Experimental manipulation reveals a trade‐off between weapons and testes

Theory predicts a trade‐off between sexually selected weapons used to secure mates and post‐copulatory traits used to maximize fertilization success. However, individuals that have a greater capacity to acquire resources from the environment may invest more in both pre‐ and post‐copulatory traits, and trade‐offs may not be readily apparent. Here, we manipulate the phenotype of developing individuals to examine allocation trade‐offs between weapons and testes in Mictis profana (Hemiptera: Coreidae), a species where the hind legs are sexually selected weapons used in contests over access to females. We experimentally prevented males from developing weapons by inducing them to autotomize their hind legs before the final moult to adulthood. We compared trait expression in this group to males where autotomy was induced in the mid‐legs, which are presumably not under sexual selection to the same extent. We found males without weapons invested proportionally more in testes mass than those with their mid‐legs removed. Males that developed to adulthood without weapons did not differ from the mid‐leg removal group in other traits potentially under precopulatory sexual selection, other post‐copulatory traits or naturally selected traits. In addition, a sample of adult males from the same population in the wild revealed a positive correlation between investment in testes and weapons. Our study presents a critical contribution to a growing body of literature suggesting the allocation of resources to pre‐ and post‐copulatory sexual traits is influenced by a resource allocation trade‐off and that this trade‐off may only be revealed with experimental manipulation.     

The evolution of symmetrical snapping in termite soldiers need not lead to reduced chemical defence

Termite soldiers use a multitude of defensive strategies, often combining mechanical weapons with defensive chemicals secreted from their frontal gland. In higher termites (Termitidae), both these weapons are situated in the head of the soldiers. Their simultaneous occurrence is supposed to be subjected to energetic and spatial constraints and the resulting trade‐off often leads to a full development of only one of the weapons and reduction of the other. Mandibular snapping represents an extreme and efficient anatomical adaptation, allowing the soldier to knock out arthropod enemies with a violent strike of elongated mandibles. The head of snapping soldiers harbours massive adductor muscles while the frontal gland is reported to be reduced and the chemical defence absent. Here we show that the symmetrical snapping soldiers of the Neotropical termite Cavitermes tuberosus possess a well‐developed frontal gland situated in the frontal projection on their head. The gland produces a blend of unbranched alkenes and alkadienes combined with three diterpene hydrocarbons, derived from a novel bicyclic hydrocarbon skeleton. We characterized the molecular structure of the most abundant of these diterpenes and proposed for it a trivial name, cavitene. We conclude that the evolution of the mandibular snapping has not necessarily been accompanied by the reduction or loss of chemical defensive adaptations and that the two defensive modes are not mutually incompatible. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●● , ●●–●●.     

Patrick Matthew's law of natural selection

Patrick Matthew is the little‐known first originator of macroevolution by natural selection. I review his ideas, and introduce some previously unnoticed writings (catalogued at a new website: http://smarturl.it/patrickmatthew ) that clarify how they differ from Darwin's and Wallace's. Matthew's formulation emphasized natural selection as an axiomatic ‘law’ rather than a ‘theory’, a distinction that could still be of use to us today. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●● , ●●–●●.     

Temperature influences host instar selection in an aphid parasitoid: support for the relative fitness rule

The relative fitness rule states that parasitoid females should adopt risk‐prone reproductive behaviours when expecting low reproductive success. Temperature influences the reproductive success of insects by affecting their basal metabolic rate during development, their egg load at emergence, and their life expectancy as adults. Using an aphid–parasitoid model system, we investigated the influence of developmental and adult temperature on the risk‐sensitive decision‐making of females. We considered the use of a low‐quality host nymphal instar by the aphid parasitoid Aphidius ervi to be a risk‐prone behaviour. Immature females were reared at 12, 20 or 28 °C and had access to the four nymphal instars of the potato aphid Macrosiphum euphorbiae for oviposition at one of these temperatures. Host selection behaviour was continuously recorded during exploitation of an aphid patch. We observed that warm‐developed females and parasitoids foraging at high temperature attacked low‐quality hosts more frequently than females from other treatments. These results support the hypothesis that a decrease in expected parasitoid reproductive success resulted in risk‐prone behaviours. To our knowledge, this is the first experimental evidence suggesting that temperature influences host stage selection and risk‐sensitive making decision in parasitoids, and the present study is the first to support the relative fitness rule. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●●, ●●–●●.     

