The role of body size in mating success of Sphenarium purpurascens in Central Mexico

1. The effect of body size on the assortative mating and reproductive behaviour of the univoltine grasshopper Sphenarium purpurascens (Charpentier) was studied in Central Mexico. 2. Assortative mating by size was observed in the field. Evidence of positive assortative mating in relation to body size was found in laboratory experiments. Female fecundity and male success in contests were also correlated with body size. 3. Larger females had a higher number of eggs per pod. Larger males usually won fights and were able to take over females from other males, and to resist takeovers by other males while guarding. 4. Individuals of both sexes were observed copulating with more than one sexual partner in the field, suggesting polygamy. Male–male contests determined access to females, and males exhibited a postcopulatory prolonged mate-guarding behaviour lasting up to 18 days. 5. In a 2-year study, sex ratio was male-biased at the beginning of the reproductive season and decreased to 1:1 by the end of the season, suggesting that the population is protandrous. 6. The results of this study indicate that assortative mating results from male–male competition and female availability, and suggests that body size is a potential target of natural and sexual selection.     

Male Gryllus bimaculatus Guard Females to Delay Them from Mating with Rival Males and to Obtain Repeated Copulations

Three hypotheses for the function of postcopulatory mate guarding were tested in the field cricket Gryllus bimaculatus De Geer. The duration of spermatophore attachment was greater in the absence than in the presence of a guarding male. The ejaculate protection hypothesis was, therefore, rejected. The duration of mate guarding was found to be equal to the interval between copulations, supporting the spermatophore renewal hypothesis. In support of the rival exclusion hypothesis, the presence of a guarding male did increase the duration of spermatophore attachment when a rival male was also present. The presence of a guarding male also delayed the female from mating with the rival male. Female mating status had a significant effect on the duration of spermatophore attachment. Females mating for the first time retained the spermatophore for a significantly longer period of time than females that had mated previously.     

Effects of Temperature and Body Size on the Swimming Speed of Larval and Juvenile Atlantic Cod (Gadus Morhua): Implications for Individual-based Modelling

Synopsis The routine swimming speed (S) of three groups of 4, 9 and 32 cm total length (L_T) juvenile cod (Gadus morhua) was quantified in the laboratory at 6 – 10 different temperatures (T) between 3.2 and 16.7°C. At temperatures between 5 and 15°C, mean group S increased exponentially with increasing T (S=a e^bT) and the effect of temperature (b = 0.082, Q₁₀ = 2.27) was not significantly different among the groups (over the 8-fold difference in fish sizes of early- and post-settlement juveniles). Differences in mean S among individuals within each group were quite large (coefficient of variation = 40 – 80%). Swimming data for juveniles and those collected for groups of 0.4, 0.7 and 0.9 cm standard length (L_S) larvae were combined to assess the effect of body size on S. At 8°C, S (mm s⁻¹) increased with L_S (mm) according to: S = 0.26L_S^Φ−5.28L_S⁻¹, where Φ = 1.55L_S^−0.08. Relative S (body lengths s⁻¹) was related to L_S by a dome-shaped relationship having a maximum value (0.49 body lengths s⁻¹) at 18.5 – 19 mm L_S corresponding to the sizes of fish at the end of larval-juvenile metamorphosis. Previous larval cod IBM’s using a cruise-predator mode likely overestimated rates of foraging (prey searching and encounters) by a factor of ~2, whereas foraging rates in pause-travel models are closer to estimates of swimming velocities obtained in this and other laboratory studies.     

Size Dependent Sexual Selection in Drosophila ananassae

Mate choice based on body size is widespread and can have numerous consequences. We present data, which show the effect of male and female body size on sexual selection in Drosophila ananassae. The relationships between wing size, locomotor activity, mating latency, courtship pattern, fertility and mating success were studied. Mating latency was negatively correlated with wing length and with locomotor activity, while wing length and locomotor activity was positively correlated in males as well as in females. In female- and male-choice, we found that mate choice influenced size-assortative mating by: (1) large and small males preferring to mate with large females, (2) large males successfully competing for large females, leaving small males to mate with small females. Males increased their reproductive success by mating with large and more fecund females. In addition, in pairs of long/short winged flies, long winged flies courted and mated more successfully than short winged flies and they also have longer duration of copulation and more progeny than short winged flies. We found sterile mating in pairs of small winged males and females.     

