Floral phenology and sex expression in functionally monoecious Rhoiptelea chiliantha (Rhoipteleaceae)

The reproductive ecology of wind-pollinated gynomonoecious species, in which the individual plant produces both female (pistillate) and perfect flowers, has rarely been studied. We examined the floral phenology and reproductive traits in Rhoiptelea chiliantha , described as gynomonoecy, to understand the adaptive significance of this sexual system. This species is a rare tree native to south-western China and northern Vietnam. The flowers are characterized by an anemophilous pollination syndrome, but no insects were observed foraging on them. Perfect flowers have larger tepals but smaller stigmas than female flowers, indicating flower size dimorphism. Floral ratios of female to perfect flowers are stable in different individuals and populations. On individual plants, perfect flowers open first, followed by female flowers, with a 1-week interval. Perfect flowers are protogynous with a 3.7-day interval (neuter phase) between the female phase (1.5 days) and expanded male phase (8.2 days). Both female and perfect flowers exhibit pronounced synchrony in flowering at the levels of inflorescences and individuals. However, flowers on different individuals show asynchronicity in timing of initial blooming. Tracking the process from pollination to fruit maturation, we found that female flowers contributed almost exclusively to seed production, but perfect flowers were sterile (functionally males). Therefore, this plant is functionally monoecious. This finding resolved a puzzle on the occurrence of female flowers in this plant, because previous reports described female flowers as being sterile. As the sex phases were completely separate between individuals, the pattern of floral phenology may ensure that outcrossing strongly predominates.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 145–151.     

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.     

Plasticity, its cost, and phenotypic selection under water and nutrient stress in two annual grasses

A comparative approach can prove to be a useful tool for studying phenotypic plasticity, if applied to specific traits involved in adaptation to particular environment in more than one species across co-located populations. The present study tested whether two annual grasses, Hordeum spontaneum and Avena sterilis , belonging to the same guild, having similar stature, seed dispersal mechanism, breeding system, and genetic variation, and sampled in exactly the same environmentally specific locations, differed with respect to: (1) plasticity in traits involved in adaptation, namely the onset of reproduction and maternal investment involving the number of inflorescences, spikelets per inflorescence, the weight of individual spikelets, and abortion rate; (2) the cost of this plasticity, and (3) the pattern of phenotypic selection on the above traits. The two species exhibited highly differing amounts of phenotypic plasticity in the onset of flowering and several reproductive traits (number of inflorescences, spikelets per inflorescence, abortion rate), but no plasticity costs in any experimental environment. The two species demonstrated a decreasing similarity in the regulation of reproduction in four experimental environments: benign, water, nutrients and water × nutrient deficient. Correlational selection appears to contribute, although not solely, to the observed species differences with respect to the regulation of reproduction.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 581–593.     

Energetics of reproduction: consequences of divergent selection on egg size, food limitation, and female age for egg composition and reproductive effort in a butterfly

Using lines artificially selected on egg size and being subjected to a restricted and an unrestricted feeding treatment, we examined the relationships between egg size, egg number, egg composition, and reproductive investment in the butterfly Bicyclus anynana . Despite a successful manipulation of egg size, correlated responses to selection in larval time, pupal mass, pupal time, longevity, fecundity, or the amount of energy allocated to reproduction were virtually absent. Thus, there was no indication for an evolutionary link between offspring size and reproductive investment. Egg composition, in contrast, was affected by selection, with larger eggs containing relatively more lipid and water, but less protein and energy compared to smaller eggs. Hence, females producing large eggs did not have to sacrifice fecundity due to adjustments in egg composition. Food limitation per se caused only minor changes in egg composition, and there was no general reduction in egg provisioning with female age. The latter was restricted to food-limited females, whereas egg quality remained remarkably similar throughout the females' life in control groups. We conclude that neglecting changes in biochemical egg composition, depending on genetic background, food availability, and female age, may introduce substantial error when estimating reproductive effort, and may ultimately lead to invalid conclusions.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 403–418.     

