Body size increase in insular rodent populations: a role for predators?

Insular mammalian populations living in areas of small size are often characterized by a drastic change in body mass compared to related continental populations or species. Generally, small mammals (less than 100 g) evolve into giant forms while large mammals (up to 100 g) evolve into dwarf forms. These changes, coupled with changes in other life, behavioural, physiological or demographic traits are referred to generally as the insular syndrome. We tested in this study the relative contribution of three factors — area of island, numbers of competitor species and number of predator species — to changes in body size of the woodmouse (Apodemus sylvaticus) in the Western Mediterranean Sea. Our results, based on a comparative analysis using the phylogenetic independent contrasts method, indicate that the increase in body size is related both to the decrease of island size and to the lower number of predator species. A decrease of competitor species does not seem to have an important effect.     

Resource utilization by two insular endemic mammalian carnivores,the island fox and island spotted skunk

We compared resource utilization of two insular endemic mammalian carnivores, the island spotted skunk and island fox, along niche dimensions of space, food, and time on Santa Cruz Island. We predicted that resource use by foxes and skunks would differ along one or more niche dimensions, and that both species would have broader niches or higher densities compared with mainland relatives. Island foxes and island spotted skunks differed to some extent in habitat use, diets, and circadian activity, which may account for their long-term coexistence. Nonetheless, substantial overlap between skunks and foxes in spatial, dietary, and temporal dimensions suggests that competition between the two species does occur. Moreover, competition may be asymmetric, affecting skunks more than foxes. Compared with mainland foxes, island foxes have smaller body size, smaller home range, increased population density, increased diurnal activity, and behavior that is more highly inquisitive and less flightprone all common features of insular faunas. Island skunks, however, apparently have not developed these changes, perhaps due to asymmetric competition with foxes in conjunction with severe ecosystem disturbances caused by feral sheep.     

Insular shifts and trade-offs in life-history traits in pond frogs in the Zhoushan Archipelago, China

Island and mainland populations of animal species often differ strikingly in life-history traits such as clutch size, egg size, total reproductive effort and body size. However, despite widespread recognition of insular shifts in these life-history traits in birds, mammals and reptiles, there have been no reports of such life-history shifts in amphibians. Furthermore, most studies have focused on one specific life-history trait without explicit consideration of coordinated evolution among these intimately linked life-history traits, and thus the relationships among these traits are poorly studied. Here we provide the first evidence of insular shifts and trade-offs in a coordinated suite of life-history traits for an amphibian species, the pond frog Rana nigromaculata . Life-history data were collected from eight islands in the Zhoushan Archipelago and neighboring mainland China. We found consistent, significant shifts in all life-history traits between mainland and island populations. Island populations had smaller clutch sizes, larger egg sizes, larger female body size and invested less in total reproductive effort than mainland populations. Significant negative relationships were found between egg size and clutch size and between egg size and total reproductive effort among frog populations after controlling for the effects of body size. Therefore, decreased reproductive effort and clutch size, larger egg size and body size in pond frogs on islands were selected through trade-offs as an overall life-history strategy. Our findings contribute to the formation of a broad, repeatable ecological generality for insular shifts in life-history traits across a range of terrestrial vertebrate taxa.     

A reserve for feral sheep on Pitt Island,Chatham group,New Zealand

Abstract

A scientific reserve of 200 ha has been created on Pitt Island, Chatham Islands for some 300 sheep from a flock of merino origin which has been feral for about 70 years. The reserve has a history of burning-off, and of grazing by cattle, sheep, and pigs. Transects have been established to follow changes in the vegetation. Although the original sheep were white, about 90% of the feral sheep have pigmented fleeces. Self-shedding of the wool is common in the population. The contribution that feral mammals can make to genetic conservation, and the problems that they pose to other conservation priorities, are discussed.     

