Pre- and postcopulatory competition affect testes mass and organization differently in two monophyletic mole-rat species, Georychus capensis and Fukomys damarensis.

Sperm competition results from postcopulatory continuation of male-male competition for paternity. The level of sperm competition is predicted to be highest in species with greater polyandry and weakest in monogamous pairs. Sperm competition levels can be indexed using traits that reflect male investment in fertilization, particularly relative testes mass (RTM). However, the relationship between RTM and levels of sperm competition may also be influenced by precopulatory competition selecting for higher levels of testosterone, also produced by the testes. To test the relationship between RTM and both pre- and postcopulatory male-male competition we compared two bathyergid mole-rat species, the promiscuous Georychus capensis and the monogamous eusocial Fukomys damarensis. The promiscuous species had not only larger RTM, but also a greater proportion of spermatogenic tissue, maximizing germ cell production as well. Conversely, the eusocial species had smaller testes, but a higher proportion of interstitial tissue (which contains the androgenic Leydig cells) and higher levels of testosterone. Consequently, testicular traits as well as testes mass may be under selection, but these are not normally measured. More research is required on relative investment in different testicular traits in relation to both pre- and postcopulatory selection pressures.

High rates of male courtship in a female-ornamented pipefish.

In species with sex-specific signalling traits that appear to be ornamental (i.e. are conspicuous and with no obvious natural selection benefit), the ornamented sex typically initiates courtship and is most active in courtship. Here, we report for the first time courtship displays in the extremely sexually dimorphic, female-ornamented wide-bodied pipefish (Stigmatopora nigra), revealing unexpected behaviours. Females use their sex-specific ornament during courtship displays, as expected, but rarely in female-female interactions. Surprisingly, males initiated 61% of reciprocated courtship bouts and chased females in 17% of the bouts. This chasing behaviour could be a form of male harassment or be indicative of female disinterest in ardent males, either of which was unexpected to be found in this female-ornamented species. Our results highlight the need to study the details of species' behaviours in considering the potential roles of sexual selection and sexual conflict in shaping sexual dimorphism.

Social learning in a nocturnal marsupial: is it a possum-ability?

Social learning can reduce the costs associated with trial-and-error learning. There is speculation that social learning could contribute to trap and bait avoidance in invasive species like the common brushtail possum (Trichosurus vulpecula)-a marsupial for which social learning has not previously been investigated. In large outdoor pens, we presented wild-caught 'demonstrator' possums with puzzle devices containing an attractive food reward; 2 of 8 demonstrators accessed the reward the first night the puzzle was presented and another three succeeded on later nights. Meanwhile, 'observer' possums in adjacent pens watched the demonstrators for five nights and then were given the opportunity to solve the puzzle themselves; 15 of 15 succeeded on their first night (a highly significant improvement). This experiment thus provides strong evidence of social learning by common brushtail possums. Future research should investigate whether information about aversive stimuli (such as traps and toxic baits) can similarly be transmitted between possums by social learning; if so, this could have important implications for possum pest control.

Endangered Australian top predator is frequently exposed to anticoagulant rodenticides.

Anticoagulant rodenticides (ARs) used to control mammalian pest populations cause secondary exposure of predatory species throughout much of the world. It is important to understand the drivers of non-target AR exposure patterns as context for assessing long-term effects and developing effective mitigation for these toxicants. In Australia, however, little is known about exposure and effects of ARs on predators. We detected AR residues in 74% of 50 opportunistically collected carcasses of the Tasmanian wedge-tailed eagle (Aquila audax fleayi), an endangered apex predator. In 22% of birds tested, or 31% of those exposed, liver concentrations of second generation ARs (SGARs) were >0.1 mg/kg ww. Eagles were exposed to flocoumafen, a toxicant only available from agricultural suppliers, at an exceptionally high rate (40% of birds tested). Liver SGAR concentrations were positively associated with the proportion of agricultural habitat and human population density in the area around where each eagle died. The high exposure rate in a species not known to regularly prey upon synanthropic rodents supports the hypothesis that apex predators are vulnerable to SGARs. Our results indicate that AR exposure constitutes a previously unrecognized threat to Tasmanian wedge-tailed eagles and highlight the importance of efforts to address non-target AR exposure in Australia.

