Crickets with extravagant mating songs compensate for predation risk with extra caution

Modern models for the evolution of conspicuous male mating displays assume that males with conspicuous displays must bear the cost of enhanced predation risk. However, if males can compensate behaviourally for their increased conspicuousness by acting more cautiously towards predators, they may be able to lower this cost. In the field cricket Gryllus integer, males call to attract females, and differ in their durations of uninterrupted trilling (calling-bout lengths). Differences among males in calling-bout lengths are heritable, and females prefer males with longer calling bouts. In this study, males with longer, more conspicuous songs behaved more cautiously than males with shorter songs on two different tests of predator avoidance. They took longer to emerge from a safe shelter within a novel, potentially dangerous environment, and they ceased calling for a longer time when their calls were interrupted by a predator cue. Thus, these males appear to compensate behaviourally for their more conspicuous mating displays. Additionally, latencies to emerge from a shelter in the novel environment were consistent over time for both individual males from the field and males that had been reared in the laboratory, indicating that the differences in latency among males may be heritable.     

Optimal time to emerge from refuge

Factors affecting emergence by prey that enter refuges when approached by predators have been studied intensively, but only two theoretical models predict how long prey should remain in a refuge before emerging. We argue that prey can make better decisions than allowed by one model; the other model describes cases in which predators wait for prey to emerge. We present optimality models that permit prey to select a time to emerge that maximizes fitness. When in a refuge, a prey cannot obtain benefits outside; emerging too soon can be catastrophic, but delaying emergence entails loss of fitness. If predators resume foraging quickly rather than engaging in strategic waiting games, current theory suggests that prey emerge when the costs of remaining in a refuge and of emerging are equal. However, prey often can do better by emerging at the time maximizing fitness rather than when benefits equal costs (i.e. when prey break even). Optimal emergence time depends on initial fitness, benefits lost by remaining in refuge, and the decay rate of predation risk. Benefits lost if a prey is killed are modelled separately from benefits that contribute to lifetime fitness, even if the prey is killed (individual reproduction, altruism). Fitness of prey emerging at the optimal emergence time may be greater than, equal to or less than initial fitness. Break-even and optimality models base predictions on the opposing effects of risk and loss of benefits. Thus, many empirically verified predictions are identical at the ordinal level although differing quantitatively. Optimality models provide novel testable predictions for the effects of initial fitness, benefits, and, for ectotherms, the rate of cooling in refuge. They predict earlier emergence for equal retainable benefits than for those lost upon death.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 375–382.     

Bite-force performance and head shape in a sexually dimorphic crevice-dwelling lizard, the common chuckwalla [Sauromalus ater (= obesus)]

The Common Chuckwalla [ Sauromalus ater (=  obesus )] is a large, sexually dimorphic lizard with a flattened head that takes refuge from predators in rock crevices. Males use their relatively large heads to bite competing males during territorial fights and to restrain females during copulation. Flattened heads with an antipredator function (i.e. seeking refuge in crevices) and enlarged heads with intrasexual competition and reproductive functions suggest possible antagonism between selective pressures on head morphology in males. To examine this hypothesis, we performed a morphometric analysis and measured the bite-force performance of 49 adult chuckwallas. Males had disproportionately wider heads than females, but did not have deeper heads. Males bit with nearly four times the force of females, consistent with the notion of sexual selection for high bite force in males. Although constrained by crevice-wedging behaviour, head depth was a good predictor of bite force in both sexes. In males, however, osteological head width also was a good predictor of bite force. These results are consistent with the hypothesis that head shape in males is under antagonistic selective pressures, which may partly explain the pattern of head shape dimorphism. The disproportionately wide head of males may reflect anatomical modifications to enhance bite force in response to sexual selection in spite of presumed constraints on head shape for crevice-wedging behaviour  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 215–222.     

A new species of Ceratozamia (Zamiaceae) from the Sierra Madre of Chiapas, Mexico, with comments on species relationships

The new species Ceratozamia vovidesii from a Pleistocene floristic refuge in southern Mexico is described and illustrated. It show an affinity with C. matudae Lundell and C. mirandae Vovides, Pérez-Farrera & Iglesias from Chiapas, but it differs from them in leaf, male and female cones, and trunk morphology.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 393–400.     

Behaviour and stress response during capture and handling of the red-billed chough Pyrrhocorax pyrrhocorax (Aves: Corvidae)

We studied the effect of capture and handling on free-living red-billed choughs Pyrrhocorax pyrrhocorax . We analysed the association between bird behavioural response and individual health, breeding status, sex, and age. Active responses (aggression towards the ringer, screaming) were more common in the breeding period, and in individuals with a high heterophils to lymphocytes ratio, indicating poor physiological condition and/or high chronic stress. Adults were more aggressive than juveniles and yearlings, and females were more aggressive than males. Sex, age, and condition differences were also recorded in the spectrotemporal output of distress calls. Birds with a screaming/active response appeared to be more stress-susceptible than passive and silent individuals, and this response was stronger during the energy demanding period of reproduction. The results obtained suggest that the response of the red-billed chough during capture might primarily reflect stress-susceptibility, although a number of potential alternative explanations are discussed.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 846–855.     

