Identifying sources of variation in reproductive and life-history traits among five populations of a Chinese lizard (Takydromus septentrionalis, Lacertidae)

Research on life-history traits of squamate reptiles has focused on North American species, while Asian taxa have been virtually ignored. In order to understand general patterns in reptile life histories, we need a broader data base. Our study on the slender-bodied lacertid lizard Takydromus septentrionalis provides the first detailed information on factors responsible for intraspecific variation in reproductive output and life history in a Chinese reptile. Clutches of recently collected lizards from five widely separated localities in China revealed major divergences in female body size at maturation, mean adult female body size, body condition after oviposition, size-adjusted fecundity, relative clutch mass, and mass and shape of eggs. Most of these geographical differences persisted when the same groups of females were maintained in identical conditions in captivity. Additionally, reproductive frequency during maintenance under laboratory conditions differed according to the animals' place of origin. Thus, the extensive geographical variation in reproductive and life-history traits that occurs within T. septentrionalis is exhibited even in long-term captives, suggesting that proximate factors that vary among localities (local conditions of weather and food supply) are less important determinants of life-history variation than are intrinsic (presumably genetic) influences. The maternal abdominal volume available to hold the clutch may be one such factor, based on low levels of variation in Relative Clutch Mass among populations, and geographical variation in the position of trade-off lines linking offspring size to fecundity.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 443–453.     

Thermoregulation during gravidity in the children's python (Antaresia childreni): a test of the preadaptation hypothesis for maternal thermophily in snakes

Pregnant squamate reptiles (i.e. lizards and snakes) often maintain higher and more stable body temperatures than their nonpregnant conspecifics, and this maternal thermophily enhances developmental rate and can lead to increased offspring quality. However, it is unclear when this behaviour evolved relative to the evolution of viviparity. A preadaptation hypothesis suggests that maternal thermophily was a preadaptation to viviparity. Oviparous squamates are unique among oviparous reptiles for generally retaining their eggs until the embryos achieve one fourth of their development. As a result, maternal thermophily by gravid squamates may provide the same thermoregulatory benefits, at least during early development, that have been associated with viviparity. Thus, the evolution of viviparity in squamates may reflect an expanded duration of a pre-existing maternal thermoregulatory behaviour. Despite its evolutionary relevance, thermoregulation during gravidity in oviparous squamates has not yet been explored in depth. In the present study, we examined whether gravidity was associated with thermoregulatory changes in the oviparous children's python, Antaresia childreni . First, we discovered that, compared to most snakes, A. childreni is at an advanced stage of embryonic development at oviposition. Second, using surgically implanted temperature loggers, we detected a significant influence of reproductive status on thermoregulation. Reproductive females maintained higher and less variable body temperatures than nonreproductive females and this difference was most pronounced during the last 3 weeks of gravidity. Overall, these results highlight the continuum between oviparity and viviparity in squamate reptiles and emphasize the importance of thermal control of early embryonic development independent of reproductive mode.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 499–508.     

Maternal influences on offspring phenotypes and sex ratios in a multi‐clutching lizard with environmental sex determination

Maternal and environmental factors are important sources of phenotypic variation because both factors influence offspring traits in ways that impact offspring and maternal fitness. The present study explored the effects of maternal factors (maternal body size, egg size, yolk‐steroid allocation, and oviposition‐site choice) and seasonally‐variable environmental factors on offspring phenotypes and sex ratios in a multi‐clutching lizard with environmental sex determination (Amphibolurus muricatus). Maternal identity had strong effects on offspring morphology, but the nature of maternal effects differed among successive clutches produced by females throughout the reproductive season (i.e. maternal identity by environment interactions). The among‐female and among‐clutch variation in offspring traits (including sex ratios) was not mediated through maternal body size, egg size, or variation in yolk steroid hormones. This lack of nongenetic maternal effects suggests that phenotypic variation may be generated by gene by environment interactions. These results demonstrate a significant genetic component to variation in offspring phenotypes, including sex ratios, even in species with environmental sex determination. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 256–266.     

