Biological Journal of the Linnean Society

BRABY, MICHAEL F.; EASTWOOD, RODNEY; MURRAY, NEIL

doi: 10.1111/j.1095-8312.2012.01909.xpmid: N/A

AbstractSubspecies lie at the interface between systematics and population genetics, and represent a unit of biological organization in zoology that is widely used in the disciplines of taxonomy and conservation biology. In this review, we explore the utility of subspecies in relation to their application in systematics and biodiversity conservation, and briefly summarize species concepts and criteria for their diagnosis, particularly from an invertebrate perspective. The subspecies concept was originally conceived as a formal means of documenting geographical variation within species based on morphological characters; however, the utility of subspecies is hampered by inconsistencies by which they are defined conceptually, a lack of objective criteria or properties that serve to delimit their boundaries, and their frequent failure to reflect distinct evolutionary units according to population genetic structure. Moreover, the concept has been applied to populations largely comprising different components of genetic diversity reflecting contrasting evolutionary processes. We recommend that, under the general lineage (unified) species concept, the definition of subspecies be restricted to extant animal groups that comprise evolving populations representing partially isolated lineages of a species that are allopatric, phenotypically distinct, and have at least one fixed diagnosable character state, and that these character differences are (or are assumed to be) correlated with evolutionary independence according to population genetic structure. Phenotypic character types include colour pattern, morphology, and behaviour or ecology. Under these criteria, allopatric subspecies are a type of evolutionarily significant unit within species in that they show both neutral divergence through the effects of genetic drift and adaptive divergence under natural selection, and provide an historical context for identifying biodiversity units for conservation. Conservation of the adaptedness and adaptability of gene pools, however, may require additional approaches. Recent studies of Australian butterflies exemplify these points.

DOCKX, CRISTINA

doi: 10.1111/j.1095-8312.2012.01916.xpmid: N/A

AbstractThe eastern North American population of the monarch butterfly (Danaus plexippus plexippus) has different migratory routes. The majority fly to overwintering colonies in Mexico and others take an eastern route through Florida and Cuba. Monarchs migrating through Florida–Cuba do not overwinter and are mostly found nectaring and flying close to vegetation. This present study explores whether Florida–Cuba versus Mexican migrants differ in (1) phenotypic traits important for migration (e.g. wing size and condition, lipid and lean mass content, and reproductive status) and (2) migratory strategies. The monarch natal grounds (e.g. migrants versus residents) were determined through thin-layer chromatography cardenolide fingerprint and stable isotopes (hydrogen δ2H and carbon δ13C). In addition, wing size and condition, lipid and lean mass, and reproductive status were determined. The results suggest that Mexican migrants are better suited for longer sustained flights and successful overwinter periods as a result of larger wings in better condition, reproductive diapause, and significant fat content. By contrast, Florida–Cuba migrants are more suited for shorter flights and opportunistic migratory strategies, given that their wings were in poor condition, as well as the active reproductive status of > 50% of these butterflies and their significantly low fat content. Eastern monarch migration is more complex and diverse than previously assumed.

IHALAINEN, EIRA; LINDSTEDT, CARITA

doi: 10.1111/j.1095-8312.2012.01922.xpmid: N/A

AbstractSeasonal polyphenism in animal colour patterns indicates that temporal variation in selection pressures maintains phenotypic plasticity. Spring generation of the polyphenic European map butterfly Araschnia levana has an orange–black fritillary-like pattern whilst individuals of the summer generation are black with white bands across the wings. What selects for the colour difference is unknown. Because predation is a major selection pressure for insect coloration, we first tested whether map butterfly coloration could have a warning function (i.e. whether the butterflies are unpalatable to birds). In a following field experiment with butterfly dummies we tested whether the spring form is better protected than the summer form from predators in the spring, and vice versa in the summer. The butterflies were palatable to birds (blue tits Cyanistes caeruleus) and in the field the spring and summer form dummies were attacked equally irrespective of season. Therefore, we found no evidence that the map butterfly is warning-coloured or that seasonal polyphenism is an adaptation to avian predation. Because insect coloration has multiple functions and map butterfly coloration is linked to morphology, life history and development it is likely that the interplay of several selection pressures explains the evolution of colour polyphenism.

YIP, ERIC C.; ROWELL, DAVID M.; RAYOR, LINDA S.

doi: 10.1111/j.1095-8312.2012.01904.xpmid: N/A

AbstractMovement among social groups interacts with the costs and benefits of group-living in complex ways. Unlike most other social spiders, the social huntsman spider, Delena cancerides, appears to enter foreign colonies, discriminates kin from non-kin, and has very limited dispersal options because their bark retreats are rare, making this species an interesting model organism with which to examine the role of inter-colony movement on group-living. We examined movement among field colonies of D. cancerides in three ways: (1) by tracking the dispersal and immigration of marked spiders into foreign colonies; (2) by recording resident spiders' behaviour toward introduced immigrants; and (3) by inferring intra-colony relatedness and immigration patterns through allozyme electrophoresis. Of the marked spiders, only young juveniles moved into neighbouring colonies, whereas subadults and adults did not. Introduced juveniles were tolerated in foreign colonies, whereas introduced adult males and subadults were usually attacked by the resident adult female, unless she had similar sized subadult/adult offspring of her own. Allozyme profiles from unmanipulated field colonies showed that 47% of sampled colonies contained at least one immigrant and that average within colony relatedness was below 0.5. These data align with previous research on the costs and benefits of group-living for D. cancerides, suggesting that spiders actively seek and regulate group membership based on interests of both the immigrant and the colony.

