Vulpeculin: a novel and abundant lipocalin in the urine of the common brushtail possum, Trichosurus vulpecula.

Lipocalins are a family of secreted proteins. They are capable of binding small lipophilic compounds and have been extensively studied for their role in chemosignalling in rodent urine. Urine of the common brushtail possum (Trichosurus vulpecula) contains a prominent glycoprotein of 20 kDa, expressed in both sexes. We have isolated this protein and determined its primary sequence by mass spectrometry, including the use of metabolic labelling to resolve the leucine/isoleucine isobaric ambiguity. The protein sequence was identified as a lipocalin, and phylogenetic analysis grouped the protein with other marsupial lipocalin sequences in a phylogenetic clade distinct from established cross-species lipocalin sub-families. The pattern of expression in possum urine and the similarity in sequence and structure to other lipocalins suggests this protein may have a role in brushtail possum chemosignalling.

Oxpeckers Help Rhinos Evade Humans.

Evolutionary theory expects social, communicative species to eavesdrop most on other species' alarm calls [e.g., 1, 2] but also that solitary-living species benefit most from eavesdropping [3, 4]. Examples of solitary species responding to the alarm calls of other species, however, are limited and unconvincing [3-5]. The Swahili name for the red-billed oxpecker (Buphagus erythrorynchus) is Askari wa kifaru, the rhinos' guard [6]. Black rhino (Diceros bicornis) are a solitary-living, non-vocal species and are critically endangered through hunting. We searched Hluhluwe-iMfolozi Park, South Africa, for rhinoceros for 27 months with and without the aid of radio telemetry and conducted 86 experimental, unconcealed approaches to 11 rhino, without or with varying numbers of resident oxpecker. Oxpeckers enabled rhinos to evade detection by us in 40% to 50% of encounters. Alarm-calling by oxpeckers significantly improved the rate and distance that rhinos detected our approach from 23% to 100% and 27 ± 6 m to 61 ± 4 m, respectively. Every additional oxpecker improved detection distance by 9 m. Rhinos alerted by oxpeckers' alarm calls never re-oriented in our direction but moved to face downwind. Thus, oxpeckers' calls communicate only threat proximity, not direction, and rhinos assume the hunter is stalking from downwind. We confirm that oxpeckers guard rhinos and the importance of depredation, not sociality, in the evolution of eavesdropping [4, 7]. Conservationists should consider reintroducing oxpeckers to rhino populations, reinstating their anti-human sentinel [8]. VIDEO ABSTRACT.

When all life counts in conservation.

Conservation science involves the collection and analysis of data. These scientific practices emerge from values that shape who and what is counted. Currently, conservation data are filtered through a value system that considers native life the only appropriate subject of conservation concern. We examined how trends in species richness, distribution, and threats change when all wildlife count by adding so-called non-native and feral populations to the International Union for Conservation of Nature Red List and local species richness assessments. We focused on vertebrate populations with founding members taken into and out of Australia by humans (i.e., migrants). We identified 87 immigrant and 47 emigrant vertebrate species. Formal conservation accounts underestimated global ranges by an average of 30% for immigrants and 7% for emigrants; immigrations surpassed extinctions in Australia by 52 species; migrants were disproportionately threatened (33% of immigrants and 29% of emigrants were threatened or decreasing in their native ranges); and incorporating migrant populations into risk assessments reduced global threat statuses for 15 of 18 species. Australian policies defined most immigrants as pests (76%), and conservation was the most commonly stated motivation for targeting these species in killing programs (37% of immigrants). Inclusive biodiversity data open space for dialogue on the ethical and empirical assumptions underlying conservation science.