Body Size and Proportions and Pelage Color of Free-Ranging Macaca mulatta from a Zone of Hybridization in Northeastern Thailand

Rhesus (Macaca mulatta) and long-tailed (M. fascicularis) macaques belong to the same species, and are parapatric within a zone that lies between 15° and 20° N on the Indochinese peninsula. Researchers have reported probable hybrids between the 2 species from that zone, but have not studied the extent of introgression. To test for phenotypic evidence of hybridization, we collected body mass, morphometrics (body size and proportions), and pelage color readings from free-ranging rhesus living close to the zonal boundary at Wat Tham Pa Mak Ho (WTPMH), Wang Saphun district, Loei province, northeastern Thailand (17°14′N, 101°47′E). Female WTPMH rhesus macaques (n =12) were 10–20% smaller, but with a greater relative tail length than the captive Chinese or Indian female rhesus. Female WTPMH were larger than the free-ranging long-tailed macaques, but with similar limb proportions and a shorter relative tail length. The WTPMH rhesus macaques also displayed the bipartite pelage color pattern typical of Macaca mulatta . The evidence suggests slight contribution of long-tailed macaques to the gene pool of the WTPMH population. Further sampling of other macaque populations within the zone and genetic analysis are essential to address better the question of hybridization. Determination of the distribution and range of biobehavioral variation of macaques within the zone is urgently needed, because their habitat is being rapidly destroyed by deforestation, and their demography and social structure are threatened by artificial disturbance.     

Molecular phylogeny of the subgenus Drosophila (Diptera, Drosophilidae) with an emphasis on Neotropical species and groups: a nuclear versus mitochondrial gene approach

The genus Drosophila has played an essential role in many biological studies during the last 100 years but much controversy and many incompletely addressed issues still remain to be elucidated regarding the phylogeny of this genus. Because information on the Neotropical species contained in the subgenus Drosophila is particularly incomplete, with this taxonomic group being underrepresented in many studies, we designed a study to answer some evolutionary questions related to these species. We subjected at least 41 Drosophilidae taxa to a phylogenetic analysis using a 516-base pair (bp) fragment of the alpha-methyldopa (Amd) nuclear gene and a 672 bp fragment of the mitochondrial cytochrome oxidase subunit II (COII) gene both individually and in combination. We found that the subgenus Drosophila is paraphyletic and subdivided into two main clusters: the first containing species traditionally placed in the virilis-repleta radiation and the second assembling species of the immigrans-Hirtodrosophila radiation. Inside the first of these clusters we could detect the monophyly of both the flavopilosa (the sister-clade of the annulimana group) and the mesophragmatica (closely related to the repleta group) species groups. Concerning the immigrans-Hirtodrosophila lineage, Zaprionus, Liodrosophila, Samoaia, and Hirtodrosophila were the early offshoots, followed by the immigrans, quinaria, testacea, and funebris species groups. The tripunctata radiation appears to be a derived clade, composed of a paraphyletic tripunctata group, intimately interposed with members of the cardini, guarani, and guaramunu species groups. Overall, the COII gene yielded a poor phylogenetic performance when compared to the Amd gene, the evolutionary hypothesis of which agreed with the total evidence tree. This phenomenon can be explained by the fast saturation of transitional substitutions in COII, due to strong biases in both base composition and substitution patterns, as also by its great among-site rate variation heterogeneity.     