Does whole‐organism performance constrain resource use? A community test with desert lizards

Discovering the mechanisms by which communities of co‐existing species exist has proven to be one of the greatest challenges for evolutionary ecologists. A recent perspective emphasizes the role of functional traits, such as whole‐organism performance, as key limiting factors in the evolution of communities, yet few studies have examined this possibility. We examine how bite force and morphology influence the ability of ten lizard species in a single community to access insect prey, as defined by prey type and prey hardness. We gathered over 3 years of data from a desert lizard community comprised of ten species and found significant variation among species for bite force and prey hardness, as well as significant differences in performance and niche breadth for each species. In general, higher levels of absolute bite force broadens resource accessibility (sizes of prey), and does not generally result in a reduced ability to access smaller prey. For example, large lizard species that are hard biters can still consume soft prey. On the other hand, small lizard species that are weak biters are more limited in their ability to access hard prey, although the overall decline in resource accessibility is modest. Our findings highlight how functional traits can influence which species can access key resources within a community of similar species. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●● , ●●–●●.     

Does developmental acclimatization reduce the susceptibility to predation in newt larvae?

Many organisms respond to the heterogeneity of abiotic environmental conditions by plastic modifications of their phenotypes (acclimation or acclimatization). Despite considerable research efforts in this area, the beneficial (adaptive) effect of acclimation or acclimatization is still debated. We examined whether the development of newt larvae (Ichthyosaura alpestris) under different natural light and thermal conditions subsequently altered their susceptibility to predation in sun‐exposed versus shaded tanks in nature. During predation trials in various light and temperature conditions, newt larvae that developed in sun‐exposed warmer tanks consistently suffered from higher predation by dragonfly nymphs (Aeshna cyanea) compared to larvae from shaded or colder tanks. We conclude that higher sun exposure during embryonic and larval development negatively affects antipredator performance even in sun‐exposed tanks: this result is inconsistent with the beneficial acclimation hypothesis. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Size and sex matter: reproductive biology and determinants of offspring survival in Gazella marica

For prey, many behavioural traits are constrained by the risk of predation. Therefore, shifts between warning and cryptic coloration have been suggested to result in parallel changes in several behaviours. In the present study, we tested whether changes in chromatic contrast among eight populations of the strawberry poison‐dart frog, Dendrobates pumilio, co‐vary with behaviour, as expected if selection is imposed by predators relying on visual detection of prey. These eight populations are geographically isolated on different island in the Bocas del Toro region of Panama and have recently diverged morphologically and genetically. We found that aggression and explorative behaviour were strongly correlated and also that males tended to be more aggressive and explorative if they belonged to populations with conspicuously coloured individuals. We discuss how evolutionary switches between predator avoidance strategies and associated behavioural divergence between populations may affect reproductive isolation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

A further look at the 1858 Wallace–Darwin mail delivery question

Recent investigations have led to a conclusion that Alfred Russel Wallace probably mailed his ‘Ternate’ paper on natural selection to Darwin 1 month later than some have assumed, thus freeing Darwin from possible accusations of plagiarism. Further examination of the question suggests that this conclusion is premature because the evidence in favour of the later mailing date appears to be weaker than first considered. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Steneosaurus edwardsi (Thalattosuchia: Teleosauridae), the largest known crocodylomorph of the Middle Jurassic

Teleosaurids were a clade of marine crocodylomorphs that were globally distributed during the Jurassic Period. They evolved a wide range of body sizes, from small (～2–3 m) to very large (> 9 m). Until now, the largest known Middle Jurassic teleosaurid was ‘Steneosaurus’ obtusidens, from the Oxford Clay Formation of the UK. Here, we re‐examine a very large Oxford Clay specimen (ilium, ischium, and femur) that had been tentatively attributed to ‘S.’ obtusidens. Based on comparative anatomical study with the ‘S.’ obtusidens holotype and referred specimens of Steneosaurus edwardsi and Steneosaurus leedsi, we conclude that this very large individual actually pertains to S. edwardsi. Based on comparisons with the Machimosaurus mosae neotype (which has a complete femur and skeleton), we estimate a total length in excess of 7 m for this large S. edwardsi individual, making it the largest known Middle Jurassic teleosaurid. Therefore, along with the closely related genus Machimosaurus, this clade of large‐bodied Middle–Late Jurassic teleosaurids were the largest species during the first 100 million years of crocodylomorph evolution. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●● , ●●–●●.     