COLLIN, RACHEL

doi: 10.1111/j.1095-8312.2012.01908.xpmid: N/A

AbstractIntraspecific variation in egg size and hatching size, and the genetic and environmental trade-offs that contribute to variation, are the basis of the evolution of life histories. The present study examined both univariate and multivariate temperature-mediated plasticity of life-history traits, as well as temperature-mediated trade-offs in egg size and clutch size, in two planktotrophic species of marine slipper limpets, Crepidula. Previous work with two species of Crepidula with large eggs and lecithotrophic development has shown a significant effect of temperature on egg size and hatching size. To further examine the effect of temperature on egg size in Crepidula, the effects of temperature on egg size and hatching size, as well as the possible trade-offs with other the life-history features, were examined for two planktotrophic species: Crepidula incurva and Crepidula cf. marginalis. Field-collected juveniles were raised at 23 or 28 °C and egg size, hatching size, capsules/brood, eggs/capsule, time to hatch, interbrood interval, and final body weight were recorded. Consistent with results for the lecithotrophic Crepidula, egg size and hatching size decreased with temperature in the planktotrophic species. The affects of maternal identity and individual brood account for more than half of the intraspecific variation in egg size and hatching size. Temperature also showed a significant effect on reproductive rate, with time to hatch and interbrood interval both decreasing with increasing temperature. However, temperature had contrasting effects on the number of offspring. Crepidula cf. marginalis has significantly more eggs/capsule and therefore more eggs per brood at 28 °C compared to 23 °C, although capsules/brood did not vary with temperature. Crepidula incurva, on the other hand, produced significantly more capsules/brood and more eggs per brood at the lower temperature, whereas the number of eggs/capsule did not vary with temperature. The phenotypic variance–covariance matrix of life-history variables showed a greater response to temperature in C. incurva than in C. cf. marginalis, and temperature induced trade-offs between offspring size and number differ between the species. These differences suggest that temperature changes as a result of seasonal upwelling along the coast of Panama will effect the reproduction and evolution of life histories of these two co-occurring species differently.

VERMEIJ, GEERAT J.

doi: 10.1111/j.1095-8312.2012.01897.xpmid: N/A

AbstractThe extent to which animal lineages achieve large body size, a trait with broad advantages in competition and defence, varies in space and time according to the supply of (and demand for) resources, as well as the magnitude and effects of extinction. Using the maximum sizes of shallow-water marine shell-bearing molluscs belonging to nineteen guilds (groups of species with similar habits and food sources) in seven temperate regions from the Early Miocene to the Recent, the present study examined the controls on productivity and predation that enable and compel large size to evolve. The North Pacific (especially its eastern sector) has been most favourable to large-bodied species from the Pliocene onward. Large productive kelps (Laminariales) evolved there in conjunction with herbivorous mammals, setting the stage through positive feedbacks between production and consumption for the evolution of large molluscan herbivores and suspension-feeders. The evolution of bottom-feeding predatory mammals together with other large predators created intense selection for large molluscan sizes. Very large molluscs in the Early Miocene were concentrated in the southern hemisphere, especially among metabolically passive species. Extinctions, which preferentially targeted the largest members of guilds in most regions, were more numerous in the southern hemisphere and the North Atlantic than in the North Pacific. Minimal disruption, together with the early evolution of metabolically-active consumers and the positive feedbacks they engendered, accounts for the evolution of molluscan gigantism in the North Pacific.

GARDUÑO-PAZ, MONICA V.; ADAMS, COLIN E.; VERSPOOR, ERIC; KNOX, DAVID; HARROD, CHRIS

doi: 10.1111/j.1095-8312.2012.01906.xpmid: N/A

AbstractThe expression of two or more discrete phenotypes amongst individuals within a species (morphs) provides multiple modes upon which selection can act semi-independently, and thus may be an important stage in speciation. In the present study, we compared two sympatric morph systems aiming to address hypotheses related to their evolutionary origin. Arctic charr in sympatry in Loch Tay, Scotland, exhibit one of two discrete, alternative body size phenotypes at maturity (large or small body size). Arctic charr in Loch Awe segregate into two temporally segregated spawning groups (breeding in either spring or autumn). Mitochondrial DNA restriction fragment length polymorphism analysis showed that the morph pairs in both lakes comprise separate gene pools, although segregation of the Loch Awe morphs is more subtle than that of Loch Tay. We conclude that the Loch Awe morphs diverged in situ (within the lake), whereas Loch Tay morphs most likely arose through multiple invasions by different ancestral groups that segregated before post-glacial invasion (i.e. in allopatry). Both morph pairs showed clear trophic segregation between planktonic and benthic resources (measured by stable isotope analysis) but this was significantly less distinct in Loch Tay than in Loch Awe. By contrast, both inter-morph morphological and life-history differences were more subtle in Loch Awe than in Loch Tay. The strong ecological but relatively weak morphological and life-history divergence of the in situ derived morphs compared to morphs with allopatric origins indicates a strong link between early ecological and subsequent genetic divergence of sympatric origin emerging species pairs. The emergence of parallel specialisms despite distinct genetic origins of these morph pairs suggests that the effect of available foraging opportunities may be at least as important as genetic origin in structuring sympatric divergence in post-glacial fishes with high levels of phenotypic plasticity.

