Mammals of Borneo – small size on a large island

Aim Island mammals have featured prominently in models of the evolution of body size. Most of these models examine size evolution across a wide range of islands in order to test which island characteristics influence evolutionary pathways. Here, we examine the mammalian fauna of a single island, Borneo, where previous work has detected that some mammal species have evolved a relatively small size. We test whether Borneo is characterized by smaller mammals than adjacent areas, and examine possible causes for the different trajectories of size evolution between different Bornean species. Location Sundaland: Borneo, Sumatra, Java and the Malay/Thai Peninsula. Methods We compared the mammalian body size frequency distributions in the four areas to examine whether the large mammal fauna of Borneo is more depauperate than elsewhere. We measured specimens belonging to 54 mammal species that are shared between Borneo and any of the other areas in order to determine whether there is an intraspecific tendency for Bornean mammals to evolve small body size. Using data on diet, body size and geographical ranges we examine factors that are thought to influence body size. Results Borneo has fewer large mammals than the other areas, but this is not statistically significant. Large Bornean mammals are significantly smaller than their conspecifics in the other regions, while there are no differences between the body sizes of mammals on Sumatra, Java and the Malay/Thai Peninsula. The finding that large mammals show the greatest size difference between Borneo and elsewhere contrasts with some models of size evolution on islands of different areas. Diet does not correlate with the degree of size reduction. Sunda region endemics show a weaker tendency to be small on Borneo than do widespread species. Main conclusions We suggest that soil quality may drive size evolution by affecting primary productivity. On Borneo, where soils are generally poor in nutrients, this may both limit biomass and cause mammals to be reduced in body size. We hypothesize that widespread species respond to low resource abundance by reducing in size, while endemic elements of the fauna have had longer to adjust to local conditions by altering their behaviour, physiology and/or ecology, and are thus similar in size across the region.     

Competition, niche specialization and the evolution of brain size in the genus Peromyscus

Previous studies of relative brain size in mammals have suggested an association with complex habitats and with low reproductive rate. In order to examine the causal relationships more thoroughly, a detailed examination of relative brain size variation in the genus Peromyscus was undertaken. Endocranial volumes were used to estimate brain weight for 32 species including 161 subspecies, and relative brain size calculated as the species deviation from the allometric relationship between brain and body size. The intrageneric allometric coefficient was higher than most values previously reported from low taxonomic levels, but intraspecific coefficients were generally lower than this.
Island species, and relict species isolated on mountain tops, which may be ecological 'islands', had consistently small relative brain sizes, but peninsular species were large brained. Among the remaining species there were significant correlations between litter size and relative brain size, and between the number of competitor species and relative brain size. Species with many competitor species have relatively large brains and small litters. It is concluded that the nature of the geographical distribution, the pattern of species formation and habitat complexity all influence relative brain size in existing forms.     

Caterpillars selected for large body size and short development time are more susceptible to oxygen‐related stress

Recent studies suggest that higher growth rates may be associated with reduced capacities for stress tolerance and increased accumulated damage due to reactive oxygen species. We tested the response of Manduca sexta (Sphingidae) lines selected for large or small body size and short development time to hypoxia (10 kPa) and hyperoxia (25, 33, and 40 kPa); both hypoxia and hyperoxia reduce reproduction and oxygen levels over 33 kPa have been shown to increase oxidative damage in insects. Under normoxic (21 kPa) conditions, individuals from the large‐selected (big‐fast) line were larger and had faster growth rates, slightly longer developmental times, and reduced survival rates compared to individuals from a line selected for small size (small‐fast) or an unselected control line. Individuals from the big‐fast line exhibited greater negative responses to hyperoxia with greater reductions in juvenile and adult mass, growth rate, and survival than the other two lines. Hypoxia generally negatively affected survival and growth/size, but the lines responded similarly. These results are mostly consistent with the hypothesis that simultaneous acquisition of large body sizes and short development times leads to reduced capacities for coping with stressful conditions including oxidative damage. This result is of particular importance in that natural selection tends to decrease development time and increase body size.     