Are viviparous lizards from cool climates ever exclusively nocturnal? Evidence for extensive basking in a New Zealand gecko

Viviparity has evolved numerous times among squamate reptiles; however, the combination of viviparity and nocturnality is apparently rare among lizards. We used time‐lapse photography to examine evidence for diurnal activity in a viviparous lizard often described as nocturnal, the gecko Woodworthia ‘Otago/Southland’ from southern New Zealand (family Diplodactylidae). Evidence for diurnal emergence was extensive. Females have a higher incidence of basking compared to males, although no difference was detected between females in different reproductive conditions. Temperature loggers inserted into calibrated copper models were used to compare the body temperatures available to geckos in two basking positions and in two retreat types. Models in basking positions reached higher mean temperatures than models in retreats, although there was no significant effect of basking position or retreat type on model temperatures. Collectively, our results indicate that pregnant geckos that bask consistently could reduce gestation length by at least 14 days compared with females that remain in retreats. Extensive basking in this species adds to the growing evidence of diurno‐nocturnality in many New Zealand lepidosaurs, including other viviparous geckos. Our results lead us to question whether viviparity in lizards is ever compatible with ‘pure’ nocturnality in a cool climate. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●● , ●●–●●.     

Once a land of big wild rivers: specialism is context‐dependent for riparian snails (Pulmonata: Valloniidae) in central Europe

Specialist species may be perceived as such because of their narrow ecological requirements, but this may be context‐dependent. The genus Vallonia (Gastropoda Pulmonata: Valloniidae) includes widespread generalist species and also two specialists endemic to Central Europe: Vallonia suevica, restricted to warm, wet meadows and riverbanks subject to seasonal flooding; and Vallonia declivis, living only in wet to humid meadows, riverbanks, and reedbeds. Both have experienced dramatic declines; as is the case with many land snails, their global conservation status has been underestimated: these species are Critically Endangered. Other congenerics are probably dispersed by birds. In contrast, the distributions of these meadow specialists appear to be the outcome of their strictly riparian habitat coupled with dispersal by fish. Thus, they have tracked drainage changes through the Pleistocene from their origin in the floodplain uplands of the Danube biodiversity hotspot in the Pliocene. Natural dispersal mechanisms have been disrupted, and riparian and river ecosystems have been destroyed throughout Europe. This has led big‐river specialist molluscs and their associated fishes to the brink of extinction. The notion of specialism thus depends on the ecological context; it is useful to stress their non‐invasive character, current restriction to scarce habitats, and evolution under quite different conditions. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●● , ●●–●●.     

Into the light: diurnality has evolved multiple times in geckos

Geckos are the only major lizard group consisting mostly of nocturnal species. Nocturnality is presumed to have evolved early in gecko evolution and geckos possess numerous adaptations to functioning in low light and at low temperatures. However, not all gecko species are nocturnal and most diurnal geckos have their own distinct adaptations to living in warmer, sunlit environments. We reconstructed the evolution of gecko activity patterns using a newly generated time‐calibrated phylogeny. Our results provide the first phylogenetic analysis of temporal activity patterns in geckos and confirm an ancient origin of nocturnality at the root of the gecko tree. We identify multiple transitions to diurnality at a variety of evolutionary time scales and transitions back to nocturnality occur in several predominantly diurnal clades. The scenario presented here will be useful in reinterpreting existing hypotheses of how geckos have adapted to varying thermal and light environments. These results can also inform future research of gecko ecology, physiology, morphology and vision as it relates to changes in temporal activity patterns. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●●, ●●–●●.     