High Frequency of Lead Exposure in the Population of an Endangered Australian Top Predator, the Tasmanian Wedge-Tailed Eagle (Aquila audax fleayi).

Lead poisoning, mainly through incidental ingestion of lead ammunition in carcasses, is a threat to scavenging and predatory bird species worldwide. In Australia, shooting for animal control is widespread, and a range of native scavenging species are susceptible to lead exposure. However, the prevalence of lead exposure in Australia's scavenging and predatory birds is largely unknown. We evaluated the degree to which the Tasmanian wedge-tailed eagle (Aquila audax fleayi), an endangered Australian raptor and facultative scavenger, showed evidence of lead exposure. We detected lead in 100% of femur and liver tissues of 109 eagle carcasses opportunistically collected throughout Tasmania between 1996 and 2018. Concentrations were elevated in 10% of 106 liver (>6 mg/kg dry wt) and 4% of 108 femur (>10 mg/kg dry wt) samples. We also detected lead in 96% of blood samples taken from 24 live nestlings, with 8% at elevated concentrations (>10 µg/dL). Of the liver samples with elevated lead, 73% had lead207/206 isotope ratios within the published range of lead-based bullets available in Tasmania. These first comprehensive data on lead exposure of an Australian raptor are comparable to those for raptor studies elsewhere that identify lead-based ammunition exposure as a conservation threat. Our findings highlight the importance of further research and efforts to address lead contamination throughout the Tasmanian ecosystem and in other Australian regions. Environ Toxicol Chem 2021;40:219-230. © 2020 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC. This article has been contributed to by US Government employees and their work is in the public domain in the USA.

Infectious disease and sickness behaviour: tumour progression affects interaction patterns and social network structure in wild Tasmanian devils.

Infectious diseases, including transmissible cancers, can have a broad range of impacts on host behaviour, particularly in the latter stages of disease progression. However, the difficulty of early diagnoses makes the study of behavioural influences of disease in wild animals a challenging task. Tasmanian devils (Sarcophilus harrisii) are affected by a transmissible cancer, devil facial tumour disease (DFTD), in which tumours are externally visible as they progress. Using telemetry and mark-recapture datasets, we quantify the impacts of cancer progression on the behaviour of wild devils by assessing how interaction patterns within the social network of a population change with increasing tumour load. The progression of DFTD negatively influences devils' likelihood of interaction within their network. Infected devils were more active within their network late in the mating season, a pattern with repercussions for DFTD transmission. Our study provides a rare opportunity to quantify and understand the behavioural feedbacks of disease in wildlife and how they may affect transmission and population dynamics in general.

Sexual conflict in action: An antagonistic relationship between maternal and paternal sex allocation in the tammar wallaby, Notamacropus eugenii.

Sex ratio biases are often inconsistent, both among and within species and populations. While some of these inconsistencies may be due to experimental design, much of the variation remains inexplicable. Recent research suggests that an exclusive focus on mothers may account for some of the inconsistency, with an increasing number of studies showing variation in sperm sex ratios and seminal fluids. Using fluorescent in-situ hybridization, we show a significant population-level Y-chromosome bias in the spermatozoa of wild tammar wallabies, but with significant intraindividual variation between males. We also show a population-level birth sex ratio trend in the same direction toward male offspring, but a weaning sex ratio that is significantly female-biased, indicating that males are disproportionately lost during lactation. We hypothesize that sexual conflict between parents may cause mothers to adjust offspring sex ratios after birth, through abandonment of male pouch young and reactivation of diapaused embryos. Further research is required in a captive, controlled setting to understand what is driving and mechanistically controlling sperm sex ratio and offspring sex ratio biases and to understand the sexually antagonistic relationship between mothers and fathers over offspring sex. These results extend beyond sex allocation, as they question studies of population processes that assume equal input of sex chromosomes from fathers, and will also assist with future reproduction studies for management and conservation of marsupials.

Biologically meaningful scents: a framework for understanding predator-prey research across disciplines.