Sexual Selection of Zorion guttigerum Westwood (Coleoptera: Cerambycidae: Cerambycinae) in Relation to Body Size and Color

Zorion guttigerum is a flower-visiting longhorned beetle endemic to New Zealand. Sexual selection of this species in relation to the body size and color form of different sexes was investigated in the field. The population sex ratio, based on censuses of feeding and mating sites (flowers), is male-biased. Females are significantly larger than males. Both sexes have antennae of similar length but the antennal length relative to the elytral length is greater in males than in females, and the antennal length of males increases more with an increase in body size than that of females. Both sexes have dark blue (DB) and yellowish-brown (YB) individuals. Both pair-bonded and solitary males are similar in elytral and antennal length. In pair-bonded males, DB individuals are significantly more numerous than YB ones, but in solitary males, the number of both color forms is similar. Males tend to have territory protection behavior, fighting with and chasing away rival males from feeding and mating sites. Larger males usually win the fight but the size-dependent fighting advantage does not translate into mating success. Male color plays an important role in mating success, with DB males having a significantly better chance to mate than YB males. Furthermore, male body size and color also have interactions in mating success: males of DB color morph obtain a greater mating advantage according to body size. Pair-bonded females are significantly larger and have longer antennae than solitary females, suggesting that males prefer larger females for mating. In addition, females of DB color morph with longer antennae are also preferred by males for mating. The significance of these findings is discussed.     

Possible Female Choice During Copulation in Ozophora baranowskii (Heteroptera: Lygaeidae): Female Behavior, Multiple Copulations, and Sperm Transfer

The remarkable behavior of female Ozophora baranowskii of lightly tapping the male with the hind legs during copulation was studied in relation to multiple copulations, copulation length, and sperm transfer. Multiple copulations did not affect female fecundity, fertility, or longevity. The incidence of tapping was higher in shorter copulations, which did not result in sperm transfer to the female. Female tapping behavior may be a mechanism for choosing between males after copulation has begun.     

The Combined Effects of Pre- and Post-Insemination Sexual Selection on Extreme Variation in Male Body Size

Orb-weaving spiders of the genus Nephila are notable for their sexual size dimorphism, with dwarf males and giant females. However, less well known is the extreme size polymorphism of males that is characteristic of some species. For example, adult male body size in N. edulis varies by an order of magnitude. Previous experiments reveal that male mating behaviour covaries with body size, suggesting the size variation is maintained by opposing pre- and post-insemination sexual selection pressures. Here, we test this idea by allowing males of different sizes to compete directly and simultaneously for access to females. Using the sterile-male technique for paternity assessment, we show that two competing males drawn from the extremes of size variations, split paternity the same way as two males of the same size drawn from the intermediate sizes. The paternity of a large male dropped from 50% to 30% on average if he competed against two instead of one small male. The large male increased his mating frequency when there were more rivals but required a much lower total duration of copulation to achieve the same paternity share. These data are consistent with the idea that opposing pre- and post-insemination selection pressures at least partly explain the variation in male body size. Co-ordinating editor: L.D. Hurst     

Multiple herbivores and coevolutionary interactions in an Ipomopsis hybrid swarm

Studies focusing on pairwise interactions between plants and herbivores may not give an accurate picture of the overall selective effect of herbivory, given that plants are often eaten by a diverse array of herbivore species. The outcome of such interactions may be further complicated by the effects of plant hybridization. Hybridization can lead to changes in morphological, phenological and chemical traits that could in turn alter plant–herbivore interactions. Here we present results from manipulative field experiments investigating the interactive effects of multiple herbivores and plant hybridization on the reproductive success of Ipomopsis aggregata formosissima X I. tenuituba. Results showed that ungulate herbivores alone had a net positive effect on plant relative fitness, increasing seed production approximately 2-fold. Caterpillars had no effect on plant relative fitness when acting alone, with caterpillar-attacked plants producing the same number of flowers, fruits and seeds as the uneaten controls. Caterpillars, however, significantly reduced flower production of ungulate browsed plants. Flower production in these plants, however, was still significantly greater (approximately 1.7-fold greater) than uneaten controls, likely leading to an increase in reproductive success through the paternal component of fitness given that fruit and seed production was not significantly different from that of herbivore-free controls. Although results suggest that herbivore imposed selection is pairwise, ungulates likely have a large influence on the abundance of, and hence the amount of damage caused by, caterpillar herbivores. Thus, because of the ecological interactions between ungulates and caterpillars, selection on Ipomopsis may be diffuse rather than pairwise, assuming such interactions translate into differential effects on plant fitness as herbivore densities vary. Plant hybridization had no significant effect on patterns of ungulate or caterpillar herbivory; i.e., no significant interactions were detected between herbivory and plant hybridization for any of the fitness traits measured in this study nor did plant hybridization have any significant effect on host preference. These results may be due to patterns of introgression or the lack of species-specific differences between I. aggregate formosissima and I. tenuituba. Plant hybridization per se resulted in lowered reproductive success of white colored morphs due in part to the effects of pollination. Although it appears that there would be strong directional selection favoring darker flower colors due to the lower reproductive success of the white colored morphs in the short run, the natural distribution of hybrids suggest that over the long run selection either tends to average out or there are no fitness differences among morphs in most years due to the additive fitness effects of hawkmoth and hummingbird pollinators.     