Suboptimal thermoregulation in male adders (Vipera berus) after hibernation imposed by spermiogenesis

In ectotherms, the main behavioural option for thermoregulation is the adjustment of daily and seasonal activity to the thermal quality of the environment. While active, ectotherms thermoregulate by shuttling in between thermally differing microhabitat patches. Here, we focused on the question of whether other behavioural or physiological processes could force ectotherms to maintain activity during thermally unfavourable periods, when accurate thermoregulation is impossible. Using laboratory experiments and field data we compared the thermoregulation of male adders ( Vipera berus ) between two periods in spring when (1) only males and (2) also females and juveniles had terminated their winter hibernation. We found that males thermoregulated actively both in the lab and in the field. Accurate thermoregulation was only possible during the second period because of the low thermal quality of the environment. Male adders maintained a lower mean body temperature in the field than in the laboratory within both periods, and in addition their body temperature during the first period was on average 4 °C lower than during the second period. The thermal qualities of the natural basking sites showed a similar pattern. We discuss the results in the context of a potential trade-off between spermiogenesis and thermoregulation, where the benefits of early spermiogenesis coupled with inaccurate thermoregulation are higher than the associated costs. The results support the contention that the earlier spring emergence of the male compared with female adders is explainable by natural selection favouring early initiation of spermiogenesis, and hence sex differences in phenology.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 19–27.     

Divergent morphologies, performance, and escape behaviour in two tropical rock-using lizards (Reptilia: Scincidae)

The present study quantified microhabitat use, morphology, performance (sprinting, climbing, clinging, and jumping), and escape behaviour of two closely related tropical rock-using lizards. Specifically, the study tested whether: (1) a flatter body and longer limbs enhance performance in rocky habitats; (2) escape behaviour supports predictions based on habitat openness; and (3) there is a trade-off between sprinting and climbing performance. Despite the occupation of generally similar rocky habitats, the habitat of Carlia scirtetis was more open and composed of larger boulders with more regular surfaces, whereas the habitat of Carlia mundivensis was composed of more undergrowth and leaf litter, consisting of smaller boulders with irregular surfaces. The longer legs, flatter body, and greater sprinting and climbing ability of C. scirtetis, supports ecomorphological predictions. By contrast to predictions based on habitat openness, C. scirtetis allowed a potential threat to approach closer and ran further to a refuge than C. mundivensis , suggesting that escape behaviour as determined by performance may be species-specific or decoupled in these two species. The increased sprint speed of C. scirtetis highlighted a performance trade-off, with climbing speed lagging behind that of sprint speed. These results suggest that subtle differences in the structural microhabitat and the degree of habitat openness may ultimately result in substantial differences in morphology, performance, and threat behaviour in closely-related lizard species.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 85–98.     

Stable isotope niche differentiation in sticklebacks with symmetric and asymmetric pectoral fins

Fluctuating asymmetry (FA) is often, but controversially, viewed as an indicator of fitness and a target of selection. In the brook stickleback, Culaea inconstans (Kirtland), FA of the pectoral fins, which are the main source of propulsion, is inversely correlated with fecundity. We examined the hypothesis that asymmetry of the pectoral fins could affect locomotion in such a way as to influence foraging and niche use in prereproductive brook stickleback. Nitrogen and carbon stable isotope analysis showed the diet of symmetric and asymmetric males diverged with increasing body size. Larger symmetric males fed at higher trophic levels and had a diet based on carbon emanating from a more pelagic source than their asymmetric counterparts. Such effects were not observed in females or smaller males. The number of chironomid larvae found in the gut was greater on average in asymmetric than symmetric fish. The results from this study strongly suggest FA of pectoral fins affects the foraging behaviour of C. inconstans and that stable isotope analyses of individual phenotypes provides a useful tool for assessing the ecological consequences of FA.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 617–623.     

Leaf teeth in certain Salicaceae and 'Flacourtiaceae'

The leaf teeth glands in four taxa from Salicaceae and six from 'Flacourtiaceae' were examined using both light and scanning electron microscopes. There appears to be a progression from glands of simple structure in the flacourtiaceous taxa and a tendency to a more complicated development in morphology and anatomy of the salicaceous species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 241–256.     