Fear of predation is a universal motivator. Because predators hunt using stealth and surprise, there is a widespread ability among prey to assess risk from chemical information - scents - in their environment. Consequently, scents often act as particularly strong modulators of memory and emotions. Recent advances in ecological research and analytical technology are leading to novel ways to use this chemical information to create effective attractants, repellents and anti-anxiolytic compounds for wildlife managers, conservation biologists and health practitioners. However, there is extensive variation in the design, results, and interpretation of studies of olfactory-based risk discrimination. To understand the highly variable literature in this area, we adopt a multi-disciplinary approach and synthesize the latest findings from neurobiology, chemical ecology, and ethology to propose a contemporary framework that accounts for such disparate factors as the time-limited stability of chemicals, highly canalized mechanisms that influence prey responses, and the context within which these scents are detected (e.g. availability of alternative resources, perceived shelter, and ambient physical parameters). This framework helps to account for the wide range of reported responses by prey to predator scents, and explains, paradoxically, how the same individual predator scent can be interpreted as either safe or dangerous to a prey animal depending on how, when and where the cue was deposited. We provide a hypothetical example to illustrate the most common factors that influence how a predator scent (from dingoes, Canis dingo) may both attract and repel the same target organism (kangaroos, Macropus spp.). This framework identifies the catalysts that enable dynamic scents, odours or odorants to be used as attractants as well as deterrents. Because effective scent tools often relate to traumatic memories (fear and/or anxiety) that cause future avoidance, this information may also guide the development of appeasement, enrichment and anti-anxiolytic compounds, and help explain the observed variation in post-traumatic-related behaviours (including post-traumatic stress disorder, PTSD) among diverse terrestrial taxa, including humans.

Developmental sexual dimorphism and the evolution of mechanisms for adjustment of sex ratios in mammals.

Sex allocation theory predicts biased offspring sex ratios in relation to local conditions if they would maximize parental lifetime reproductive return. In mammals, the extent of the birth sex bias is often unpredictable and inconsistent, leading some to question its evolutionary significance. For facultative adjustment of sex ratios to occur, males and females would need to be detectably different from an early developmental stage, but classic sexual dimorphism arises from hormonal influences after gonadal development. Recent advances in our understanding of early, pregonadal sexual dimorphism, however, indicate high levels of dimorphism in gene expression, caused by chromosomal rather than hormonal differences. Here, we discuss how such dimorphism would interact with and link previously hypothesized mechanisms for sex-ratio adjustment. These differences between males and females are sufficient for offspring sex both to be detectable to parents and to provide selectable cues for biasing sex ratios from the earliest stages. We suggest ways in which future research could use the advances in our understanding of sexually dimorphic developmental physiology to test the evolutionary significance of sex allocation in mammals. Such an approach would advance our understanding of sex allocation and could be applied to other taxa.

Spotted hyaena space use in relation to human infrastructure inside a protected area.

Increasing human population growth has led to elevated levels of human-carnivore conflict. However, some carnivore populations have adapted to urban environments and the resources they supply. Such associations may influence carnivore ecology, behaviour and life-history. Pockets of urbanisation sometimes occur within protected areas, so that anthropogenic influences on carnivore biology are not necessarily confined to unprotected areas. In this study we evaluated associations between human infrastructure and related activity and space use of spotted hyaenas within one of the largest protected areas in South Africa, the Kruger National Park. Home range size was smaller for the dominant female of a clan living in close proximity to humans than that of the dominant female of a clan without direct access to human infrastructure. The home range including human infrastructure was also used less evenly during the night, presumably when the animals were active. Within this home range, a village area was preferred during the night, when the least modified areas within the village were preferred and administration and highly modified areas were avoided. During the day, however, there were no preference or avoidance of the village area, but all habitats except unmodified habitats within the village area were avoided. We suggest that human infrastructure and associated activity influenced hyaena space use, primarily through alterations in the spatial distribution of food. However, these effects may have been indirectly caused by habitat modification that generated favourable hunting habitat rather than a direct effect caused by access to human food such as garbage. Because of the often pivotal effects of apex predators in terrestrial ecosystems, we encourage further work aimed to quantify how human presence influences large carnivores and associated ecosystem processes within protected areas.

Devil declines and catastrophic cascades: is mesopredator release of feral cats inhibiting recovery of the eastern quoll?