Reproductive biology in Anagyris foetida L. (Leguminosae), an autumn–winter flowering and ornithophilous Mediterranean shrub

As most plants of the Mediterranean region bloom in spring, there have been few studies of the reproductive biology of species with autumn–winter flowering. In this study, we investigate the breeding system of Anagyris foetida , one of the few shrubs that blooms at this time. The floral, phenological, and reproductive aspects of two populations of this Mediterranean legume from south-west Spain were studied via field and laboratory experiments. The variability of fruit and seeds was studied in another 12 Iberian populations with respect to certain meteorological parameters (temperature and rainfall). Anagyris foetida shows cauliflory, marked floral longevity, and adichogamy. The peak of flowering is in January–February. It is self-compatible, with no clear advantage of cross- over self-pollination, and with virtually no autonomous self-pollination. This is because the stigma, like some other legumes, prevents the germination of pollen if its surface is not ruptured by pollinators. The number of seeds per fruit under natural pollination was positively correlated with the total rainfall during the fruiting period (from January to May), and significantly influenced the percentage of fruit weight represented by the pericarp, in the sense that the smaller the number of viable seeds in the fruit, the greater the percentage of pericarp weight.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 519–532.     

Seed ecology of a Mediterranean perennial herb with an exceptionally extended flowering and fruiting season

Lobularia maritima is a Mediterranean short-lived herb with a flowering and fruiting season that lasts for ten months. Previous studies have shown that recruitment in periods other than autumn of the flowering season has few demographic implications; that is contributes little to the population growth rate. Since environmental conditions in periods other than autumn would allow recruitment, we examined to what extent the seed ecology of L. maritima accounts for recruitment shortage for the greater part of the year. To this end, we studied the effects of selfing and outcrossing on seed production and germination, within- and between-year variation in seed mass and germination, seed characteristics in the soil seed bank throughout the year, and the effect of temperature as a factor controlling seed germination. Results indicate that selfing does not decrease recruitment, and thus the observed changes in visitation rate and pollinator composition throughout the year cannot account for differences in recruitment. Germinability decreases throughout the year, suggesting a possible cost in reproduction associated with extended flowering. L. maritima has a transient seed bank whose seeds also experience a decrease in their germination throughout the year. Finally, temperature affects seed germination patterns, indicating the existence of quiescence mechanisms that prevent germination in the months prior to the summer drought. Overall, the results obtained support and, at least partly, explain the recruitment patterns of L. maritima observed in the field.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 273–280.     

Dissecting components of flowering pattern: size effects on female fitness

Flowering synchrony is essential for plant reproductive success, especially in the case of small‐sized populations of self‐incompatible species. Closely related to synchrony, flowering intensity influences pollinator attraction and pollinator movements. Thus, a high flowering intensity may increase pollinator attraction but, at the same time, may also increase the probability of geitonogamous pollinations. Depending on the mating system, the female fitness of plants in small populations may be affected by both the positive effects of higher flowering synchrony and pollinator attraction and the negative effects of geitonogamous pollinations induced by a high flowering intensity. It was hypothesized that different‐sized plants in a population would show contrasting flowering patterns, resulting in differences in pollinator behaviour. These influences could result in differences in mating and female reproductive success. This hypothesis was tested by studying the flowering pattern of Erodium paularense (Geraniaceae), a rare and endangered endemic of central Spain. The temporal distribution of flower production was explored throughout the reproductive season, and the probability of xenogamy and geitonogamy and their relationship to plant size and fitness components were calculated. The analysis of this partially self‐compatible species showed diverse flowering patterns related to different plant sizes. Small plants produced a larger number of seeds per fruit in spite of having lower values of flowering synchrony. By contrast, large plants produced a larger number of seeds from geitonogamous pollinations. The effect of different flower displays and outcrossing rates on seed set varied throughout the season in the different groups. Our findings highlight the relevance of individual plant size‐dependent phenology on female reproductive success and, in particular, on the relationship between flowering synchrony and fitness. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156 , 227–236.     

The effect of flower position on male and female reproductive success in a deceptively pollinated tropical orchid

In many plants, including orchids, differential fruit set along the inflorescence has been attributed to pollinator behaviour. For instance, the pollinator, moving up the inflorescence, becomes satiated with the resources and leaves before visiting the upper flowers. Consequently, the pollinators do not visit flowers as frequently higher up the inflorescence. Alternatively, flower size may vary along the inflorescence, making pollination ineffective as flowers decrease in size. I tested for the presence of differential pollination along the inflorescence in a pollinator-limited tropical epiphyte, Lepanthes rupestris Stimson, and determined the likely cause of the observed pattern. As this species has inflorescences with sequential flowering, pollinator behaviour, moving up the inflorescence as in synchronous multiflowering inflorescences, can be discounted as an explanation for differential fruit set. Fruit set is shown to be more frequent at the base of the inflorescence, but male reproductive success through pollinarium removal is basically independent of flower position. Moreover, cross-pollination by hand at variable flower positions along the inflorescence results in equal fruit set, suggesting that resources are not limiting and cannot explain the cause of differential fruit production along the inflorescence in natural populations. Furthermore, flower size is shown to diminish along the inflorescence, suggesting that the pollinator(s) may be ineffective at depositing the pollinarium in the smaller higher flowers. Consequently, pollinator behaviour and its interaction with flower size, and not resource limitation, is likely to be the main cause of differential fruit set along the inflorescence in L. rupestris .  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 405–410.     