Survival of male scarlet-tufted malachite sunbirds (Nectarinia johnstoni) on Mount Kenya and the influence of ornamentation

A prediction of handicap and signalling theories is that there should be a positive relationship between ornament size and viability. This prediction was tested in scarlet-tufted malachite sunbirds ( Nectarinia johnstoni ). These birds are endemic to the high-altitude zones of East African mountains. They are bright iridescent green with elongated central tail feathers and red pectoral tufts. The elongated tail is believed to be a mate choice cue, and the red tufts play a role in aggressive encounters between males. As adults, they feed almost exclusively on the nectar of Lobelia telekii flowers, which males defend from conspecifics. The long term survival of individually identifiable males colour-marked in 1989 and 1990 was investigated. Territorial adult males had an average annual survival rate of 72%, which is at the top of the range reported for tropical birds. The effects of morphology and territory characteristics on male survival were examined by adding covariates to the survival model. This analysis showed that male N. johnstoni with longer tails had a higher probability of survival than did short-tailed males. There was no apparent relation between tuft size or the number of L. telekii inflorescences on the territory and survival. The results presented here suggest that long-tailed males may be of intrinsically higher quality than are shorter tailed males. If this is true then females may benefit by using this trait as a mate choice cue through increased viability of their offspring.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 125–133.     

Direct and indirect selection in moth pheromone evolution: population genetical simulations of asymmetric sexual interactions

Female moths generally use pheromones to attract males. Normally, all females in a population produce a specific chemical blend with only a limited variance, and the local males are highly attracted to this blend. To better understand the direct and indirect selective forces acting on this communication system, where, unusually, it is the reproductively limited sex that signals for matings, a population genetical model has been constructed and numerically analysed. Basic to the model is the assumption that the pheromone attraction system functions asymmetrically, leading to strong sexual selection between males but no direct sexual selection between females. Evolutionary simulations using the model show that sexual selection in males causes an indirect stabilizing selection on the pheromone blends produced by females. Thus, a more narrow range of pheromone variation is selected for, even in the absence of female sexual selection. The strength of the selection is analysed, and it is suggested that this indirect stabilizing selection becomes particularly important in situations where geographically adjacent populations have evolved different pheromone blends.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 117–123.     

An inquiline fig wasp using seeds as a resource for small male production: a potential first step for the evolution of new feeding habits?

The processes allowing evolutionary transitions in resources used by parasitic wasps are largely unknown. Microhymenopteran communities associated with figs could provide a model system to investigate such transitions. We investigate here a species of genus Idarnes . The larvae generally develop as inquilines of the pollinating wasp larvae. However, in figs where the parasitic pressure is high, eggs are laid in developing seeds. These eggs turn into small males. This is the first report of seed consumption by a fig wasp. Using an alternative resource to produce small males could provide a pathway to select for increased ability to use this resource and hence provide an intermediate step for evolving the capacity to use new resources.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 9–17.     

Chromosome number, karyotype, and taxonomic considerations on the enigmatic Sellocharis paradoxa Taubert (Leguminosae, Papilionoideae, Genisteae)

Until recently, Sellocharis paradoxa Taubert, the only species of this Genisteae genus, was known solely by the isotypes. Recent new collections in Rio Grande do Sul, southern Brazil, have enabled data on chromosome number and karyotype morphology to be obtained for the first time. S. paradoxa has 2 n  = 20 chromosomes, and a bimodal asymmetrical karyotype, composed of one pair of long ( c . 6.3 µm) metacentric, five pairs of shorter acrocentric, and four pairs of shorter telocentric chromosomes ranging from c . 3.7 to 2.7 µm. The chromosome number and karyotype morphology of S. paradoxa do not fit into the Genisteae pattern. Existing information is so far insufficient to answer evolutionary questions about its origin and phylogenetic relationships, but the uniqueness of this taxon, first indicated by its peculiar leaf arrangement, and now supported by its uncommon karyological constitution, strongly suggests that the suprageneric taxonomic position of S. paradoxa should be re-evaluated.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 223–226.     

Functional basis for sexual differences in bite force in the lizard Anolis carolinensis

In many species of lizards, males attain greater body size and have larger heads than female lizards of the same size. Often, the dimorphism in head size is paralleled by a dimorphism in bite force. However, the underlying functional morphological basis for the dimorphism in bite force remains unclear. Here, we test whether males are larger, and have larger heads and bite forces than females for a given body size in a large sample of Anolis carolinensis . Next, we test if overall head shape differs between the sexes, or if instead specific aspects of skull shape can explain differences in bite force. Our results show that A. carolinensis is indeed dimorphic in body and head size and that males bite harder than females. Geometric morphometric analyses show distinct differences in skull shape between males and females, principally reflecting an enlargement of the jaw adductor muscle chamber. Jaw adductor muscle mass data confirm this result and show that males have larger jaw adductors (but not jaw openers) for a given body and head size. Thus, the observed dimorphism in bite force in A. carolinensis is not merely the result of an increase in head size, but involves distinct morphological changes in skull structure and the associated jaw adductor musculature.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 111–119.     