Adaptive size modification by dominant female meerkats

In species of cooperative insects that live in large groups, selection for increased fecundity has led to the evolution of an increased body size among female reproductives, but whether this is also true of cooperative vertebrates is unknown. Among vertebrates, morphological modification of female breeders has only been documented in a single species; in naked mole rats (Heterocephalus glaber), acquisition of alpha status is associated with a significant increase in body size through an elongation of the lumbar vertebrae. Here we provide evidence of morphological modification among breeding females of a cooperative carnivore, the meerkat (Suricata suricatta), and demonstrate that this modification is likely to be adaptive. The same female meerkats were significantly larger when they were dominant than when they were subordinate. This increased body size was not explained by differences in age, foraging efficiency, or investment in offspring care, but may have arisen, in part, through increased levels of hormone that govern bone growth. Increases in body size are likely to result in fitness benefits, for large females delivered larger litters and had heavier offspring, both of which are known to correlate positively with measures of breeding success in meerkats. Our results suggest that the acquisition of alpha status in female meerkats is associated with an adaptive increase in body size and hence that morphological modification of female vertebrates may be more widespread than has been previously supposed.     

Competition,niche specialization and the evolution of brain size in the genus Peromyscus

Previous studies of relative brain size in mammals have suggested an association with complex habitats and with low reproductive rate. In order to examine the causal relationships more thoroughly, a detailed examination of relative brain size variation in the genus Peromyscus was undertaken. Endocranial volumes were used to estimate brain weight for 32 species including 161 subspecies, and relative brain size calculated as the species deviation from the allometric relationship between brain and body size. The intrageneric allometric coefficient was higher than most values previously reported from low taxonomic levels, but intraspecific coefficients were generally lower than this. Island species, and relict species isolated on mountain tops, which may be ecological ‘islands’, had consistently small relative brain sizes, but peninsular species were large brained. Among the remaining species there were significant correlations between litter size and relative brain size, and between the number of competitor species and relative brain size. Species with many competitor species have relatively large brains and small litters. It is concluded that the nature of the geographical distribution, the pattern of species formation and habitat complexity all influence relative brain size in existing forms.     

Little effect of climate change on body size of herbivorous beetles

Ongoing climate change affects various aspects of an animal's life, with important effects on distribution range and phenology. The relationship between global warming and body size changes in mammals and birds has been widely studied, with most findings indicating a decline in body size over time. Nevertheless, little data exist on similar size change patterns of invertebrates in general and insects in particular, and it is unclear whether insects should decrease in size or not with climate warming. We measured over 4000 beetle specimens, belonging to 29 beetle species in 8 families, collected in Israel during the last 100 years. The sampled species are all herbivorous. We examined whether beetle body size had changed over the years, while also investigating the relationships between body size and annual temperature, precipitation, net primary productivity (NPP) at the collection site and collection month. None of the environmental variables, including the collection year, was correlated with the size of most of the studied beetle species, while there were strong interactions of all variables with species. Our results, though mostly negative, suggest that the effect of climate change on insect body size is species‐specific and by no means a general macro‐ecological rule. They also suggest that the intrapopulation variance in body size of insects collected as adults in the field is large enough to conceal intersite environmental effects on body size, such as the effect of temperature and NPP.     

Measuring the relationship between dietary quality and body size in primates

The feeding niche and the body size of any species are fundamental parameters that constrain the evolution of many other phenotypic characters. Moreover, previous work has shown that body size and diet are correlated, as a consequence of the negative allometry of metabolic rate. Unfortunately, the precise form of the association between body size and diet has never been specified, principally because no suitable cross-species measure of diet has been advanced. Here we develop a measure of diet that is sensitive over the whole spectrum of primate feeding niches, and use this measure to define the relationship between body size and diet for a sample of 72 primate species. Subsequently, we present several examples of how behavioral and ecological hypotheses can be tested by examining the extent to which particular species deviate from the general diet-body size pattern.     