Cuando Toda la Vida Importa en la Conservación Resumen La ciencia de la conservación involucra la recolección y el análisis de datos. Estas prácticas científicas emergen de los valores que forman quién y qué se cuenta. Actualmente, los datos de conservación son filtrados a través de un sistema de valores que considera a la vida nativa como el único sujeto apropiado para el interés de la conservación. Examinamos cómo cambian las tendencias de riqueza de especies, distribución y amenazas cuando se considera a toda la vida silvestre con la adición de las poblaciones denominadas como no nativas y ferales a la Lista Roja de la Unión Internacional para la Conservación de la Naturaleza y a las evaluaciones de riqueza local de especies. Nos enfocamos en las poblaciones de vertebrados que cuentan con miembros fundadores llevados y extraídos de Australia (es decir, migrantes). Identificamos 87 especies inmigrantes de vertebrados y 47 especies emigrantes. Los informes formales de conservación subestimaron los rangos globales por un promedio del 30% para las especies inmigrantes y del 7% para las especies emigrantes; las inmigraciones rebasaron las extinciones en Australia por 52 especies; las especies migrantes estuvieron amenazadas de manera desproporcionada (33% de las especies inmigrantes y 29% de las especies emigrantes estaban amenazadas o declinando en sus distribuciones nativas); y la incorporación de las poblaciones migrantes a las evaluaciones de riesgo redujeron el estado mundial de amenaza para 15 de las 18 especies. Las políticas australianas definen a la mayoría de las especies inmigrantes como plagas (76%) y se citó a la conservación como la principal motivación para enfocarse en estas especies durante los programas de erradicación (37% de las especies inmigrantes). La información inclusiva de conservación genera un espacio para el diálogo sobre las suposiciones éticas y empíricas subyacentes en la ciencia de la conservación.

Mesoscale movement and recursion behaviors of Namibian black rhinos.

BACKGROUND: Understanding rhino movement behavior, especially their recursive movements, holds significant promise for enhancing rhino conservation efforts, and protecting their habitats and the biodiversity they support. Here we investigate the daily, biweekly, and seasonal recursion behavior of rhinos, to aid conservation applications and increase our foundational knowledge about these important ecosystem engineers. METHODS: Using relocation data from 59 rhinos across northern Namibia and 8 years of sampling efforts, we investigated patterns in 24-h displacement at dawn, dusk, midday, and midnight to examine movement behaviors at an intermediate scale and across daily behavioral modes of foraging and resting. To understand recursion patterns across animals' short and long-term ranges, we built T-LoCoH time use grids to estimate recursive movement by each individual. Comparing these grids to contemporaneous MODIS imagery, we investigated productivity's influence on short-term space use and recursion. Finally, we investigated patterns of recursion within a year's home range, measuring the time to return to the most intensively used patches. RESULTS: Twenty four-hour displacements at dawn were frequently smaller than 24-h displacements at dusk or at midday and midnight resting periods. Recursion analyses demonstrated that short-term recursion was most common in areas of median rather than maximum NDVI values. Investigated across a full year, recursion analysis showed rhinos most frequently returned to areas within 8-21 days, though visits were also seen separated by months likely suggesting seasonality in range use. CONCLUSIONS: Our results indicate that rhinos may frequently stay within the same area of their home ranges for days at a time, and possibly return to the same general area days in a row especially during morning foraging bouts. Recursion across larger time scales is also evident, and likely a contributing mechanism for maintaining open landscapes and browsing lawns of the savanna.

Author Correction: Single compounds elicit complex behavioural responses in wild, free-ranging rats.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has not been fixed in the paper.

Single compounds elicit complex behavioural responses in wild, free-ranging rats.

There is mounting evidence that single compounds can act as signals and cues for mammals and that when presented at their optimal concentration they can elicit behavioural responses that replicate those recorded for complex mixtures like gland secretions and foods. We designed a rapid bioassay to present nine compounds that we had previously identified in foods, each at seven different concentrations (63 treatments), to wild, free-ranging rats and scored each treatment for attraction and three behavioural responses. Nine treatments (taken from five compounds) statistically outperformed the current standard rat attractant, peanut butter. Attraction to treatments was highest at the two lowest concentrations (0.1 and 0.01 µg g-1) and a statistically significant relationship of increasing attraction with decreasing treatment concentration was identified. Our study identified five compounds not previously associated with behavioural responses by rats that elicit equivalent or more intense behavioural responses than those obtained with peanut butter. Moreover, attraction to treatments was driven by a concentration-dependent relationship not previously reported. This is the first study to identify isopentanol, 1-hexanol, acetoin, isobutyl acetate and 2-methylbutyl acetate as possible semiochemicals/cues for rats. More broadly, our findings provide important guidance to researchers in the ongoing search for mammalian semiochemicals and cues.

Effects of age and sex ratios on offspring recruitment rates in translocated black rhinoceros.