The eastern quoll (Dasyurus viverrinus) is a medium-sized Australian marsupial carnivore that has recently undergone a rapid and severe population decline over the 10 years to 2009, with no sign of recovery. This decline has been linked to a period of unfavourable weather, but subsequent improved weather conditions have not been matched by quoll recovery. A recent study suggested another mechanism: that declines in Tasmanian devil (Sarcophilus harrisii) populations, due to the spread of the fatal Devil Facial Tumour Disease, have released feral cats (Felis catus) from competitive suppression, with eastern quoll declines linked to a subsequent increase in cat sightings. Yet current evidence of intraguild suppression among devils, cats and quolls is scant and equivocal. We therefore assessed the influences of top-down effects on abundance and activity patterns among devils, feral cats and eastern quolls. Between 2011 and 2013, we monitored four carnivore populations using longitudinal trapping and camera surveys, and performed camera surveys at 12 additional sites throughout the eastern quoll's range. We did not find evidence of a negative relationship between devil and cat abundance, nor of higher cat abundance in areas where devil populations had declined the longest. Cats did not appear to avoid devils spatially; however, there was evidence of temporal separation of cat and devil activity, with reduced separation and increasing nocturnal activity observed in areas where devils had declined the longest. Cats and quolls used the same areas, and there was no evidence that cat and quoll abundances were negatively related. Temporal overlap in observed cat and quoll activity was higher in summer than in winter, but this seasonal difference was unrelated to devil declines. We suggest that cats did not cause the recent quoll decline, but that predation of juvenile quolls by cats could be inhibiting low density quoll populations from recovering their former abundance through a 'predator pit' effect following weather-induced decline. Predation intensity could increase further should cats become increasingly nocturnal in response to devil declines.

Non-invasive assessment of adrenocortical function in captive Nile crocodiles (Crocodylus niloticus).

The occurrence of stress-inducing factors in captive crocodilians is a concern, since chronic stress can negatively affect animal health and reproduction, and hence production. Monitoring stress in wild crocodiles could also be beneficial for assessing the state of health in populations which are potentially threatened by environmental pollution. In both cases, a non-invasive approach to assess adrenocortical function as a measure of stress would be preferable, as animals are not disturbed during sample collection, and therefore sampling is feedback-free. So far, however, such a non-invasive method has not been established for any crocodilian species. As an initial step, we therefore examined the suitability of two enzyme-immunoassays, detecting faecal glucocorticoid metabolites (FGMs) with a 11ß,21-diol-20-one and 5ß-3α-ol-11-one structure, respectively, for monitoring stress-related physiological responses in captive Nile crocodiles (Crocodylus niloticus). An adrenocorticotropic hormone (ACTH) challenge was performed on 10 sub-adult crocodiles, resulting in an overall increase in serum corticosterone levels of 272% above the pre-injection levels 5h post-injection. Saline-treated control animals (n=8) showed an overall increase of 156% in serum corticosterone levels 5h post-administration. Faecal samples pre- and post-injection could be obtained from three of the six individually housed crocodiles, resulting in FGM concentrations 136-380% above pre-injection levels, always detected in the first sample collected post-treatment (7-15 days post-injection). FGM concentrations seem comparatively stable at ambient temperatures for up to 72 h post-defaecation. In conclusion, non-invasive hormone monitoring can be used for assessing adrenocortical function in captive Nile crocodiles based on FGM analysis.

Forgotten fathers: paternal influences on mammalian sex allocation.

Sex allocation research in mammals has focussed almost exclusively on mothers under the assumption that the male contribution is genetically determined during meiosis and therefore not under adaptive control. Although early studies on sperm traits suggested that sex ratios were at parity, technological advances have made analysis more reliable and cheaper. Subsequently, more studies have shown variation in the production of X-/Y-chromosome-bearing spermatozoa. We review the evidence that the X/Y ratio varies between and within individuals in mammals, and test whether there are consistent predictors of these sex ratio biases and finally discuss their implications for future studies on sex allocation.

Removal experiments indicate that subordinate stallions are not helpers.