Incisor size and diet revisited: The view from a platyrrhine perspective

Allometric relationships between incisor size and body size were determined for 26 species of New World primates. While previous studies have suggested that the incisors of Old World primates, and anthropoids in general, scale isometrically with body size, the data presented here indicate a negative allometric relationship between incisor size and body size among New World species. This negative allometry was exhibited by platyrrhines when either upper or lower incisor row length was regressed against body weight, and when either least-squares or bivariate principal axis equations were used. When upper incisor length was plotted against skull length, negative allometry could be sustained using both statistical techniques only when the full sample of 26 species was plotted. The choice of variables to represent incisor size and body size, and the choice of a statistical technique to effect the allometric equation, had a more pronounced impact on the location of individual species with regard to lines of best fit. Platyrrhines as a group have smaller incisors relative to body size than do catarrhines, regardless of diet. Among New World primates, small incisors represent a plausible primitive condition; species with relatively large incisors manifest a phyletic change associated with a dietary shift to foods that require increased incisal preparation. The opposite trend characterizes Old World primates. In spite of the taxonomic differences in relative incisor size between platyrrhine and catarrhine primates, inferences about diet derived from an allometric equation for all anthropoids should prove reliable as long as the species with unknown diet does not lie at the upper end of the body size range for platyrrhines or catarrhines.     

Sociality, ecology, and relative brain size in lemurs

The social brain hypothesis proposes that haplorhine primates have evolved relatively large brains for their body size primarily as an adaptation for living in complex social groups. Studies that support this hypothesis have shown a strong relationship between relative brain size and group size in these taxa. Recent reports suggest that this pattern is unique to haplorhine primates; many nonprimate taxa do not show a relationship between group size and relative brain size. Rather, pairbonded social monogamy appears to be a better predictor of a large relative brain size in many nonprimate taxa. It has been suggested that haplorhine primates may have expanded the pairbonded relationship beyond simple dyads towards the evolution of complex social groups. We examined the relationship between group size, pairbonding, and relative brain size in a sample of 19 lemurs; strepsirrhine primates that last share a common ancestor with monkeys and apes approximately 75 Ma. First, we evaluated the social brain hypothesis, which predicts that species with larger social groups will have relatively larger brains. Secondly, we tested the pairbonded hypothesis, which predicts that species with a pairbonded social organization will have relatively larger brains than non-pairbonded species. We found no relationship between group size or pairbonding and relative brain size in lemurs. We conducted two further analyses to test for possible relationships between two nonsocial variables, activity pattern and diet, and relative brain size. Both diet and activity pattern are significantly associated with relative brain size in our sample. Specifically, frugivorous species have relatively larger brains than folivorous species, and cathemeral species have relatively larger brains than diurnal, but not nocturnal species. These findings highlight meaningful differences between Malagasy strepsirrhines and haplorhines, and between Malagasy strepsirrhines and nonprimate taxa, regarding the social and ecological factors associated with increases in relative brain size. The results suggest that factors such as foraging complexity and flexibility of activity patterns may have driven selection for increases in brain size in lemurs.     

Body size explains interspecific variation in size–latitude relationships in geographically widespread beetle species

1. In most birds and mammals, larger individuals of the same species tend to be found at higher latitudes, but in insects, body size–latitude relationships are highly variable. 2. Recent studies have shown that larger‐bodied insect species are more likely to decrease in size when reared at increased temperature, compared with smaller‐sized species. These findings have led to the prediction that a positive relationship between body size and latitude should be more prevalent in larger‐bodied insect species. 3. This study measured the body size of > 4000 beetle specimens (12 species) collected throughout North America. Some beetle species increased in size with latitude, while others decreased. Importantly, mean species body size explained c. 30% of the interspecific variation in the size–latitude response. 4. As predicted, larger‐bodied beetle species were more likely to show a positive relationship between body size and latitude (Bergmann's rule), and smaller‐bodied species were more likely to show a negative body size–latitude relationship (inverse Bergmann's rule). 5. These body size–latitude patterns suggest that size‐specific responses to temperature may underlie global latitudinal distributions of body size in Coleoptera, as well as other insects.     