Mainland size variation informs predictive models of exceptional insular body size change in rodents

The tendency for island populations of mammalian taxa to diverge in body size from their mainland counterparts consistently in particular directions is both impressive for its regularity and, especially among rodents, troublesome for its exceptions. However, previous studies have largely ignored mainland body size variation, treating size differences of any magnitude as equally noteworthy. Here, we use distributions of mainland population body sizes to identify island populations as ‘extremely’ big or small, and we compare traits of extreme populations and their islands with those of island populations more typical in body size. We find that although insular rodents vary in the directions of body size change, ‘extreme’ populations tend towards gigantism. With classification tree methods, we develop a predictive model, which points to resource limitations as major drivers in the few cases of insular dwarfism. Highly successful in classifying our dataset, our model also successfully predicts change in untested cases.     

Classification tree methods provide a multifactorial approach to predicting insular body size evolution in rodents

Many hypotheses have been proposed to explain size changes in insular mammals, but no single variable suffices to explain the diversity of responses, particularly within Rodentia. Here in a data set on insular rodents, we observe strong consistency in the direction of size change within islands and within species but (outside of Heteromyidae) little consistency at broader taxonomic scales. Using traits of islands and of species in a classification tree analysis, we find the most important factor predicting direction of change to be mainland body mass (large rodents decrease, small ones increase); other variables (island climate, number of rodent species, and area) were significant, although their roles as revealed by the classification tree were context dependent. Ecological interactions appear relatively uninformative, and on any given island, the largest and smallest rodent species converged or diverged in size with equal frequency. Our approach provides a promising framework for continuing examination of insular body size evolution.     

Comparison of life-history characteristics of island and mainland populations of the swamp antechinus

The reproductive ecology of the swamp antechinus Antechinus minimus , a small dasyurid (Dasyuridae) marsupial with obligate male semelparity, was investigated in populations inhabiting the mainland coast and on a nearby offshore island in south-eastern Australia. The size and sex ratios of litters, individual body mass and size, timing of births and female longevity were determined from live-trapped animals. The island population had significantly smaller litter sizes and greater adult body mass in comparison with the mainland population. This is consistent with features of the 'island syndrome', which predicts directional selection for these traits in high-density populations with reduced extrinsic mortality. However, inter-annual variability in litter sizes in the island population suggests that litter size is more responsive to fluctuating local conditions, such as population density, which is likely to affect food availability, rather than directional genetic changes. In contrast with other antechinus species, biased sex ratios were not evident. In addition, large variations of the timing of births were estimated at both sites and these appear to be related to seasonal conditions such as autumn rainfall and female body mass before mating.     

Body size evolution in insular vertebrates: generality of the island rule

Aim My goals here are to (1) assess the generality of the island rule – the graded trend from gigantism in small species to dwarfism in larger species – for mammals and other terrestrial vertebrates on islands and island‐like ecosystems; (2) explore some related patterns of body size variation in insular vertebrates, in particular variation in body size as a function of island area and isolation; (3) offer causal explanations for these patterns; and (4) identify promising areas for future studies on body size evolution in insular vertebrates. Location Oceanic and near‐shore archipelagos, and island‐like ecosystems world‐wide. Methods Body size measurements of insular vertebrates (non‐volant mammals, bats, birds, snakes and turtles) were obtained from the literature, and then regression analyses were conducted to test whether body size of insular populations varies as a function of body size of the species on the mainland (the island rule) and with characteristics of the islands (i.e. island isolation and area). Results The island rule appears to be a general phenomenon both with mammalian orders (and to some degree within families and particular subfamilies) as well as across the species groups studied, including non‐volant mammals, bats, passerine birds, snakes and turtles. In addition, body size of numerous species in these classes of vertebrates varies significantly with island isolation and island area. Main conclusions The patterns observed here – the island rule and the tendency for body size among populations of particular species to vary with characteristics of the islands – are actually distinct and scale‐dependent phenomena. Patterns within archipelagos reflect the influence of island isolation and area on selective pressures (immigration filters, resource limitation, and intra‐ and interspecific interactions) within particular species. These patterns contribute to variation about the general trend referred to as the island rule, not the signal for that more general, large‐scale pattern. The island rule itself is an emergent pattern resulting from a combination of selective forces whose importance and influence on insular populations vary in a predictable manner along a gradient from relatively small to large species. As a result, body size of insular species tends to converge on a size that is optimal, or fundamental, for a particular bau plan and ecological strategy.     