Offspring performance is linked to parental identity and male breeding ornamentation in whitefish

The 'good genes' hypothesis predicts that males advertise their quality with different sexual ornaments and that females are able to recognize the genetic quality of males by evaluating these characteristics. In the present study, we investigated the parental effects on offspring performance (feeding and swimming ability of newly-hatched larvae) and examined whether male ornamentation indicates offspring success in performance trials of whitefish ( Coregonus lavaretus Linnaeus). Offspring first-feeding success had a strong paternal effect and it was also positively correlated with the size of male breeding tubercles, indicating that breeding ornamentation of males can function as an honest indicator of their genetic quality. In addition, the observed positive correlation between male tubercle size and condition factor suggests that highly ornamented males are efficient foragers and that this trait may have a heritable basis. By contrast to feeding success, only a maternal effect was found in the swimming ability of the larvae. Clear family-specific differences observed in both measures of performance strongly suggest that parental identity may have important effects on larval survival in the wild.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 532–539.     

A new spineless species of Vella (Brassicaceae) from the high mountains of south-eastern Spain

Vella castrilensis sp. nov. is described from the high mountains of Granada and Jaén provinces (south-eastern Iberian Peninsula), where it grows on calcareous soils. It is a hexaploid (2 n  = 68) spineless dwarf shrub, woody at the base, with oblong-lanceolate, entire to shallowly dentate leaves and fruit with an acute tongue-shaped stylar segment and strongly reticulate-nerved valves. The characteristic unique combination of vegetative, karyological and reproductive features of V. castrilensis is not present in any described taxon of the genus, and warrants recognition at the species rank. Affinities and differences with other related taxa are discussed. Phylogenetic, biogeographical, bioclimatic, ecological data and conservation proposals are also reported.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 121–128.     

Sex-dependent effects of sibling cannibalism on life history traits of the ladybird beetle Harmonia axyridis (Coleoptera: Coccinellidae)

To investigate the sex-dependent effects of sibling cannibalism on variations in life history traits, I analysed body size, weight and instar interval in relation to the occurrence of sibling cannibalism in the ladybird beetle Harmonia axyridis. Sibling cannibalism at the time of hatching significantly affected the body size and weight of adults. There was a 2.32% and 1.05% increase in the body size of males and females, respectively, and a 3.55% increase and a 2.30% decrease in their respective body weights. Sibling cannibalism also significantly shortened the total and larval instar intervals, by 4.24% in males and by 1.22% in females, mainly due to shortening of the first instar. These results suggest that the effects of sibling cannibalism on life history traits are sex-differentiated and are greater in males than in females. A simulation of aphid density indicated that shortening the instar interval affected larval survival; the aphid density when the larvae completed development was 39.71% and 10.52% larger for cannibalistic males and females, respectively, than for non-cannibals. These results suggest that sibling cannibalism promotes more rapid development and larger adult size, although the effect was more pronounced in males than in females. Faster development may be adaptive for resource tracking, and the large adult size may increase fecundity in females and mating success in males through female mate choice, both resulting in an increase in the fitness of cannibals.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 349–360.     

Size-dependent mating strategies and the risk of cannibalism

The role of sexual selection in determining the nature and direction of sexual size dimorphism may depend upon the timing of sexual selection, and this may also influence the variation in male size. For example, selection through sperm competition favours smaller males in the highly sexually size dimorphic orb-weaving spider Nephila edulis , whereas larger males are better able to exclude their smaller rivals from the central hub of the web where mating takes place. We investigate experimentally the role of body size and hub tenure in determining male fertilization success when males of different sizes compete for a single female over a 24-h period that includes a period of darkness. Our results confirm that small and large males obtain similar paternity share but that, in contrast with previous studies, hub tenure does not translate into greater paternity share. Unexpectedly, smaller males are at greater risk of postmating sexual cannibalism than larger males, suggesting that natural selection through sexual cannibalism may place a lower limit on male size.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 355–363.     