Males that drop a sexually selected weapon grow larger testes

Costly sexually selected weapons are predicted to trade off with postcopulatory traits, such as testes. Although weapons can be important for achieving access to females, individuals of some species can permanently drop (i.e. autotomize) their weapons, without regeneration, to escape danger. We capitalized on this natural behavior to experimentally address whether the loss of a sexually selected weapon leads to increased testes investment in the leaf‐footed cactus bug, Narnia femorata Stål (Hemiptera: Coreidae). In a second experiment, we measured offspring production for males that lost a weapon during development. As predicted, males that dropped a hind limb during development grew significantly larger testes than the control treatments. Hind‐limb autotomy did not result in the enlargement of other nearby traits. Our results are the first to experimentally demonstrate that males compensate for natural weapon loss by investing more in testes. In a second experiment we found that females paired with males that lost a hind limb had 40% lower egg hatching success than females paired with intact males, perhaps because of lower mating receptivity to males with a lost limb. Importantly, in those cases where viable offspring were produced, males missing a hind limb produced 42% more offspring than males with intact limbs. These results suggest that the loss of a hind‐limb weapon can, in some cases, lead to greater fertilization success.     

Post‐glacial colonization of eastern Europe from the Carpathian refugium: evidence from mitochondrial DNA of the common vole Microtus arvalis

There is now considerable evidence for the survival of temperate species within glacial refugia that were situated at relatively high latitudes, notably the Carpathian Basin and Dordogne region in Europe. However, the prevalence of fossil remains in such locations is rarely matched by molecular evidence for their contribution to subsequent geographical and demographic expansion of the species in question. One obstacle to this has been insufficient analysis of modern samples from the relevant areas, in particular the parts of eastern Europe that surround the Carpathian refugium. In the present study, we examine the patterns of variation in mitochondrial DNA of the common vole (Microtus arvalis), obtained from existing museum specimens and from newly‐collected samples obtained in this area. We show that common voles from one of six extant mitochondrial DNA lineages have colonized most of the species' range in eastern Europe. We contend that the post‐glacial dispersal of this lineage most likely originated from the Carpathian refugium, adding support to the argument that such northern refugia made an important contribution to existing genetic diversity in Europe. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●● , ●●–●●.     

The effects of substratum on locomotor performance in lacertid lizards

Locomotion is important to animals because it has direct implications for fitness through its role in predator escape, prey capture, and territory defence. Despite significant advances in our understanding of animal locomotion, studies exploring how substrate properties affect locomotor performance remain scant. In the present study, we explore how variation in substrate (sand, slate, cork) affects locomotor performance in lacertid lizards that differ in morphology. Moreover, we explore whether substrate effects are the same for different types of locomotor performance (speed, acceleration, and stamina). Our results show that the substrate affected most types of locomotor performance studied but not always in the same way. Although substrate effects were species‐dependent for the maximal speed over 50 cm and the distance run to exhaustion, this was not the case for acceleration capacity. These results suggest that substrate texture differentially affects burst performance vs. longer duration measures of locomotor performance. Finally, straightforward relationships between habitat use and the substrate on which performance was maximized were not observed. This suggests that the evolution of locomotor capacity is complex and that animals may show compromise phenotypes allowing them to deal with a variety of substrates in their natural environment. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●●, ●●–●●.     

Ontogenetic shifts in resource allocation: colour change and allometric growth of defensive and reproductive structures in the Caribbean spiny lobster Panulirus argus

Resource allocation theory predicts a disproportionately large allocation of resources to defensive structures during early ontogeny in organisms that are subject to more intense predation at smaller than at larger body sizes. We tested this prediction on the Caribbean spiny lobster Panulirus argus, which exhibits a negative relationship between predation risk and body size with a high natural mortality of smaller individuals. Independent allometric growth analyses demonstrated that numerous defensive structures (e.g. orbital horns, segments supporting the antenna, the tail fan) display negative allometric growth throughout ontogeny. We interpret these findings as lobsters investing disproportionately more resources to defensive structures when small to improve survivorship. Similarly, we observed an ontogenetic shift in lobster colour pattern; small individuals (< 23 mm carapace length) that inhabit nursery grounds (preferably among red algae) displayed a disruptive pattern (camouflage), whereas larger juveniles displayed a bicolour pigmentation typical of adult lobsters. This shift in colour pattern further suggests that small lobsters employ cryptic coloration throughout their asocial algal stage. However, this cryptic coloration offers no advantage when lobsters grow larger and start dwelling in crevices. Other structures linked to reproduction (e.g. female pleopods and male pereopods) experienced either isometric or positive allometric growth throughout ontogeny. Our results support one of the main predictions of resource allocation theory and demonstrate ontogenetic shifts in defensive structures and coloration concomitantly with changes in lobster mortality risk mediated by size‐dependent predation risk. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Resource quality affects weapon and testis size and the ability of these traits to respond to selection in the leaf‐footed cactus bug,Narnia femorata