Range size in North American Enallagma damselflies correlates with wing size

1. Cross-species macroecological comparisons in freshwater invertebrates have been restricted by a lack of large-scale distributional data, and robust phylogenies. Here, we use data from the OdonataCentral database to explore body length–range size and wing length–range size relationships in damselflies from the genus Enallagma ; the recent publication of a phylogeny for this group meant that, as well as a cross-species analysis, we were able to assess relationships in a phylogenetically controlled manner.
2. For cross-species comparisons, only wing length showed significant (positive) regression relationships with range size and occupancy, although the inclusion of body length in multiple regressions increased the fit of the models. Damselflies with larger wings relative to their body length had larger distributions, a result confirmed by a significant positive relationship between range size and residuals from the regression of wing size on body size.
3. For the phylogenetically controlled analyses, only wing length contrast scores were significantly related to distribution patterns and entered into regression models; the significant positive relationships between wing length contrasts and both range size and occupancy contrasts suggested that evolutionary increases in wing length had occurred alongside range expansions.
4. Together these results suggest that species of Enallagma with larger wings (both absolute and relative to body length) tend to be more widely distributed in North America and that the evolution of wing size may have played a role in range expansion. No such relationships were evident for body size. We discuss the potential importance of wing morphometrics for studying the evolutionary ecology of freshwater insects.     

Atmospheric oxygen level and the evolution of insect body size

Insects are small relative to vertebrates, possibly owing to limitations or costs associated with their blind-ended tracheal respiratory system. The giant insects of the late Palaeozoic occurred when atmospheric PO₂ (aPO₂) was hyperoxic, supporting a role for oxygen in the evolution of insect body size. The paucity of the insect fossil record and the complex interactions between atmospheric oxygen level, organisms and their communities makes it impossible to definitively accept or reject the historical oxygen-size link, and multiple alternative hypotheses exist. However, a variety of recent empirical findings support a link between oxygen and insect size, including: (i) most insects develop smaller body sizes in hypoxia, and some develop and evolve larger sizes in hyperoxia; (ii) insects developmentally and evolutionarily reduce their proportional investment in the tracheal system when living in higher aPO₂, suggesting that there are significant costs associated with tracheal system structure and function; and (iii) larger insects invest more of their body in the tracheal system, potentially leading to greater effects of aPO₂ on larger insects. Together, these provide a wealth of plausible mechanisms by which tracheal oxygen delivery may be centrally involved in setting the relatively small size of insects and for hyperoxia-enabled Palaeozoic gigantism.     

Sexually antagonistic selection on primate size

Abstract Male intrasexual selection in haplorhine primates has previously been shown to increase male size and to a lesser degree also female size. I address the following questions: (1) why does female size increase when the selection is on males, and (2) why does female size not increase to the same extent as that of males. The potential for correlational selection on females through increased resource competition was analysed with independent contrasts analyses. No such effect was found, nor did matched pairs comparisons reveal females to increase in size because of selection to bear larger male offspring. Instead further matched pairs analyses revealed higher female postpartum investment, as indicated by a longer lactation period, in more sexually selected species, also after correcting for body weight. Concerning the second question, independent contrast analyses showed that large size has had negative effects on female reproductive rate across the primate order. Matched‐pairs analyses on haplorhines revealed that females of species in more polygynous clades have lower reproductive rates than females of species in less polygynous clades. This is also true after the effects of body weight are removed. These results, both when correcting for body weight and when not, suggest that sexual selection has shifted female size from one favouring female lifetime fecundity to one favouring male success in competition. This depicts antagonistic selection pressures on female size and a trade‐off for females between the ecologically optimal size of their foremothers and the larger size that made their forefathers successful.     