The effect of prey availability on offspring survival depends on maternal food resources

Maternal nutrition can strongly influence embryo development and offspring fitness. The environmental matching hypothesis posits that developmental conditions affect offspring in ways that enable them to appropriately deal with similar post‐developmental conditions, although mismatches between developmental and post‐developmental environments will reduce fitness. To test this hypothesis, reproductive lizards (Anolis sagrei) were reared in environments with high versus low prey availability. The resultant offspring were then reared reciprocally under the same two prey conditions that their mothers experienced. High levels of prey available to mothers increased egg production, although the survival of eggs was low compared to those produced by mothers in the low‐prey treatment. Low prey availability to offspring reduced growth, regardless of the amount of prey available to their mothers. Low prey availability also compromised offspring survival, although this negative effect was only present when mothers experienced high‐prey conditions, whereas matching of low‐prey conditions in maternal and offspring stages resulted in high survival. However, because the mismatch of low maternal and high offspring prey availability resulted in similar offspring survival to the matched treatments, our results do not fully support the environmental matching hypothesis. Nevertheless, the present study highlights the interactive role of maternal and post‐hatching environments in generating variation in offspring fitness. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 437–447.     

Facultative apomixis and hybridization in arctic Potentilla section Niveae (Rosaceae) from Svalbard

The degree of sexual seed set and the ability to cross were investigated in three taxa of Potentilla section Niveae ( P. chamissonis , P. insularis and P. nivea ) from the Svalbard Archipelago. Emasculated and bagged flowers had little or no seed set, while 71% of the emasculated, bagged and pollinated flowers had some. The taxa are interpreted as pseudogamous apomicts. Parental plants and offspring were subjected to isozyme analysis. Most of the offspring were clones of the mother plant, but 1.7% had bands from the pollen donor. The sexual offspring were all produced by mother plants of P. insularis , which is interpreted as a facultative apomict. Surprisingly, crosses between different taxa gave higher seed to ovule ratios and numbers of sexually produced offspring than crosses within taxa. Some of the sexually produced hybrid offspring had the same kind of hairs on the petioles as their mother taxon, showing that hybrids may not be intermediate in hair characters, which are considered important in the delimitation of these taxa. The results of the present study indicate very close relationships among the investigated plants.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 373−381.     

Functional Value of Matriphagy in the Spider Amaurobius ferox

The adaptive value of matriphagy, the consumption of the mother by her offspring, in the sub-social spider Amaurobius ferox (Araneae, Amaurobiidae) was experimentally evaluated in terms of the benefits to the offspring and the costs to the mother. Matriphagy resulted in a 2.5-fold weight gain in the offspring over their initial weight, advancement of their moulting time, and larger body mass at dispersal in comparison with clutches deprived of matriphagy, but otherwise well provisioned with prey. Matriphagous offspring were also more successful at capturing large prey items, had a more extended social period, and a higher survival rate at dispersal. Mothers separated from their broods just prior to matriphagy were able to produce second clutches, effectively producing 33% more viable offspring (compared with the first clutches). However, the estimated reproductive outputs of the alternative maternal strategies (being devoted to the first clutch vs. deserting the first and producing a second one) suggest that mothers of A. ferox that are cannibalized by their broods enjoy greater reproductive success than those that escape cannibalism and produce second clutches.     

A test of sexual cannibalism models, using a sit-and-wait predator

Sexual cannibalism, the capture and consumption of one sex by the other resulting from sexual interactions, provides an intriguing set of conditional fitness payoffs. Usually these payoffs will only benefit the cannibal, but under certain circumstances advantages might accrue to the cannibalized individual as well. Here I test three models of sexual cannibalism with data from a sit-and-wait predator, the crab spider Misumena vatia , whose lifestyle differs from the species for which these models were generated. The models include both precopulatory mechanisms (no gametes exchanged) and postcopulatory ones (gametes exchanged) perceived to generate adult fitness payoffs and situations in which cannibalism is not adaptive in the adult stage (aggressive spillover). In M. vatia , females sometimes cannibalize males before mating can occur. Precopulatory cannibalism is unlikely to reward female M. vatia significantly because of the diminutive size of the males, but it could be part of a broader syndrome that provides the females with large food payoffs earlier in the life cycle. Although the frequency of cannibalism is low (3.8–7.6%), this level suffices to be an important selective factor, a point accentuated by the extremely cautious behaviour of the males toward the females. I then compare the traits associated with these models to the species for which they were designed and certain others for which adequate data also exist. None of the sets of responses to these traits closely resembles each other in the different species. These results suggest that sexual cannibalism has developed in response to a heterogeneous collection of lifestyles and phylogenetic constraints.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 427–437.     