Of mice and mammoths: evaluations of causal explanations for body size evolution in insular mammals

Aim We investigated the hypothesis that the insular body size of mammals results from selective forces whose influence varies with characteristics of the focal islands and the focal species, and with interactions among species (ecological displacement and release). Location Islands world‐wide. Methods We assembled data on the geographic characteristics (area, isolation, maximum elevation, latitude) and climate (annual averages and seasonality of temperature and precipitation) of islands, and on the ecological and morphological characteristics of focal species (number of mammalian competitors and predators, diet, body size of mainland reference populations) that were most relevant to our hypothesis (385 insular populations from 98 species of extant, non‐volant mammals across 248 islands). We used regression tree analyses to examine the hypothesized contextual importance of these factors in explaining variation in the insular body size of mammals. Results The results of regression tree analyses were consistent with predictions based on hypotheses of ecological release (more pronounced changes in body size on islands lacking mammalian competitors or predators), immigrant selection (more pronounced gigantism in small species inhabiting more isolated islands), thermoregulation and endurance during periods of climatic or environmental stress (more pronounced gigantism of small mammals on islands of higher latitudes or on those with colder and more seasonal climates), and resource subsidies (larger body size for mammals that utilize aquatic prey). The results, however, were not consistent with a prediction based on resource limitation and island area; that is, the insular body size of large mammals was not positively correlated with island area. Main conclusions These results support the hypothesis that the body size evolution of insular mammals is influenced by a combination of selective forces whose relative importance and nature of influence are contextual. While there may exist a theoretical optimal body size for mammals in general, the optimum for a particular insular population varies in a predictable manner with characteristics of the islands and the species, and with interactions among species. This study did, however, produce some unanticipated results that merit further study – patterns associated with Bergmann’s rule are amplified on islands, and the body size of small mammals appears to peak at intermediate and not maximum values of latitude and island isolation.     

Insular shifts in body size of rice frogs in the Zhoushan Archipelago, China

1. Differences in body size between mainland and island populations have been reported for reptiles, birds and mammals. Despite widespread recognition of insular shifts in body size in these taxa, there have been no reports of such body size shifts in amphibians. 2. We provide the first evidence of an insular shift in body size for an amphibian species, the rice frog Rana limnocharis. We found significant increases in body size of rice frogs on most sampled islands in the Zhoushan archipelago when compared with neighbouring mainland China. 3. Large body size in rice frogs on islands was significantly related to increased population density, in both breeding and non-breeding seasons. Increases in rice frog density were significantly related to higher resource availability on islands. Increased resource availability on islands has led to higher carrying capacities, which has subsequently facilitated higher densities and individual growth rates, resulting in larger body size in rice frogs. We also suggest that large body size has evolved on islands, as larger individuals are competitively superior under conditions of harsh intraspecific competition at high densities. 4. Increases in body size in rice frogs were not related to several factors that have been implicated previously in insular shifts in body size in other taxa. We found no significant relationships between body size of rice frogs and prey size, number of larger or smaller frog species, island area or distance of islands from the mainland. 5. Our findings contribute to the formation of a broad, repeatable ecological generality for insular shifts in body size across a range of terrestrial vertebrate taxa, and provide support for recent theoretical work concerning the importance of resource availability for insular shifts in body size.     

Primate brains,the ‘island rule’ and the evolution of Homo floresiensis

The taxonomic status of the small bodied hominin, Homo floresiensis, remains controversial. One contentious aspect of the debate concerns the small brain size estimated for specimen LB1 (Liang Bua 1). Based on intraspecific mammalian allometric relationships between brain and body size, it has been argued that the brain of LB1 is too small for its body mass and is therefore likely to be pathological. The relevance and general applicability of these scaling rules has, however, been challenged, and it is not known whether highly encephalized primates adapt to insular habitats in a consistent manner. Here, an analysis of brain and body size evolution in seven extant insular primates reveals that although insular primates follow the ‘island rule’, having consistently reduced body masses compared with their mainland relatives, neither brain mass nor relative brain size follow similar patterns, contrary to expectations that energetic constraints will favour decreased relative brain size. Brain:body scaling relationships previously used to assess the plausibility of dwarfism in H. floresiensis tend to underestimate body masses of insular primates. In contrast, under a number of phylogenetic scenarios, the evolution of brain and body mass in H. floresiensis is consistent with patterns observed in other insular primates.     