The size of weapons and testes can be central to male reproductive success. Yet, the expression of these traits is often extremely variable. Studies are needed that take a more complete organism perspective, investigating the sources of variation in both traits simultaneously and using developmental conditions that mimic those in nature. In this study, we investigated the components of variation in weapon and testis sizes using the leaf‐footed cactus bug, Narnia femorata (Hemiptera: Coreidae) on three natural developmental diets. We show that the developmental diet has profound effects on both weapon and testis expression and scaling. Intriguingly, males in the medium‐quality diet express large weapons but have relatively tiny testes, suggesting complex allocation decisions. We also find that heritability, evolvability, and additive genetic variation are highest in the high‐quality diet for testis and body mass. This result suggests that these traits may have an enhanced ability to respond to selection during a small window of time each year when this diet is available. Taken together, these results illustrate that normal, seasonal fluctuations in the nutritional environment may play a large role in the expression of sexually selected traits and the ability of these traits to respond to selection.     

Very fine‐scale population genetic structure of sympatric asterinid sea stars with benthic and pelagic larvae: influence of mating system and dispersal potential

The present study investigated the fine‐scale population genetic structure of sympatric asterinid sea stars with contrasting modes of larval development (benthic versus pelagic). Parvulastra exigua lacks a dispersive life phase yet is one of the worlds most widely distributed and abundant sea stars, whereas Meridiastra calcar, a sea star with a dispersive larva, has a more limited regional scale distribution. Populations of P. exigua sampled from tide pools on three adjacent headlands showed significant genetic substructure (mitochodrial DNA control region) at fine spatial scales (tide pools < 300 m apart: F_ST = 0.249, P < 0.01; headlands 5–15 km apart: F_ST = 0.125, P = 0.04). As expected, M. calcar populations sampled from the same headlands did not exhibit significant genetic structuring (F_ST = 0.029, P = 0.14). The life‐history traits of P. exigua, a mixed mating system (selfing + outcrossing), pseudocopulation among closely‐related conspecifics, and an entirely benthic life cycle with a philopatric larva, undoubtedly influence its strong genetic structure across fine spatial scales. Localized genetic structure, especially at the very fine‐scale of tide pools, would not be detected in the more typical regional scale approaches adopted by most studies of marine invertebrate populations. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, ●●, ●●–●●.     

Phylogenetic systematics of Erythronium (Liliaceae): morphological and molecular analyses

For a nearly complete set of species of Erythronium (Liliaceae), we examined two plastid loci (the rps16 intron and the 5′ trnK intron, excluding the matK exon), one nuclear locus (nrITS) and morphology to evaluate species relationships and that of Erythronium to Amana, the putatively most closely allied genus. A matrix of morphological characters was developed through observation of around 900 living and herbarium specimens; evolution of these was examined using character optimization on the combined (total‐evidence) tree. Parsimony methods were used to examine the morphological and molecular data sets produced, both separately and in combination, with Bayesian methods also used on the molecular data sets. These established that the genus is probably sister to Amana (although most analyses placed Amana inside Erythronium) and that Tulipa is sister to the pair of Erythronium and Amana. Within Erythronium, there are three strongly supported geographically distinct clades: (1) Eurasian and (2) eastern and (3) western North American. Separation of species in these three clades is less clear, particularly among the western North American taxa. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ●●, ●●–●●.     

Length–mass allometry in snakes

Body size and body shape are tightly related to an animal's physiology, ecology and life history, and, as such, play a major role in understanding ecological and evolutionary phenomena. Because organisms have different shapes, only a uniform proxy of size, such as mass, may be suitable for comparisons between taxa. Unfortunately, snake masses are rarely reported in the literature. On the basis of 423 species of snakes in 10 families, we developed clade‐specific equations for the estimation of snake masses from snout–vent lengths and total lengths. We found that snout–vent lengths predict masses better than total lengths. By examining the effects of phylogeny, as well as ecological and life history traits on the relationship between mass and length, we found that viviparous species are heavier than oviparous species, and diurnal species are heavier than nocturnal species. Furthermore, microhabitat preferences profoundly influence body shape: arboreal snakes are lighter than terrestrial snakes, whereas aquatic snakes are heavier than terrestrial snakes of a similar length. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.