Hybrids between Chorthippus brunneus and C. jacobsi (Orthoptera: Acrididae) do not show endogenous postzygotic isolation

Studies of prezygotic and postzygotic isolation in Drosophila have shown in general that species in sympatry tend to evolve prezygotic barriers earlier than do species in allopatry. However, postzygotic barriers tend to evolve at the same evolutionary rate in both sympatric and allopatric species. In contrast to these observations, the grasshoppers Chorthippus parallelus parallelus and C. p. erythropus show complete hybrid male sterility but only limited prezygotic isolation after an estimated 0.5 millions years of divergence. Like their congeners, C. brunneus and C. jacobsi form a hybrid zone where their ranges meet in northern Spain. However, the hybrid zone is mosaic and bimodal and, in contrast to the high levels of postzygotic isolation between C. parallelus subspecies, these two species showed no significant reduction in hybrid fitness in F₁ or backcross generations relative to the parental generations. The level of prezygotic isolation in laboratory tests was comparable to that between C. parallelus subspecies. These results suggest that endogenous postzygotic isolation does not play an important role in the reproductive isolation between C. brunneus and C. jacobsi , or in determining the structure of the hybrid zone. Exogenous postzygotic isolation may be present and should be tested in future studies.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 195–203.     

Unexpected insularity effects in invasive plant mating systems: the case of Carpobrotus (Aizoaceae) taxa in the Mediterranean Basin

Although biological invasions are currently recognized as being especially severe on islands, little attention has been given to detecting parameters influencing this pattern. This study tested the common perceptions that uniparental reproductive modes are associated with invasive and/or island plant populations due to a lack of pollinator vectors and/or small initial population sizes, and that pollinator services and biparental seed production modes will subsequently be associated with adjacent mainland sites. Using controlled pollination and germination experiments on invasive Carpobrotus edulis and C.  aff. acinaciformis populations in both island and mainland habitats in south-east France, we found no evidence to support these hypotheses. All significant mean differences found between locations for reproductive indices describing uniparental reproductive modes were significantly smaller in island populations. In contrast, seedling sizes issuing from manual outcrossing and manual hybridization experiments, regardless of taxon, as well as C.  aff.  acinaciformis hybridization indices, were larger in island populations. The presence of significant, though limited, pollinator service was detected for all populations, and stressed the importance of such mutualisms. We suggest that invasive plant reproductive traits could be highly contextual, and that islands may have a potential role in the acceleration of the invasion process through the production of highly variable populations.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 65–79.     

Facultative polygyny in the plant-ant Petalomyrmex phylax (Hymenoptera: Formicinae): sociogenetic and ecological determinants of queen number

In polygynous ants it has been proposed that the coexistence of several queens in a colony evolved as a response to ecological, social and genetic parameters. We present demographic, histological and genetic data showing that the plant-ant Petalomyrmex phylax is facultatively and secondarily polygynous. Polygyny is functional, lowers the reproductive output per queen, and is a kin-selected trait as new queens accepted in polygynous colonies are highly related females that never left their natal colony. The degree of polygyny varies according to a geographical gradient. Northern colonies can be strongly polygynous, while at the southern edge of the species' distribution, colonies are almost exclusively monogynous. However, ecological studies of the host-plant populations revealed that this cline could not be explained by variations in the degree of nest site limitation. We discuss selective costs and benefits associated with these social structures, and propose that this cline may result from historical processes such as selection of a more dispersive strategy along a colonization front.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 133–151.     

A new species of Ferula (Apiaceae) from South Anatolia, Turkey

Ferula coskunii H. Duman & M. Sağıroğlu sp. nov. from the Amanos Mountains (C6: Hatay-Turkey) is described and illustrated. It is closely related to F. drudeana Korovin, from which it differs mainly in habit, basal leaves, stem and mericarp features.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 357–361.     