The selective basis for increased egg retention: early incubation temperature determines hatchling phenotype in wall lizards (Podarcis muralis)

The thermal environment during development influences many aspects of the phenotype of hatchling reptiles. We hypothesized that temperature should differentially affect early incubation stages, in which differentiation dominates over growth, and late incubation stages, characterized by high growth rates. To test this idea, we incubated eggs of wall lizard ( Podarcis muralis ) under three regimes with the same mean temperature (29 °C), one constant and two variable with opposite sequences: first cold (25 °C) and then hot (32 °C), and vice versa. Hatchlings incubated at high temperature during the initial period had shorter hindlimbs and tails than those incubated under the other two temperature regimes and shorter heads than those incubated initially at low temperature. Thus, temperature experienced by embryos during the early external incubation period produced similar phenotypic responses compared to those reported in previous studies for the same constant temperature applied over the whole incubation period. Because female wall lizards select lower body temperatures during pregnancy, an increase of intrauterine retention would extend the time of exposure of developing embryos to suitable temperatures. Diminution of body temperature during pregnancy is contrary to the expected pattern under the hypothesis that egg retention has evolved to accelerate development, as proposed by the cold-climate model for evolution of viviparity in squamates, and the results of the present study support the alternative hypothesis of developmental optimization as a special case of the broader maternal manipulation view.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 441–447.     

Sarcinomyces sideticae, a new black yeast from historical marble monuments in Side (Antalya, Turkey)

A new species of Sarcinomyces , a melanized fungus with meristematic development, is described. On the basis of small subunit and internal transcribed sequence regions of ribosomal DNA sequencing, the new species was accommodated in the anamorph genus Sarcinomyces (Herpotrichiellaceae, Chaetothyriales) close to S. petricola .  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 373–380.     

A new genus and species of Cyphophthalmi (Arachnida: Opiliones) from the north-eastern states of India

We describe a new genus and new species of Cyphophthalmi from north-eastern India, from the unexplored and extremely wet region lying between China, Bangladesh, Myanmar, and Bhutan. Meghalaya annandalei   sp. nov. , from the Mehao Wildlife Sanctuary, Arunāchal Pradesh, is an interesting Cyphophthalmi, with eyes located anterior to the ozophore, a modified tibia of leg III of males, and a robust subtriangular adenostyle. The new genus has overall resemblance to the members of the family Stylocellidae, but also has important differences for other characters, which resemble those of the members of the other tropical families Ogoveidae and Troglosironidae. The discovery of additional specimens belonging to other undescribed species indicates the need for arachnological research in that part of south-eastern Asia.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 663–670.     

Euphrasia amurensis, E. hirtella var. ramosa and var. karoiana (Orobanchaceae)

The lectotype for Euphrasia hirtella var. ramosa is selected. This taxon and E. hirtella var. karoiana are compared with E. amurensis and confirmed as synonyms, corroborating Juzepcuk (1955 ).  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 215–223.     

Two new taxa and a new combination for Stipa (Gramineae: Stipeae) in Tunisia

Two new taxa in Stipa L. Sect. Leiostipa Dumort., S. alba and S. letournexii ssp. ignea , are described from southern Tunisia (northern Africa). In addition, the S. letournexii complex is reviewed and a new combination, S. letournexii ssp. tunetana (H.Scholz) F.M. Vázquez, is presented.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 439–444.     

Covariation between predation risk, body size and fin elaboration in the green swordtail, Xiphophorus helleri

Natural and sexual selection can have either opposing or synergistic effects on the evolution of traits. In the green swordtail Xiphophorus helleri , sexual selection arising from female choice is known to favour larger males and males with longer swords. We examined variation in male and female size and fin morphology among 15 populations that varied in their predation environments. Males and females from populations in which piscivorous fishes were present had longer and deeper bodies than did males and females from populations in which piscivorous fishes were absent. Controlling for a positive effect of body size on sword length, males from populations in which piscivores were present had relatively shorter swords than did males from populations in which piscivores were absent. The associations between morphology and predation environment may be due to direct effects of predation, indirect effects of predation, other sources of selection that covary with predator presence, or other environmental effects on trait expression. These results suggest that while sexual selection favours longer swords, natural selection may have an opposing effect on sword length in populations with predators. Natural selection on body size, however, may act synergistically with sexual selection in populations with predators; both may favour the evolution of larger body size. The body size results for X. helleri contrast with related taxa that have become model systems for the study of life history evolution.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 87–100.