Success of animal translocations depends on improving postrelease demographic rates toward establishment and subsequent growth of released populations. Short-term metrics for evaluating translocation success and its drivers, like postrelease survival and fecundity, are unlikely to represent longer-term outcomes. We used information theory to investigate 25 years of data on black rhinoceros (Diceros bicornis) translocations. We used the offspring recruitment rate (ORR) of translocated females-a metric integrating survival, fecundity, and offspring recruitment at sexual maturity-to detect determinants of success. Our unambiguously best model (AICω = 0.986) predicted that ORR increases with female age at release as a function of lower postrelease adult rhinoceros sex ratio (males:females). Delay of first postrelease reproduction and failure of some females to recruit any calves to sexual maturity most influenced the pattern of ORRs, and the leading causes of recruitment failure were postrelease female death (23% of all females) and failure to calve (24% of surviving females). We recommend translocating older females (≥6 years old) because they do not exhibit the reproductive delay and low ORRs of juveniles (<4 years old) or the higher rates of recruitment failure of juveniles and young adults (4-5.9 years old). Where translocation of juveniles is necessary, they should be released into female-biased populations, where they have higher ORRs. Our study offers the unique advantage of a long-term analysis across a large number of replicate populations-a science-by-management experiment as a proxy for a manipulative experiment, and a rare opportunity, particularly for a large, critically endangered taxon such as the black rhinoceros. Our findings differ from previous recommendations, reinforce the importance of long-term data sets and comprehensive metrics of translocation success, and suggest attention be shifted from ecological to social constraints on population growth and species recovery, particularly when translocating species with polygynous breeding systems.

Standardising Home Range Studies for Improved Management of the Critically Endangered Black Rhinoceros.

Comparisons of recent estimations of home range sizes for the critically endangered black rhinoceros in Hluhluwe-iMfolozi Park (HiP), South Africa, with historical estimates led reports of a substantial (54%) increase, attributed to over-stocking and habitat deterioration that has far-reaching implications for rhino conservation. Other reports, however, suggest the increase is more likely an artefact caused by applying various home range estimators to non-standardised datasets. We collected 1939 locations of 25 black rhino over six years (2004-2009) to estimate annual home ranges and evaluate the hypothesis that they have increased in size. A minimum of 30 and 25 locations were required for accurate 95% MCP estimation of home range of adult rhinos, during the dry and wet seasons respectively. Forty and 55 locations were required for adult female and male annual MCP home ranges, respectively, and 30 locations were necessary for estimating 90% bivariate kernel home ranges accurately. Average annual 95% bivariate kernel home ranges were 20.4 ± 1.2 km(2), 53 ± 1.9% larger than 95% MCP ranges (9.8 km(2) ± 0.9). When home range techniques used during the late-1960s in HiP were applied to our dataset, estimates were similar, indicating that ranges have not changed substantially in 50 years. Inaccurate, non-standardised, home range estimates and their comparison have the potential to mislead black rhino population management. We recommend that more care be taken to collect adequate numbers of rhino locations within standardized time periods (i.e., season or year) and that the comparison of home ranges estimated using dissimilar procedures be avoided. Home range studies of black rhino have been data deficient and procedurally inconsistent. Standardisation of methods is required.

Reducing sampling error in faecal egg counts from black rhinoceros (Diceros bicornis).

Faecal egg counts (FECs) are commonly used for the non-invasive assessment of parasite load within hosts. Sources of error, however, have been identified in laboratory techniques and sample storage. Here we focus on sampling error. We test whether a delay in sample collection can affect FECs, and estimate the number of samples needed to reliably assess mean parasite abundance within a host population. Two commonly found parasite eggs in black rhinoceros (Diceros bicornis) dung, strongyle-type nematodes and Anoplocephala gigantea, were used. We find that collection of dung from the centre of faecal boluses up to six hours after defecation does not affect FECs. More than nine samples were needed to greatly improve confidence intervals of the estimated mean parasite abundance within a host population. These results should improve the cost-effectiveness and efficiency of sampling regimes, and support the usefulness of FECs when used for the non-invasive assessment of parasite abundance in black rhinoceros populations.

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.

Behavior and Characteristics of Sap-Feeding North Island kaka (Nestor meridionalis septentrionalis) in Wellington, New Zealand.