Body size of insular carnivores: evidence from the fossil record

Aim Our goals here are to: (1) assess the generality of one aspect of the island rule – the progressive trend towards decrease in size in larger species – for fossil carnivores on islands; (2) offer causal explanations for this pattern and deviations from it – as far as fossil carnivores are concerned; and (3) estimate the speed of this trend. Location Oceanic and oceanic‐like islands world‐wide. Methods Body size estimates of fossil insular carnivores and of their phylogenetically closest mainland relative were obtained from our own data and the published literature. Our dataset consisted of 18 species from nine islands world‐wide. These data were used to test whether the body size of fossil insular carnivores varies as a function of body size of the mainland species in combination with characteristics of the island ecosystem. Results Dwarfism was observed in two canid species. Moderate decrease in body mass was observed in one hyena species. Gigantism was observed in one otter species. Moderate body mass increase was observed in two otter species, one galictine mustelid and perhaps one canid. Negligible or no change in body mass at all was observed in five otter species, three galictine mustelids and one genet. Size changes in teeth do not lag behind in comparison to skeletal elements in the dwarfed canids. The evolutionary speed of dwarfism in a canid lineage is low. Main conclusions Size change in fossil terrestrial insular carnivores was constrained by certain ecological conditions, especially the availability of prey of appropriate body size. When such alternative prey was not available, the carnivores retained their mainland size. The impact of competitive carnivores seems negligible. The case of (semi‐)aquatic carnivores is much less clear. The species that maintained their ancestral body mass may have changed their diet, as is evidenced by their dentition. Among the otters, one case of significant size increase was observed, perhaps best explained as being due to it entering the niche of an obligate aquatic otter. Dwarfism was not observed in otters. The island rule seems to apply to fossil carnivores, but with exceptions. The dependency of the island rule on resource availability is emphasized by the present study.     

‘On being the right size’ – Do aliens follow the rules?

Aim

To assess whether mammalian species introduced onto islands across the globe have evolved to exhibit body size patterns consistent with the ‘island rule,’, and to test an ecological explanation for body size evolution of insular mammals.

Location

Islands worldwide.

Methods

We assembled data on body mass, geographical characteristics (latitude, maximum elevation) and ecological communities (number of mammalian competitors, predators and prey) for 385 introduced populations across 285 islands, comprising 56 species of extant, non‐volant mammals. We used linear regression, ANCOVA and regression tree analyses to test whether introduced populations of mammals exhibit the island rule pattern, whether the degree of body size change increased with time in isolation and whether residual variation about the general trend can be attributed to the geographical and ecological characteristics of the islands.

Results

Introduced populations follow the predicted island rule trend, with body size shifts more pronounced for populations with greater residence times on the islands. Small mammals evolved to larger body sizes in lower latitudes and on islands with limited topographic relief. Consistent with our hypothesis on the ecology of evolution, body size of insular introduced populations was influenced by co‐occurring species of mammalian competitors, predators and prey.

Conclusion

The island rule is a pervasive pattern, exhibited across a broad span of geographical regions, taxa, time periods and, as evidenced here, for introduced as well as native mammals. Time in isolation impacts body size evolution profoundly. Body size shift of introduced mammals was much more pronounced with increasing residence times, yet far less than that exhibited by native, palaeo‐insular mammals (residence times > 10,000 years). Given the antiquity of many species introductions, it appears that much of what we view as the natural character and ecological dynamics of recent insular communities may have been rendered artefacts of ancient colonizations by humans and commensals.     