Age-related changes in thoracic mass: possible reallocation of resources to reproduction in butterflies

In nectar-feeding butterflies, reproductive potential is usually thought to depend on the size of the reproductive reserves in the abdomen, the adult food quality and, for females, the amount of resources received in the spermatophores at mating. Recent findings show that thorax mass and nitrogen content decrease with age in some butterfly species, and that thorax resources may be used for reproduction in the butterfly Pieris napi , just as in some other insects. In order to determine whether this is a general pattern and ascertain how it relates to the investment of resources in reproduction we studied the dynamics of thorax and abdomen mass changes in 11 Swedish butterfly species. By regressing thorax and abdomen mass on age of field-collected specimens, we show that loss of mass from both the thorax and the abdomen is a common phenomenon among nectar-feeding temperate zone butterflies under natural conditions. We argue that our results indicate that resources from flight muscles can be reallocated to reproduction by these butterflies, thus increasing their reproductive potential. Within species, females use proportionately more resources from the thorax than do males, as expected from the difference in investment of resources in reproduction. Among males we expect species with a higher reproductive investment to have a larger decrease in thorax and abdomen mass, and our data indicate that this is the case. Looking at the change in relative thorax mass, our results suggest that the use of resources from the thorax does not affect flight performance negatively, something that could constrain the use of muscle resources.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 363–380.     

Morphological,genetic and behavioural analyses of a hybrid zone between the ground beetles Carabus lewisianus and C. albrechti (Coleoptera,Carabidae): asymmetrical introgression caused by movement of the zone?

We analysed the putative hybrid zone between the ground beetles Carabus lewisianus Breuning and C. albrechti Morawitz from the Kanto and Tanzawa Mountains in central Honshu, Japan, using morphological, genetic, and behavioural data. Canonical discriminant analysis of three external and five genital morphological characters revealed an apparent morphological gap, suggesting restricted sympatry in the contact zone. RFLP and molecular phylogenetic analyses of the mitochondrial gene ND5 revealed that haplotypes originating from C. lewisianus introgressed extensively into C. albrechti . Fitting tanh curves to morphological (external and genital) and genetic characters showed differential cline widths and centres, suggesting stronger selection for genital morphology with narrower cline widths, and extensive introgression of the nuclear genes responsible for external characters relative to those controlling genital morphology into the range of C. albrechti . Linkage disequilibria between the morphological characters did not differ from zero, suggesting sufficient recombination occurred in the zone but the effect of small sample sizes was not negligible. Mate choice trials demonstrated that males showed a weak but not significant preference for conspecific mates. We interpret these patterns as evidence for (1) partial reproductive isolation presumably by postmating processes; (2) selection against intermediate genital morphologies, and (3) independent responses of the characters to different evolutionary forces that may cause movement of genital clines and introgression of the mitochondrial gene and external morphology. The movement of the boundaries may have resulted in the particular species distribution patterns in the study area.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 79–94.     

A new species of Oxalis section Corniculatae (Oxalidaceae) from the Balearic islands

A new species, Oxalis ferae Llorens, Gil & Cardona (Oxalidaceae) from the Balearic islands (Spain) is described and illustrated. It is considered to be endemic to Mallorca. The morphological differences between the more closely related taxa of Oxalis section Corniculatae are discussed. IUCN threatened category and observations on the ecology of the populations are noted.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 489–493.     

Temporal patterns of geographic parthenogenesis in a freshwater snail

Geographic parthenogenesis describes the observation that parthenogenetic organisms tend to occupy environments different from those of their close, sexually reproducing relatives. These environments are often described as extreme or disturbed habitats. We examined whether patterns of geographical parthenogenesis persist over time, by conducting a 3-year life-history survey and comparing two very proximate habitats of the freshwater snail Melanoides tuberculata : Nahal Arugot, a desert stream naturally disturbed by flash floods, and Or Ilan, a stable freshwater pond. Both sites occur in a xeric environment and are subject to otherwise similar biotic (e.g. parasites, predators) and climatic conditions. In the stable habitat, male frequencies and snail densities were significantly higher than in the disturbed one, whereas infection levels, mean embryo counts, and water temperatures were similar at both sites. Additionally, male frequencies declined after density decreased, thereby providing evidence for geographical parthenogenesis via reproductive assurance. Infection prevalence was very low regardless of reproduction mode. Although further genetic work is required, the apparent metapopulation structure of M. tuberculata in the Judean desert may be suitable for evaluating other possible explanations of geographical parthenogenesis.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 711–718.