and responses to future rapid climate change may be premature.
thermal tolerance; range size; rarity; climate change; diving beetle; ecophysiology
Wolves from the Great Lakes area were historically decimated due to habitat loss and predator control programmes. Under the protection of the US Endangered Species Act, the population has rebounded to approximately 3000 individuals. We show that the pre-recovery population was dominated by mitochondrial DNA haplotypes from an endemic American wolf referred to here as the Great Lakes wolf. In contrast, the recent population is admixed, and probably derives also from the grey wolf (Canis lupus) of Old World origin and the coyote (Canis latrans). Consequently, the pre-recovery population has not been restored, casting doubt on delisting actions.
hybridization; coyote; introgression; aDNA