Sexual performance and production traits in white-faced yearling rams

Poor mating behavior results in increased ram costs, an extended lambing season, and decreased genetic progress from sires with desired production traits. Producers have recognized the importance of ram libido, but constraints regarding time, labor, and facilities generally limit the routine evaluation of ram breeding behavior. Easily measurable production traits would be an usable selection tool if correlated to reproductive performance. To determine the discriminating value of production traits as indicators of reproductive performance, white-faced rams (n = 82; 10–12 months of age) consigned to the University of Wyoming ram test were fed using the Grow-Safe^® feeding system. Rams were categorized based on sexual behavior as low, intermediate or mounting. Body weight gain performance, feed intake and feeding behavior were analyzed. Overall production performance was determined using a calculated index ratio based on body weight gain adjusted for wool characteristics. Sexual behavior classification did not influence the index ratio or feed consumed per day. However, numbers of no-intake feeding episodes (P < 0.01) differed according to ram sexual behavior. Rams expressing sexual mounting behavior visited the feed bunk more often than the intermediate performing rams (P < 0.01) and tended to differ from low sexually performing rams. The numbers of visits to the feed bunk in which feed was consumed did not differ according to ram sexual behavior. Average daily gain was not correlated (r ≤ 0.18; P = 0.1) to sexual behavior. These easily attainable measures of performance did not correlate strongly with any measure of sexual behavior. This study provides evidence that rams that exhibit a high level of sexual behavior most likely investigate their environment more thoroughly, which could facilitate the identification of ewes in estrus.     

Auction price of Texel, Suffolk and German white-headed mutton rams: A genetic-statistical study

In this study, the impact of various traits and effects on auction price of Texel, Suffolk and German white-headed mutton rams was determined. Furthermore, (co)variance components between auction price and performance traits recorded at licensing were estimated. Data from 1988 to 2007 were extracted from the recording database of the Sheep Breeding Organisation in Schleswig-Holstein, Germany. Auction prices of 1133 Texel, 373 Suffolk and 341 German white-headed mutton rams from the year 2003 to 2007 were analysed. General Linear Models and Multiple Regression Analysis were used to identify the impact of traits and effects on the auction price. Estimation of (co)variance components was carried out using multivariate animal model. From the traits recorded at licensing, live weight, followed by type traits of conformation and muscle mass, had the highest impact on the auction price. Also the effects of breed, PrP (Prion Protein) genotype, and owner of the ram influenced the auction price of rams significantly. A moderate genetic contribution of auction price, with favourable genetic correlations to all collected performance traits (average daily gain from birth until licensing, muscle depth, fat depth, conformation, muscle mass and wool quality), was estimated. Genetic correlations were 0.42, 0.34, 0.17, 0.49, 0.39 and 0.23, respectively. The results showed a high influence of live weight of animal on auction price of the rams contrary to very low influence of carcass quality traits measured with ultrasound technology.     

The island syndrome in isolated populations of a tropical forest rodent

I examined population traits of eight isolated populations of a tropical forest rodent (Proechimys semispinosus, the Central American spiny rat) for 1 year in central Panamá. Populations were sampled by monthly live-trapping, and seven traits (density, population growth rate, adult survival, reproductive effort, age structure, sex ratio, and body mass) were compared among populations. I also compared results with published data from nearby mainland populations. Each isolated population showed characteristics typical of island populations when compared with mainland populations, including higher and more stable densities, reduced reproductive effort, and greater body mass. Densities were the highest yet recorded for this species, and biomass of these island populations was among the highest of any tropical rodent yet studied. Population traits varied not only between island and mainland populations but also among island populations. P. semispinosus have traits that allow individuals in a population to rapidly respond to temporal changes in habitat quality or resource abundance. These traits include a high reproductive rate and an ability to adjust reproductive effort to changes in density. P. semispinosus are therefore able to quickly reach and maintain high densities under favorable conditions, thereby allowing close tracking of temporally and spatially varying resources. This flexibility is predicted for habitat generalists and presumably promotes abundance and persistence in temporally and spatially heterogeneous environments. P. semispinosus, often the most abundant and widely distributed species of rodent in forests throughout their geographic range, therefore have traits that are similar to those of generalist rodents in temperate forests.     