The North Island kaka (Nestor meridionalis septentrionalis), a threatened New Zealand native parrot, was successfully reintroduced to an urban sanctuary in Wellington, New Zealand. Conflict has recently begun to emerge with Wellington City residents due to tree damage caused by kaka sap foraging. Little is known about sap foraging behavior of kaka, and this study aimed to gain a greater understanding of this behavior, and to test hypotheses that sap feeding is predominantly a female activity and that one technique, forming transverse gouges through bark, may be restricted to adult kaka. We used instantaneous scan sampling to record the behavior of kaka during 25 60-100 minute observation periods at Anderson Park, Wellington Botanic Garden, and during 13 opportunistic observations of sap feeding kaka in Wellington City. Forty-one observations of sap feeding were made of 21 individually-identified birds. Sap feeding birds were predominantly young and, based on estimated sex, females were no more likely to sap feed than males (exact binomial test p = 0.868). Twenty of the 21 identified sap feeding kaka utilized supplementary feeding stations at Zealandia-Karori Wildlife Sanctuary. Kaka were observed defending sap feeding sites from tui (Prosthemadera novaeseelandiae) and conspecifics. Sap appears to be an important resource for kaka across sexes and life stages, and provision of supplementary food is unlikely to reduce sap feeding and tree damage in Wellington City.

Translocations as experiments in the ecological resilience of an asocial mega-herbivore.

Species translocations are remarkable experiments in evolutionary ecology, and increasingly critical to biodiversity conservation. Elaborate socio-ecological hypotheses for translocation success, based on theoretical fitness relationships, are untested and lead to complex uncertainty rather than parsimonious solutions. We used an extraordinary 89 reintroduction and 102 restocking events releasing 682 black rhinoceros (Diceros bicornis) to 81 reserves in southern Africa (1981-2005) to test the influence of interacting socio-ecological and individual characters on post-release survival. We predicted that the socio-ecological context should feature more prominently after restocking than reintroduction because released rhinoceros interact with resident conspecifics. Instead, an interaction between release cohort size and habitat quality explained reintroduction success but only individuals' ages explained restocking outcomes. Achieving translocation success for many species may not be as complicated as theory suggests. Black rhino, and similarly asocial generalist herbivores without substantial predators, are likely to be resilient to ecological challenges and robust candidates for crisis management in a changing world.

Men's preferences for women's breast morphology in New Zealand, Samoa, and Papua New Guinea.

Sexual selection via mate choice may have influenced the evolution of women's breast morphology. We conducted an image-based questionnaire quantifying and comparing the preferences of men from Papua New Guinea (PNG), Samoa, and New Zealand (NZ) for images of women's breast size, breast symmetry, areola size, and areolar pigmentation. Results showed that men from PNG preferred larger breasts to a greater extent than men from Samoa and NZ, providing some support for the hypothesis that men from subsistence living cultures have a greater preference for morphological cues indicative of caloric reserves. Symmetrical breasts were most attractive to men in each culture. However, preferences were highest among NZ men, followed by men from Samoa, and were lowest among men from PNG. These results did not support the hypothesis that people living in higher pathogen environments have a greater preference for traits indicative of pathogen resistance and developmental stability. Large areolae were preferred among men from PNG, and to a lesser extent in Samoa, while in NZ men preferred medium-sized areolae. Thus, men's preferences for women's areolar size appear to be highly culturally specific. Darkly pigmented areolae were most attractive to men from Samoa and PNG, whereas men from NZ preferred areolae with medium pigmentation. These findings suggest that areolar pigmentation indicative of sexual maturity is preferred by men rather than lighter pigmentation, which may signal that a woman is in the early years of reproductive maturity. This study highlights the importance of cross-cultural research when testing the role of morphological cues in mate choice.

Eye tracking of men's preferences for female breast size and areola pigmentation.

Sexual selection via male mate choice has often been implicated in the evolution of permanently enlarged breasts in women. While questionnaire studies have shown that men find female breasts visually attractive, there is very little information about how they make such visual judgments. In this study, we used eye-tracking technology to test two hypotheses: (1) that larger breasts should receive the greatest number of visual fixations and longest dwell times, as well as being rated as most attractive; (2) that lightly pigmented areolae, indicative of youth and nubility, should receive most visual attention and be rated as most attractive. Results showed that men rated images with medium-sized or large breasts as significantly more attractive than small breasts. Images with dark and medium areolar pigmentation were rated as more attractive than images with light areolae. However, variations in breast size had no significant effect on eye-tracking measures (initial visual fixations, number of fixations, and dwell times). The majority of initial fixations during eye-tracking tests were on the areolae. However, areolar pigmentation did not affect measures of visual attention. While these results demonstrate that cues indicative of female sexual maturity (large breasts and dark areolae) are more attractive to men, patterns of eye movements did not differ based on breast size or areolar pigmentation. We conclude that areolar pigmentation, as well as breast size, plays a significant role in men's judgments of female attractiveness. However, fine-grained measures of men's visual attention to these morphological traits do not correlate, in a simplistic way, with their attractiveness judgments.