Relationships between males defending the same harem are described as cooperative or competitive and explained by mutualism, reciprocal altruism, and reproductive concessions or limited control between unequal contestants. These alternate hypotheses can be tested by removing males from harems. Some feral horse (Equus caballus) harems are defended by more than one and up to five stallions. We temporarily removed the subordinate stallion from two of six multi-stallion bands (a harem and its stallions) for three weeks during the breeding season. We monitored harems for changes in composition, and measured and compared rates of (i) intra- and inter-band stallion aggression, (ii) stallion aggression towards mares, and (iii) stallion-mare proximity before subordinate stallion removal and after his return with rates during his absence. Harems were successfully defended during the subordinate's absence and stallion-mare aggression was substantially reduced. Dominant stallions did not require assistance in harem defence, and heightened harassment of mares is directly attributable to subordinate stallion's residence, not characteristics of the dominant stallion or mares. Cooperative hypotheses do not appear to explain multi-stallion bands but the experiments in this study should be replicated further. The limited control hypothesis (e.g., mate-parasitism) appears better supported but we outline its limitations too.

Is publication rate an equal opportunity metric?

Publication quantity is frequently used as a ranking metric for employment, promotion, and grant success, and is considered an unbiased metric for comparing applicants. However, research suggests that women publish fewer papers, such that the measure may not be equitable. We suggest reasons for the disparity, and potential future remedies. Publication quality and impact provide more equitable metrics of research performance and should be stressed above publication quantity.

Validation of noninvasive monitoring of adrenocortical endocrine activity in ground-feeding aardwolves (Proteles cristata): exemplifying the influence of consumption of inorganic material for fecal steroid analysis.

Biologically inert material in feces may confound interpretations of noninvasive fecal endocrine data, because it may induce variance related to differences in foraging behavior rather than to differences in endocrine activity. We evaluated two different enzyme immunoassays (EIAs) for the noninvasive evaluation of adrenocortical activity in ground-feeding aardwolves (Proteles cristata) and tested the influence of soil content in aardwolf feces on the interpretation of fecal glucocorticoid metabolite data. Using adrenocorticotropic hormone (ACTH) challenges for validation, we successfully identified a cortisol EIA suitable for assessing adrenocortical activity in aardwolves. An alternatively tested 11-oxoetiocholanolone EIA failed to detect a biologically relevant signal after ACTH administration. Although the proportion of inorganic content in aardwolf feces did not alter qualitative conclusions from the endocrine data, the data related to mass of organic content had a larger amount of variance attributed to relevant biological contrasts and a lower amount of variance attributed to individual variation, compared with data related to total dry mass of extracted material. Compared with data expressed as dry mass of extracted material, data expressed as mass of organic content may provide a more refined and statistically powerful measure of endocrine activity in species that ingest large amounts of indigestible material.

Pattern of faecal 20-oxopregnane and oestrogen concentrations during pregnancy in wild plains zebra mares.

Regulative endocrine mechanisms influence the reproductive behaviour and success of mammals, but they have been studied predominantly in domestic and captive animals. The study aims at describing the pattern of faecal 20-oxopregnane and oestrogen concentrations during pregnancy in wild plains zebra Equus quagga chapmani. Data were collected during wet and dry seasons 2007-2009. Enzyme Immunoassays were used to determine 20-oxopregnane and oestrogen concentrations in faecal samples (n=74) collected from individual mares (n=32) whose dates of foaling were known through long-term monitoring. Hormonal profiles were described with a General Additive Model (GAM: Hormone ∼ Days to Foaling). Faecal 20-oxopregnanes have a complex cycle during pregnancy (GAM, n=70, R(2)=0.616, p<0.001). From -250 days to foaling, faecal 20-oxopregnane concentrations were above the baseline levels found in non-pregnant mares, peaking in the last 50 days. Faecal oestrogen levels showed a clear peak in mid-pregnancy (GAM, n=62, R(2)=0.539, p<0.001). The sex of the foetus and season had no detectable effect on hormone concentrations during pregnancy. High levels (>200 ng/g DW) of faecal 20-oxopregnanes associated with high (>160 ng/g DW) faecal oestrogen levels indicate mid-pregnancy in c.90% of cases (16/17). High faecal 20-oxopregnanes (>200 ng/g DW) and low faecal oestrogen levels (<160 ng/g DW) indicate late pregnancy, again in c.90% of cases. Two faecal samples would allow the stage of pregnancy to be determined with confidence.