Measuring temporal variation in reproductive output reveals optimal resource allocation to reproduction in the northern grass lizard, Takydromus septentrionalis

We measured the reproductive output of Takydromus septentrionalis collected over 5 years between 1997 and 2005 to test the hypothesis that reproductive females should allocate an optimal fraction of accessible resources in a particular clutch and to individual eggs. Females laid 1–7 clutches per breeding season, with large females producing more, as well as larger clutches, than did small females. Clutch size, clutch mass, annual fecundity, and annual reproductive output were all positively related to female size (snout–vent length). Females switched from producing more, but smaller eggs in the first clutch to fewer, but larger eggs in the subsequent clutches. The mass-specific clutch mass was greater in the first clutch than in the subsequent clutches, but it did not differ among the subsequent clutches. Post-oviposition body mass, clutch size, and egg size showed differing degrees of annual variation, but clutch mass of either the first or the second clutch remained unchanged across the sampling years. The regression line describing the size–number trade-off was higher in the subsequent clutch than in the first clutch, but neither the line for first clutch, nor the line for the second clutch varied among years. Reproduction retarded growth more markedly in small females than in large ones. Our data show that: (1) trade-offs between size and number of eggs and between reproduction and growth (and thus, future reproduction) are evident in T. septentrionalis ; (2) females allocate an optimal fraction of accessible resources in current reproduction and to individual eggs; and (3) seasonal shifts in reproductive output and egg size are determined ultimately by natural selection.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 315–324.     

Seed dormancy and germination behaviour in two Euphrasia species (Orobanchaceae) occurring in the Swiss Alps

The optimal time for germination of a seed depends on environmental conditions of its habitat, the life cycle of the germinating plant, and the conditions for successful establishment, growing and reproduction. We studied the germination behaviour of the alpine annual Euphrasia minima and an alpine ecotype of E. salisburgensis in a lowland garden experiment. Seeds of both species and their hybrids germinated at constant (5 °C) and at varying temperatures (3–10 °C), and never before spring after seed ripening. Germination was spread over 3 years, which suggests that the seeds formed a persistent seed bank. The two species together with E. minima and E. minima hybrids differed significantly in the germination rate in the first and second spring.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 649–656.     

Clonality and recombination in the arctic plant Saxifraga cernua

The circumarctic clonal plant Saxifraga cernua reproduces efficiently via bulbils, largely depends on insects for pollination and appears to set seed very rarely. However, high levels of genotypic variation observed at small spatial scales in the arctic archipelago of Svalbard have been taken as evidence of occasional sexual reproduction. Here we assess the relative contributions of mutation and recombination to random amplified polymorphic DNA variation in four populations in East Greenland and re-analyse the Svalbard data. Greater variation due to recombination was predicted in Greenland than in Svalbard, because the higher summer temperatures and longer growing season likely increase the chances for sexual reproduction. Although we observed higher levels of genotypic diversity in Greenland than in Svalbard, matrix incompatibility and linkage disequilibrium measures provided no evidence of more sexuality, suggesting differences in glacial/postglacial history. The genetic structure and spatial distribution of clones suggest that clonal migration may increase variability in local populations, which is consistent with frequent large-scale migration in this species inferred from a circumarctic analysis of chloroplast DNA haplotypes. We conclude that a combination of somatic mutations and sexual reproduction has contributed to the observed patterns of genotypic diversity in the Greenland and Svalbard populations of S. cernua , and that sexual reproduction is important in maintaining genotypic diversity, despite the rarity of observations of seed setting.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 209–217.     

Character divergences and convergences in canopy-dwelling Loranthaceae

The Loranthaceae is the largest plant family with aerial branch parasites termed mistletoes. Three genera of Loranthaceae are terrestrial root parasites and the remaining 72 genera are aerial parasites. Several characters, including habit, haustorial type, germination pattern, pollen morphology, chromosome number, inflorescence morphology and flower merosity, fusion, symmetry and size, are considered to reflect evolutionary relationships within the family. Convergence is a common evolutionary pattern and can confound interpretations of evolution. We investigated character evolution by mapping character states onto a phylogenetic tree based on the nuclear ITS and chloroplast trnL–trnF regions. Convergences in form were found in several characters, including habit, haustorial type, flower symmetry and merosity. These convergences typically correspond to ecological parameters such as pollination syndrome or stresses associated with the canopy habit. Other characters such as chromosome number and germination pattern illustrate divergent evolution among clades.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 101–113.     