Importance of Conserving Alternative Pollinators: Assessing the Pollination Efficiency of the Squash Bee, Peponapis limitaris in Cucurbita moschata (Cucurbitaceae)

Although the honey bee, Apis mellifera, has been considered the best pollinator for crops needing insect pollination, the current pandemic of varroatosis among honeybees highlights the need to find additional or alternative species as managed crop pollinators. Moreover, there is evidence that A. mellifera may not always be the most efficient pollinator. Introduction of A. mellifera into crops may be unnecessary, and even detrimental to non-Apis bee populations, which should be considered as an alternative for crop production improvement. Evaluating the pollination efficiency of non-Apis bees is one of the first steps in planning successful strategies for their conservation. In this study, we evaluated the pollination efficiency of Peponapis limitaris and A. mellifera in plots of Cucurbita moschata: pollen removal and deposition; pollinator visit frequency; and the pollinator visit–nectar production relationship. The results show P. limitaris to be the most efficient pollinator as: (1) both males and females remove and deposit almost four times as much pollen as A. mellifera; (2) they make significantly more floral visits than A. mellifera; and (3) their visit frequency shows a strong relationship to C. moschata nectar production during anthesis. Recommendations arising from this study are: (1) the introduction of A. mellifera be avoided in C. moschata crops; and (2) basic research be done on the biology of P. limitaris that contribute to its conservation and greater exploitation.     

Group Composition, Home Range Size, and Diet of Three Sympatric Bamboo Lemur Species (Genus Hapalemur) in Ranomafana National Park, Madagascar

This is the first long-term, simultaneous, comparative study of three bamboo lemur species (Hapalemur griseus, H. aureus, and H. simus) at a site in southeastern-central Madagascar where they occur in sympatry. At Talatakely, Ranomafana National Park, the three Hapalemur spp. share overlapping home ranges. Hapalemur griseus has flexible group sizes, varying from three to nine individuals (n = 6). The home range of Hapalemur griseus averages 15 ha (n = 2). Hapalemur aureus forms family groups of 4 individuals (n = 3); they have a home range on average of 26 ha (n = 2). The single group of Hapalemur simus is composed of one or three adult males, two adult females, and their offspring; they occupy a home range of 62 ha. The three species of Hapalemur are year-round bamboo specialists: >88% of their diets consist of bamboo and grass in the Family Poaceae. Contrary to earlier findings, all three Hapalemur spp. consume the cynogenic parts—young leaf bases, young pseudopetioles, and young shoots—of the giant bamboo, Cathariostachys madagascariensis. They rely heavily on this plant, which comprises 72–95% of their diets. Hapalemur griseus and H. aureus consume similar proportions of bamboo vs. nonbamboo plants, though they differ in the species of bamboo they prefer. Hapalemur simus has the most distinct diet of the three bamboo lemurs. They exploit the young shoots of Cathariostachys madagascariensis during the austral summer rainy season, between November and April. From June to November, Hapalemur simus shifts its diet to eating the mature culm pith of Cathariostachys madagascariensis; the proportion of pith represented in their diet reaches a maximum of 89% in October. Seasonal availability of food resources, feeding competition, and factors related to body size may provide clues to the understanding of diet selection among the three sympatric Hapalemur spp.     