Eye-tracking of men's preferences for waist-to-hip ratio and breast size of women.

Studies of human physical traits and mate preferences often use questionnaires asking participants to rate the attractiveness of images. Female waist-to-hip ratio (WHR), breast size, and facial appearance have all been implicated in assessments by men of female attractiveness. However, very little is known about how men make fine-grained visual assessments of such images. We used eye-tracking techniques to measure the numbers of visual fixations, dwell times, and initial fixations made by men who viewed front-posed photographs of the same woman, computer-morphed so as to differ in her WHR (0.7 or 0.9) and breast size (small, medium, or large). Men also rated these images for attractiveness. Results showed that the initial visual fixation (occurring within 200 ms from the start of each 5 s test) involved either the breasts or the waist. Both these body areas received more first fixations than the face or the lower body (pubic area and legs). Men looked more often and for longer at the breasts, irrespective of the WHR of the images. However, men rated images with an hourglass shape and a slim waist (0.7 WHR) as most attractive, irrespective of breast size. These results provide quantitative data on eye movements that occur during male judgments of the attractiveness of female images, and indicate that assessments of the female hourglass figure probably occur very rapidly.

Male preferences for female waist-to-hip ratio and body mass index in the highlands of Papua New Guinea.

One hundred men, living in three villages in a remote region of the Eastern Highlands of Papua New Guinea were asked to judge the attractiveness of photographs of women who had undergone micrograft surgery to reduce their waist-to-hip ratios (WHRs). Micrograft surgery involves harvesting adipose tissue from the waist and reshaping the buttocks to produce a low WHR and an "hourglass" female figure. Men consistently chose postoperative photographs as being more attractive than preoperative photographs of the same women. Some women gained, and some lost weight, postoperatively, with resultant changes in body mass index (BMI). However, changes in BMI were not related to men's judgments of attractiveness. These results show that the hourglass female figure is rated as attractive by men living in a remote, indigenous community, and that when controlling for BMI, WHR plays a crucial role in their attractiveness judgments.

Social bonds between unrelated females increase reproductive success in feral horses.

In many mammals, females form close social bonds with members of their group, usually between kin. Studies of social bonds and their fitness benefits have not been investigated outside primates, and are confounded by the relatedness between individuals in primate groups. Bonds may arise from kin selection and inclusive fitness rather than through direct benefits of association. However, female equids live in long-term social groups with unrelated members. We present 4 years of behavioral data, which demonstrate that social integration between unrelated females increases both foal birth rates and survival, independent of maternal habitat quality, social group type, dominance status, and age. Also, we show that such social integration reduces harassment by males. Consequently, social integration has strong direct fitness consequences between nonrelatives, suggesting that social bonds can evolve based on these direct benefits alone. Our results support recent studies highlighting the importance of direct benefits in maintaining cooperative behavior, while controlling for the confounding influence of kinship.

Extreme sex ratio variation in relation to change in condition around conception.

Adaptive theory predicts that mothers would be advantaged by adjusting the sex ratio of their offspring in relation to their offspring's future reproductive success. Studies investigating sex ratio variation in mammals have produced notoriously inconsistent results, although recent studies suggest more consistency if sex ratio variation is related to maternal condition at conception, potentially mediated by changes in circulating glucose level. Consequently, we hypothesized that change in condition might better predict sex ratio variation than condition per se. Here, we investigate sex ratio variation in feral horses (Equus caballus), where sex ratio variation was previously shown to be related to maternal condition at conception. We used condition measures before and after conception to measure the change in condition around conception in individual mothers. The relationship with sex ratio was substantially more extreme than previously reported: 3% of females losing condition gave birth to a son, whereas 80% of those females that were gaining condition gave birth to a son. Change in condition is more predictive of sex ratio than actual condition, supporting previous studies, and shows the most extreme variation in mammals ever reported.