Photoperiod as a reproductive cue in the marsupial genus Antechinus: ecological and evolutionary consequences

Species in the Australian marsupial genus Antechinus exhibit a short annual mating period which is concluded by the abrupt death of all males. The timing of the annual rut within each of the ten described species varies little from year to year at any given locality, but for some species can differ by up to four months between locations. To determine the influence of photoperiod in regulating the precise interannual synchrony of mating and ovulation, we first investigated populations of each species at over 300 localities throughout their geographical ranges to identify the time of reproduction. We then compared the absolute photoperiod and the rate of change of photoperiod prevailing at the time of reproduction in all population localities. A different, and characteristic, rate of change of photoperiod was correlated strongly with the reproductive timing of four species; there was probably a correlation with reproduction in four more species, but sample sizes were small. For two species, there was no obvious photoperiodic correlation with time of reproduction. There was no evidence that absolute photoperiod or ambient temperature explained the synchrony or narrow timespan of reproduction among any species of Antechinus . Different species-specific ovulatory responses to photoperiod appear to separate the timing of reproduction in sympatric species, with the larger member of species pairs usually breeding first. We suggest that photoperiodic cues (1) allow females to produce young during seasons when food is most reliable and abundant and their energetic demands are maximal; (2) facilitate allochronic isolation between sympatric congeners, and (3) maximize body size differences and hence ecological separation between species.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 365–379.     

Reproductive performance of otters Lutra lutra (Linnaeus, 1758) in Eastern Germany: low reproduction in a long-term strategy

The reproductive status of female otters ( Lutra lutra ) was determined from an examination of 518 carcasses collected from Eastern Germany between 1950 and 2001. In Germany otters mate throughout the year. Significant seasonal differences are evident with a distinct peak in summer. Females were in breeding condition between the ages of three and 15 years, with the majority between six and nine years. The mean litter size at birth was 2.36, which was the lowest when compared with other inland populations. There were no significant differences in litter size for regions, seasons or age. Prenatal losses account for 26.31%. The mortality between birth and first appearance of cubs following their mother is about 29%. An effective reproduction rate of 78.4% was deduced from the known age structure of the sample and the proportion of breeding females. Reproductively active females show a significantly higher body condition than non-reproductive females. Following these investigations into breeding status, suggestions on reproductive output and the life history of the otter are discussed.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 329–340.     

Effective population size estimation on Sardina pilchardus in the Bay of Biscay using a temporal genetic approach

Over three consecutive years, we surveyed the temporal variability in genetic structure of sardine populations in the Bay of Biscay and effective population size. Based on individual age, the genetic structure of year classes of the fishes was also surveyed, showing that populations of sardines have weak but significant genetic differences between sampling years and between year classes. We used two different methods to assess effective population size. The methods resulted in different values but a similar range, indicating a low effective population for Sardina pilchardus . Effective population size decreased over the 3 years, probably resulting from an abundance of fish in the Bay. Based on these results, we conclude that temporal variability in the genetic structure of the sardine population and effective size are likely related to environmental conditions in the Bay. Finally, we propose to use effective population size to estimate biomass of sardines in the Bay.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 591–602.     

Distribution and genetic divergence of two parapatric sibling ant species in Central Europe

The two sibling ant species Temnothorax nylanderi and Temnothorax crassispinus are widely distributed throughout deciduous forests in Europe. Their resemblance in morphology and similar ecological requirements suggest that they evolved from the same ancestral species in different glacial refugia and re-immigrated into Central Europe after the last ice age. Here, we show that the two species are parapatrically distributed in south-eastern Germany and hybridize along a narrow contact zone close to the continental divide. Phylogeographical data based on the mitochondrial genes cytochrome oxidase subunit I and cytochrome b suggest that the dominant haplotypes for T. nylanderi and T. crassispinus might have diverged already 1.5–2 Mya. Intraspecific variability is extremely low in both species, which might be explained by severe bottlenecks during rapid postglacial expansion into Central Europe.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 223–234.     