Size and fecundity hierarchies in an herbaceous perennial

Summary Inequalities in size in populations have potentially important effects on fitness but have rarely been examined in natural populations. I measured size (number of culms) and fecundity (number of spikelets) in five populations of the grass Danthonia spicata from 1981 to 1985. The populations were in sites in a Pinus-Quercus-Populus forest in northern lower Michigan, USA comprising a secondary succession sequence. The sites had been burned in 1980, 1954, 1948, 1936, and 1911, respectively. Mean sizes and fecundities and the amount of hierarchy in size and fecundity, measured by the Gini coefficient, were compared between the YOUNG population, 1980 burn site, and the OLD populations, 1954, 1948, 1936, and 1911 burn sites. I found large differences in mean size and fecundity between the YOUNG and OLD populations with much larger individuals in the YOUNG population. No differences in size hierarchies were found in either the first year of measurement or after five years. The fecundity hierarchies showed no significant difference among the populations in the first year but after five years the YOUNG population showed a significant decrease in amount of inequality. The longterm patterns of size and fecundity hierarchies differed because fecundity was a cumulative trait while size was not. Size inequalities may not always be a good measure of fecundity inequalities. Short-term measures of inequality in perennials may not be a good indicator of long-term values. In contrast to greenhouse studies, habitat light levels did not affect size hierarchies although they did affect fecundity hierarchies.     

Ecological and selective effects of stigma-anther separation in the self-incompatible tropical tree Ipomoea wolcottiana (Convolvulaceae)

We evaluated the effect of stigma-anther separation (SAS) in flowers of the self-incompatible tropical tree Ipomoea wolcottiana (Convolvulaceae). We measured the deposition of self pollen on the stigma and the growth of pollen tubes. We also evaluated phenotypic selection on female fertility using univariate regressions of SAS with fruit production. SAS varied among individuals in two years, with a range of 0–6 mm. There was a negative correlation between SAS and the number of self pollen grains on the stigma; trees with SAS of 0–2 mm received averages of ca 80–130 self grains. The effect of loading the stigma with self-pollen was tested by allowing isolated flowers to self-pollinate, then cross-pollinating by hand: there was a negative correlation between the pollen load and the proportion of grains forming pollen tubes. The highest proportion of grains forming tubes was found with loads of <10 grains, while the minimum frequency of tubes occurred with loads of >50 grains. The results suggest that more self pollen deposition leads to fewer tubes from outcross pollen. There was a significant effect of SAS on fruit production in both years. The analyses suggested stabilizing selection; very small SAS is disadvantageous due to the high load of self pollen, while very large SAS probably led to damage to the style during pollinator visits. We suggest that herkogamy may be a key character for determining the success of fruit production in self-incompatible hermaphroditic species.     

Effects of behavioral and morphological plasticity on risk of predation in a Neotropical tadpole

Predator-induced phenotypic plasticity is widespread among aquatic animals, however the relative contributions of behavioral and morphological shifts to reducing risk of predation remain uncertain. We tested the phenotypic plasticity of a Neotropical tadpole (Rana palmipes) in response to chemical cues from predatory Belostoma water bugs, and how phenotype affects risk of predation. Behavior, morphology, and pigmentation all were plastic, resulting in a predator-induced phenotype with lower activity, deeper tail fin and muscle, and darker pigmentation. Tadpoles in the predator cue treatment also grew more rapidly, possibly as a result of the nutrient subsidy from feeding the caged predator. For comparison to phenotypes induced in the experiment, we quantified the phenotype of tadpoles from a natural pool. Wild-caught tadpoles did not match either experimentally induced phenotype; their morphology was more similar to that produced in the control treatment, but their low swimming activity was similar to that induced by predator cues. Exposure of tadpoles from both experimental treatments and the natural pool to a free-ranging predator confirmed that predator-induced phenotypic plasticity reduces risk of predation. Risk of predation was comparable among wild-caught and predator-induced tadpoles, indicating that behavioral shifts can substantially alleviate risk in tadpoles that lack the typical suite of predator-induced morphological traits. The morphology observed in wild-caught tadpoles is associated with rapid growth and high competition in other tadpole species, suggesting that tadpoles may profitably combine a morphology suited to competition for food with behaviors that minimize risk of predation.     