HEINS, DAVID C.

doi: 10.1111/j.1095-8312.2012.01907.xpmid: N/A

AbstractCausal explanations for host reproductive phenotypes influenced by parasitism fit into three broad evolutionary models: (1) non-adaptive side effect; (2) adaptive parasitic manipulation; and (3) adaptive host defence. This study demonstrates fecundity compensation, an adaptive non-immunological host defence, in the three-spined stickleback fish (Gasterosteus aculeatus) infected by the diphyllobothriidean cestode Schistocephalus solidus. Both infected and uninfected female sticklebacks produced egg clutches at the same age and size. The reproductive capacity of infected females decreased rapidly with increased parasite : host body mass ratio. Body condition was lower in infected females than uninfected females and decreased with increasing parasite : host mass ratio. Females with clutches had greater body condition than those without clutches. A point biserial correlation showed that there was a body condition threshold necessary for clutch production to occur. Host females apparently had the capacity to produce egg clutches until the prolonged effects of nutrient theft by the parasite and the drain on resources from reproduction precluded clutch formation. Clutch mass, adjusted for female body mass, did not differ significantly between infected and uninfected females. Infected females apparently maintained the same level of reproductive allotment (egg mass as proportion of body mass) as uninfected females. Infected females produced larger clutches of smaller eggs than uninfected females, revealing a trade-off between egg mass and egg number, consistent with the fecundity compensation hypothesis. The rapid loss of reproductive capacity with severity of infection probably reflects the influence of the parasite combined with a trade-off between current and future reproduction in the host. Inter-annual differences in reproductive performance may have reflected ecological influences on host pathology and/or intra-annual seasonal changes.

HETTYEY, ATTILA; RÖLLI, FRANZISKA; THÜRLIMANN, NINA; ZÜRCHER, ANNE-CATHERINE; VAN BUSKIRK, JOSH

doi: 10.1111/j.1095-8312.2012.01923.xpmid: N/A

AbstractThe ability of prey to detect predators directly affects their probability of survival. Chemical cues are known to be important for predator detection in aquatic environments, but the role of other potential cues is controversial. We tested for changes in behaviour of Rana temporaria tadpoles in response to chemical, visual, acoustic, and hydraulic cues originating from dragonfly larvae (Aeshna cyanea) and fish (Gasterosteus aculeatus). The greatest reduction in tadpole activity occurred when all cues were available, but activity was also significantly reduced by visual cues only. We did not find evidence for tadpoles lowering their activity in response to acoustic and hydraulic cues. There was no spatial avoidance of predators in our small experimental containers. The results show that anuran larvae indeed use vision for predator detection, while acoustic and hydraulic cues may be less important. Future studies of predator-induced responses of tadpoles should not only concentrate on chemical cues but also consider visual stimuli.

BARATA, MAFALDA; PERERA, ANA; MARTÍNEZ-FREIRÍA, FERNANDO; HARRIS, D. JAMES

doi: 10.1111/j.1095-8312.2012.01903.xpmid: N/A

AbstractQuedenfeldtia (Boettger, 1883) is a genus of diurnal geckos, endemic to the Atlas Mountains in Morocco, with two species being recognized: Quedenfeldtia moerens and Quedenfeldtia trachyblepharus. Quedenfeldtia moerens is found across a wide variety of habitats, from sea level to 3000 m a.s.l., whereas Q. trachyblepharus occupies exclusively high mountain regions reaching up to 4000 m a.s.l. This differentiation, offers an interesting model for study biogeographical patterns and evolutionary scenarios in a North African endemic. Analysis of two mitochondrial (12S rRNA and ND4) and four nuclear (ACM4, MC1R, PDC, and Rag1) DNA markers revealed high genetic variation, consistent with other recent phylogeographical studies, and with the two currently described species. However, within each species, a subdivision into two groups with geographical consistence was found. Multivariate morphological analyses confirmed the existence of two main phenotypes, whereas ecological niche modelling identified various environmental variables associated with the distribution of each species, and helped to predict occurrences outside the confirmed ranges. The results obtained in the present study indicate the possible existence of additional ‘cryptic’ species within this genus, a condition found in many North African reptiles, and particularly common in geckos. In general, North African montane fauna appears to reflect the occurrence of diverse palaeoendemics, as seen in Central Africa Mountain systems, rather than the pattern of recent postglacial recolonization observed in Europe.