Fitness, developmental instability, and the ontogeny of fluctuating asymmetry in Daphnia magna

Fluctuating asymmetry (FA) is the most commonly used measure of developmental instability. The relation between FA and individual fitness remains controversial, partly due to limited knowledge on the mechanisms behind variation in FA. To address this, we investigated the associations between FA, growth and reproduction as well as the ontogeny of FA in a clonal population of Daphnia magna . FA was not correlated with growth and reproduction, either at the between- or the within-individual level, in a high ( N  = 48 individuals) or in a low ( N  = 52 individuals) food-quantity regime. There were therefore no indications of functional effects of FA or of phenotypic trade-offs between developmental stability, growth and reproduction. Individual asymmetries varied randomly in sign and magnitude between subsequent molts ( N  = 19 individuals, 9–11 instars), but the levels of FA were generally lowest at intermediate ages. No feedback between right and left sides was detected. This suggests that FA only reflects the most recent growth history, that developmental instability may increase in old age, and that FA depends on processes operating on each side of the body independently. The results also suggest that FA differences within and among individual Daphnia are largely random, with limited biological significance.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 179–192.     

Reproductive effort and sex allocation strategy in Commelina benghalensis L., a common monsoon weed

Commelina benghalensis L. exhibits variability in both foliar and floral features; every plant bears three types of branches and four types of flowers. The branches are negatively geotropic, positively geotropic and diageotropic. The flowers are uni- or bisexual, chasmogamous and cleistogamous. This variability influences the breeding system as well as resource allocation to male and female functions. The plants allocate c. 15% of their total resources to reproduction, the major part of which (68.9%) is devoted to production of aerial branches. The proportion of reproductive effort (RE) allocated to various branch systems is correlated with the availability of resources at the time of their differentiation. The pollen/ovule (P/O) ratio, female : male biomass ratio and reproductive output vary between different flower and branch types; variation is more pronounced in the latter. These variations notwithstanding, the results are in line with Charnov's sex allocation theory. The cleistogamous flowers of aerial branches are, however, an exception, being male- rather than female-biased. The reason behind the deviation is, in all probability, their recent evolution from chasmogamous flowers.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 403−413.     

Genital variation in a dimorphic moth Selenia tetralunaria (Lepidoptera, Geometridae)

Insect genitals vary greatly among species and provide a key tool for species-level taxonomy. Insects differing in the genitalia are often treated as discrete, reproductively isolated species. This principle dates back to the lock-and-key hypothesis, which states that genitalia vary between species in order to provide a mechanical reproductive isolation system. Thus, the hypothesis assumes low within-species variability in genital traits. However, recent studies suggest that sexual selection may be responsible for the evolution of insect genitalia. We studied allometry and genital size and shape variation in a dimorphic moth Selenia tetralunaria . We found that the genitalia showed negative allometry in relation to body size as reported in many insect and spider species. This allometry was stronger in internal genital structures than it was in external genitalia. We also found that there was more variation in internal compared with external genitalia. Finally, we found that the shape of genital structures differed between morphs in all three examined areas. S. tetralunaria is among the first reported cases of genitally dimorphic insect species. Considerable variation in internal genitalia and especially the presence of genital shape differences between morphs were not consistent with the predictions of the lock-and-key hypothesis.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 297–307.     

Locomotor capacity and dominance in male lizards Lacerta monticola: a trade‐off between survival and reproductive success?

Trade-offs between reproduction and survival are important determinants of life-history characteristics of lizards. Organisms cannot increase the allocation of limited resources to reproduction without diverting a proportional amount of energy from another trait. Locomotor performance is an ecologically relevant trait that potentially influences survival by affecting the ability to escape from predators. Most studies have used female lizards as subjects because pregnancy is known to reduce their locomotor abilities, whereas little is known on costs of reproduction in males. In this study we suggest that in males of the lizard Lacerta monticola reproductive investment in morphological traits that confer dominance (i.e. head size) might lead to a low probability of survival by decreasing investment in other traits that affect locomotor performance (i.e. limb symmetry). We staged laboratory agonistic encounters between males and measured their morphology and burst speed on a race track to examine possible relationships between morphology, social dominance and locomotor capacity. Our results indicate that social dominance was positively related to relative head height, and that escape speed was negatively related to levels of fluctuating asymmetry in femur length, but also negatively related to relative head height. Males with greater relative head height also had more asymmetrical femurs, thus dominant males suffered a decrease in locomotor performance. Males with higher heads tend to dominate male–male interactions and hence may gain access to reproductive females, thus increasing their current reproduction success. However, this might occur at the expense of future survivorship mediated by a decrease in escape speed. Therefore, in male L. monticola there might be a trade-off between current reproductive success and survival.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002; 77 , 201–209.