Genetic and phenotypic variation for flight ability in the cactophilicDrosophila species,D. aldrichi andD. buzzatii

The flight ability ofDrosophila aldrichi (Patterson & Crow) andD. buzzatii (Patterson & Wheeler) using tethered flights, was measured with respect to age-related changes, genetic variation and adult body size variation induced by rearing at different larval densities.Drosophila buzzatii flew for much longer thanD. aldrichi, especially females, but age-related changes in flight duration were significant only forD. aldrichi. Effects of body size on flight ability were significant inD. buzzatii, but not inD. aldrichi. InD. buzzatii, there was a significant genotype-environment interaction (larval density × line) for flight duration, with short and average flight duration isofemale lines showing longer flights, but a long flight duration line shorter flights as body size decreased (i.e., as larval density increased). Heritability estimates for flight duration were similar in the two species, but flight duration showed no significant genetic correlations with developmental time, body size or wing dimensions (except for one wing dimension inD. buzzatii). Although not significantly different between the species, heritabilities for life-history traits (adult size and developmental time) showed contrasting patterns — with higher heritability for body size (body weight and thorax length) inD. buzzatii, and higher for developmental time inD. aldrichi. In agreement with limited previous field evidence,D. buzzatii is better adapted for colonization than isD. aldrichi.     

Changes in Baboon Feeding Behavior: Maturity-dependent Fruit and Seed Size Selection within a Food Plant Species

Despite considerable inter- and intraindividual variation in fruit and seed size in many plant species, researchers have given little attention to the relevance of the traits for primate fruit choice within a food plant species and its implications for tree regeneration. We studied feeding behavior and selectivity of olive baboons (Papio anubis) in the African locust bean (Parkia biglobosa, Mimosaceae), via direct observations of habituated groups and indirect evidence from leftovers of pods after feeding events. Olive baboons acted as both seed predators and dispersers for Parkia biglobosa. They fed on and destroyed unripe seeds, and swallowed intact ripe seeds when consuming mature fruit pulp. Predation rate was high, and only 10% of the seeds were dispersed. Predation and dispersal of seeds is linked to seed number and size. Digestible unripe seeds accounted for 10% of the unripe fruit mass, while indigestible ripe seeds made up 28% of the mature fruit mass. With these constraints, olive baboons increased food gain per fruit by selecting unripe pods containing a high number of large and heavy seeds. Consequently, only pods with fewer and smaller seeds remained for maturation. Thereafter, baboons fed on mature pods containing the smallest seeds, and exploited pods with more seeds to a greater extent than those with fewer seeds. Thus, fruits with small seeds and an intermediate seed number contributed the most to dispersal by baboons.     

青藏高原露蕊乌头不同海拔居群的开花持续期与繁殖分配

对青藏高原东部3个不同海拔居群的露蕊乌头(Aconitum gymnandrum)(毛茛科)开花持续期和繁殖分配进行了野外观察和检测试验。结果表明:(1)随着海拔的升高,露蕊乌头的开花持续期时间延长,且不同居群间存在显著差异。(2)露蕊乌头的植株个体越大,繁殖投入越高。(3)露蕊乌头的繁殖分配在地上总生物量中所占的比例随海拔升高而增加。研究认为,露蕊乌头在高海拔地区较高的繁殖分配比例强调了有性繁殖在高山恶劣环境中的重要性,而高海拔地区开花持续期的延长补偿了传粉昆虫的减少并增加了对传粉昆虫的吸引能力。     

Body size and proportions in chimpanzees,with special reference toPan troglodytes schweinfurthii from Gombe National Park,Tanzania

Body weight, cranial capacity, linear and joint area data from ten free-ranging adult chimpanzees from Gombe National Park, Tanzania with known life histories allow study of variation in a local population and comparison to other populations ofPan troglodytes and toPan paniscus. Because individuals in the Gombe population are small compared to other common chimpanzees, they provide a useful comparison toPan paniscus. Body weight and some linear dimensions overlap withPan paniscus. However, cranial capacity, tooth size, and body proportions of Gombe individuals lie within the range of otherPan troglodytes and are distinct fromPan paniscus.