Body size and resource partitioning in ladybirds

Aphidophagous insects all exploit aphids as food, but there appears to be no association between the size of the aphidophagous predator and that of the species or the age structure of the aphid colonies they exploit. Aphid colonies generally increase, peak and decline in abundance, and are exploited by a sequence of predators, which is consistent from year to year. The objective of this study was to determine the rules underlying this temporal pattern. For example, in the field, aphid colonies are often first attacked by a small and then a larger species of ladybird. Theory based on the geometry and physiology of ladybirds predicts that the quantity of food required for oviposition and the area searched per unit time should scale with body weight, with exponents of 1 and 0.66, respectively. An analysis of empirical data supports these predictions. Thus, in relative terms a 35 mg ladybird requires 1.5 times more aphids per unit area for oviposition than a 10 mg ladybird. That is, the temporal pattern in oviposition is possibly mainly determined by geometrical and physiological constraints associated with body size, with small species of ladybird able to lay eggs at lower aphid population densities than large species. Cannibalism is thought to be the mechanism by which these predators are able to coexist.     

Body size and sexual size dimorphism in calanoid copepods

Patterns of sexual size dimorphism and body size in calanoid copepods are examined. We hypothesize that favorable conditions for development will result in large body size and high sexual size dimorphism among populations of a given species and that differences in this allometric relationship among species is governed by the male's role in insemination. We confirm that there is a greater advantage to large female size, normally the larger sex, when compared to males, hence leading to selection for developmental patterns favoring high size dimorphism. Individuals from populations of four centropagid copepod species were measured; other sizes were obtained from published sources. In the four species we examined, the relationships between prosome length and both clutch size and the ability to produce multiple clutches with one insemination were determined. Results show a trend toward hyperallometry in all centropagid species examined: sexual size dimorphism increases with increasing size. Large females produce larger clutches and more additional clutches on one insemination. That hyperallometry is not observed in diaptomid copepods may result from the greater role the male plays in reproduction. Males are needed for each clutch produced, hence the selective pressure to be larger is greater than that in the centropagidae.     

Explaining the seasonal decline in litter size in European ground squirrels

In European ground squirrels Spermophilus citellus as in many ground squirrel species. late born litters are composed of fewer young than early born litters. Two alternative though not mutually exclusive hypotheses may explain this seasonal pattern of change in litter size. On the one hand. the production of few large young late in the season may be an adaptation to time limitations on the offspring. that have to complete growth and fattening prior to hibernation. Then one would expect a trade-off between offspring number and size as the breeding season progresses. At its extreme. this hypothesis would predict that total maternal effort should be equal independent of litter size. Alternatively. litter size may be determined by physiological limitations on the mother. in that highly constrained mothers breed later and produce smaller litters. Then one would expect reduced overall maternal effort in highly constrained mothers of smaller litters. In this case. a trade-off between litter size and offspring size would not be expected. We found that total maternal effort in terms of gestation length and the duration of lactation increased with increasing litter size. thus supporting the second hypothesis. Lactation was not terminated at natal emergence. It extended a relatively long period of time beyond the time of first litter emergence depending on litter size. During prolonged lactation. individual young of large litters made up body mass to young of small litters. As a consequence. juvenile weaning body mass was unaffected by litter size although offspring body mass at natal emergence was inversely related to litter size. This additional weight gain in young of large litters compensated for initial survival disadvantages and presumably affected fecundity at yearling age.     