Water homeostasis in desert-dwelling horses

This study set out to investigate tolerance of the body water pool to short-term water deprivation in horses and, in particular, to assess whether feral horses from the Namib Desert showed tolerance to dehydration superior to Transvaal. Hydration status was compared in six feral horses from the Namib Desert and in six Boerperd farm horses under conditions of normal hydration and after 72 h of dehydration. Under normal hydration, the two groups did not differ significantly in water intake, plasma sodium and potassium concentrations, plasma osmolality, hematocrit, total plasma protein, body water content, or water turnover (ml.kg-0.82.day-1). The Namib horses were significantly smaller (P less than 0.0001) and turned over 5 liters less water per day than the Boerperd during normal hydration and 4 liters less during dehydration. Increases in plasma sodium concentration after 72 h of dehydration were greater (P less than 0.05) in the Namib horses. It was concluded that horses can easily tolerate water deprivation that results in a 12% reductions in body mass. The feral horses of the Namib desert were not significantly different per unit mass from domestic horses with regard to indexes of total body water content under conditions of normal hydration and after 72 h of dehydration. Their smaller size and, hence, lower water turnover might be mechanisms they use for survival in the Namib Desert.     

Selection Practices of Bonga Sheep Reared in Southern Ethiopia

The study was conducted to assess the traits traditionally used to select the Bonga sheep reared in Southern Ethiopia. The breed was included for improvement under the community based breeding program (CBBP) launched in the year 2009. The results are based on the data collected from the project between 2009 till 2012. The findings are based on focus group discussions with the community elders who have been rearing the Bonga sheep for more than 20 years. The present findings are based on information obtained from 50% of the total respondents who were involved in CBBP. The Bonga sheep is a mutton type breed and the ewes are moderately prolific. In the past this breed of sheep were bartered to settle legal disputes and were traded against household commodities.. However, over the years (and also after the intervention of CBBP) the sheep have been selected for their body weight as well as for their distribution in many parts of Ethiopia The respondents selected the lambs at both pre weaning and post weaning stages. Traditionally the traits of rams are selected based on their body length, canon circumference, broad face, enlarged thyroid, while for the ewes, traits prolificacy, skin thickness and pelvic width are determining characters. Fat tail and brown coat color were preferred irrespective of the sexes in trait selection. The ram lambs selected for breeding purpose are locally known as “Dookoo”. These rams are selected based on some predetermined phenotypic traits and are initially selected at preweaning stage and further, again at the post weaning stage. These rams are preferentially cared and are provided with supplementary feed and comfortable housing.     

Relative allocation to horn and body growth in bighorn rams varies with resource availability

Males may allocate a greater proportion of metabolic resourcesto maintenance than to the development of secondary sexual characterswhen food is scarce, to avoid compromising their probabilityof survival. We assessed the effects of resource availabilityon body mass and horn growth of bighorn rams (Ovis canadensis)at Ram Mountain, Alberta, Canada over 30 years. The number ofadult ewes in the population tripled during our study, and theaverage mass of yearling females decreased by 13%. We used theaverage mass of yearling females as an index of resource availability.Yearling female mass was negatively correlated with the bodymass of rams of all ages, but it affected horn growth only duringthe first three years of life. Yearly horn growth was affectedby a complex interaction of age, body mass, and resource availability.Among rams aged 2–4 years, the heaviest individuals hadsimilar horn growth at high and at low resource availability,but as ram mass decreased, horn growth for a given body massbecame progressively smaller with decreasing resource availability.For rams aged 5–9 years, horn growth was weakly but positivelycorrelated with body mass, and rams grew slightly more hornfor a given body mass as resource availability decreased. Whenfood is limited, young rams may direct more resources to bodygrowth than to horn growth, possibly trading long-term reproductivesuccess for short-term survival. Although horn growth of olderrams appeared to be greater at low than at high resource availability,we found no correlation between early and late growth in hornlength for the same ram, suggesting that compensatory horn growthdoes not occur in our study population. Young rams with longerhorns were more likely to be shot by sport hunters than thosewith shorter horns. Trophy hunting could select against ramswith fast-growing horns.     

Explaining adaptive shifts in body size on islands: a life history approach

The tendency for island populations to differ in body size from their mainland relatives has been well documented, but the mechanisms for these size changes remain speculative. Explanations have typically been based on ecological interactions that directly favor either an increase or decrease in body size. While it is clear that direct ecological interactions can influence body size, life history shifts present an alternative explanation for observed insular size trends across phylogenetic groups and trophic levels. Here I describe how decreased resource availability and reduced predation pressure, the same selective forces invoked by previous hypotheses, can operate to produce body size changes via the evolution of life history traits. This mechanism is more generally applicable than previous explanations and is consistent with much of the available data.