DEGREE OF SPECIALIZATION IS RELATED TO BODY SIZE IN HERBIVOROUS INSECTS: A PHYLOGENETIC CONFIRMATION

Numerous studies have suggested a general relationship between the degree of host specialization and body size in herbivorous animals. In insects, smaller species are usually shown to be more specialized than larger‐bodied ones. Various hypotheses have attempted to explain this pattern but rigorous proof of the body size–diet breadth relationship has been lacking, primarily because the scarceness of reliable phylogenetic information has precluded formal comparative analyses. Explicitly using phylogenetic information for a group of herbivores (geometrid moths) and their host plant range, we perform a comparative analysis to study the body size–diet breadth relationship. Considering several alternative measures of body size and diet breadth, our results convincingly demonstrate without previous methodological issues—a first for any taxon—a positive association between these traits, which has implications for evaluating various central aspects of the evolutionary ecology of herbivorous insects. We additionally demonstrate how the methods used in this study can be applied in assessing hypotheses to explain the body size–diet breadth relationship. By analyzing the relationship in tree‐feeders alone and finding that the positive relationship remains, the result suggests that the body size–diet breadth relationship is not solely driven by the type of host plant that species feed on.     

Experiments on clutch size and nest size in passerine birds

Summary Results of experiments on three passerine species suggest that brood size may be constrained by nest size, since the breeding success of pairs provided with large nestcups was greater than that of those provided with small artificial nestcups. These results may have important implications, e.g. to the design of experiments involving manipulation of clutch and brood size. A small nestcup is requisite for successful hatching during the incubation period, but a large one for successful rearing during the nestling period. In nature this difference may select for types of nesting materials that are elastic, such as mosses and lichens. However, experiments showed that such materials rapidly absorb rainwater but only slowly dry out. In addition, because large nests dry out more slowly than small nests, selection will favour small nests among those open-nesting species that have exposed nests. A further possible nest size constraint on open-nesters is nest predation. However, no difference in the predation rate was found in experiments with small and large artificial nests.     

Problems of body-weight estimation in fossil primates

Body-weight estimates of fossil primates are commonly used to infer many important aspects of primate paleobiology, including diet, ecology, and relative encephalization. It is important to examine carefully the methodologies and problems associated with such estimates and the degree to which one can have confidence in them. New regression equations for predicting body weight in fossil primates are given which provide body-weight estimates for most nonhominid primate species in the fossil record. The consequences of using different subgroups (evolutionary “grades”) of primate species to estimate fossil-primate body weights are explored and the implications of these results for interpreting the primate fossil record are discussed. All species (fossil and extant) were separated into the following “grades”: prosimian grade, monkey grade, ape grade, anthropoid grade, and all-primates grade. Regression equations relating lower molar size to body weight for each of these grades were then calculated. In addition, a female-anthropoid grade regression was also calculated for predicting body weight infernales of extinct, sexually dimorphic anthropoid species. These equations were then used to generate the fossil-primate body weights. In many instances, the predicted fossil-primate body weights differ substantially from previous estimates.     

Mass extinctions do not explain skew in interspecific body size distributions

In several higher animal taxa, such as mammals and birds, the distribution of species body sizes is heavily skewed towards small size. Previous studies have suggested that small‐bodied organisms are less prone to extinction than large‐bodied species. If small body size is favourable during mass extinction events, a post mass extinction excess of small‐bodied species may proliferate and maintain skewed body size distributions sometime after. Here, we modelled mass extinctions and found that even unrealistically strong body mass selection has little effect on the skew of interspecific body size distributions. Moreover, selection against large body size may, counter intuitively, skew size distributions towards large body size. In any case, subsequent evolutionary diversification rapidly erases these rather small effects mass extinctions may have on size distributions. Next, we used body masses of extant species and phylogenetic methods to investigate possible changes in body size distributions across the Cretaceous–Paleogene (K‐Pg) mass extinction. Body size distributions of extant clades that originated during the Cretaceous are on average more skewed than their subclades that originated during the Paleogene, but the difference is only minor in mammals, and in birds, it can be explained by a positive relationship between species richness and skewness that is also present in clades that originated after the transition. Hence, we cannot infer from extant species whether the K‐Pg mass extinctions were size‐selective, but they are not the reason why most extant bird and mammal species are small‐bodied.