Host acceptance and sex allocation of Nasonia wasps in response to conspecifics and heterospecifics

Species recognition is an important aspect of an organism''s biology. Here, we consider how parasitoid wasps vary their reproductive decisions when their offspring face intra- and interspecific competition for resources and mates. We use host acceptance and sex ratio behaviour to test whether female Nasonia vitripennis and Nasonia longicornis discriminate between conspecifics and heterospecifics when ovipositing. We tested pairs of conspecific or heterospecific females ovipositing either simultaneously or sequentially on a single host, using strains varying in their recent history of sympatry. Both N. vitripennis and N. longicornis rejected parasitized hosts more often than unparasitized hosts, although females were more likely to superparasitize their own species in the sequential treatment. However, sex ratio behaviour did not vary, suggesting similar responses towards conspecifics and heterospecifics. This contrasts with theory predicting that heterospecifics should not influence sex ratios as their offspring do not influence local mate competition, where conspecifics would. These non-adaptive sex ratios reinforce the lack of adaptive kin discrimination in N. vitripennis and suggest a behavioural constraint. Discrimination between closely related species is therefore context dependent in Nasonia. We suggest that isolating mechanisms associated with the speciation process have influenced behaviour to a greater extent than selection on sex ratios.     

Etude de l'influence des facteurs visuels sur les comportements sexuels et agressifs de Mollienesia (Poecilia) velifera (Pisces,Poeciliidae)

One of the most striking features of cave fishes is their pronounced ocular regression. They are, however, perfectly adapted since they multiply and display the behaviour patterns which are biologically important in their surroundings. It is therefore of interest to study the relative role of visual factors in the corresponding behaviour patterns in related epigean species.Other authors have previously demonstrated the dominant role of chemical stimuli in the releasing of sexual behaviour patterns in M. sphenops, of which some forms are cavernicolous. In M. velifera and M. latipinna, on the contrary, visual factors seem more important. The releasers of the aggressive behaviour patterns of these three species seem to be predominantly visual. In the present study, the importance of visual factors has been verified experimentally in M. velifera by comparing the observed behavioural differences in experimentally blinded individuals and normal controls.The results of the pre-experimental phase show that there is no difference between these two groups before blinding, neither in the frequency of the patterns, nor in their numerical values.The results of the first experimental phase confirm the importance of visual releasers in the sexual behaviour patterns of M. velifera. Thus, sexual approaches, mouth—abdomen contacts and copulation attempts are totally absent in blinded pairs. Sexual motivation however is still present since the gonopodal flexions are equally frequent in both groups. On the other hand, it seems that aggressive behaviour patterns are maintained in blinded individuals but only when the partners are very close to each other or when an actual contact has taken place.The comparison of pairs in which only one of the partners is blinded (second experimental phase), as well as results of the first phase, suggest that experimental blinding does not cause hormonal changes liable to influence the observed patterns. In addition to this, the role of the ♀ in courtship has been specified. When a ♀ individual is intact and receptive, it will initiate courtship when it is placed in the company of an inactive blinded partner.     

Mating behaviour in the endangered Sonoran topminnow: speciation in action

Two species of the endangered Sonoran topminnow, Poeciliopsis occidentalis and P. sonoriensis, occur in two isolated drainage systems in southeast Arizona, U.S.A., and are allopatric throughout their range. Although these two taxa are morphologically very similar, and have been previously described as the same species or subspecies, several molecular studies have since indicated that they differ more than their morphology suggests. To determine whether the behaviours of the two species function as premating barriers to reproduction, we investigated their mating preferences and behavioural patterns in a laboratory setting. Results from no-choice mating observations showed that the mating behaviours of the two species differ. Observations conducted during multiple-choice mating trials provided evidence of assortative mating, suggesting an early stage of premating reproductive isolation.     

Fluid physico-chemical properties influence capture and diet in Nepenthes pitcher plants

Background and Aims Nepenthes pitcher plants have evolved modified leaves with slippery surfaces and enzymatic fluids that trap and digest prey, faeces and/or plant detritus. Although the fluid’s contribution to insect capture is recognized, the physico-chemical properties involved remain underexplored and may vary among species, influencing their diet type. This study investigates the contributions of acidity and viscoelasticity in the fluid’s capture efficiency of two ant and two fly species in four Nepenthes species with different nutrition strategies.Methods Four Nepenthes species were studied, namely N. rafflesiana, N. gracilis, N. hemsleyana and N. ampullaria. Fluid was collected from pitchers of varying ages from plants growing in the field and immediately transferred to glass vials, and individual ants (tribe Campotini, Fomicinae) and flies (Calliphora vomitoria and Drosophila melanogaster) were dropped in and observed for 5 min. Water-filled vials were used as controls. Survival and lifetime data were analysed using models applied to right-censored observations. Additional laboratory experiments were carried out in which C. vomitoria flies were immersed in pH-controlled aqueous solutions and observed for 5 min.Key Results Pitcher fluid differed among Nepenthes species as regards insect retention capacity and time-to-kill, with differences observed between prey types. Only the fluids of the reputedly insectivorous species were very acidic and/or viscoelastic and retained significantly more insects than the water controls. Viscoelastic fluids were fatal to flies and were able to trap the broadest diversity of insects. Younger viscoelastic fluids showed a better retention ability than older fluids, although with less rapid killing ability, suggesting that a chemical action follows a mechanical one. Insect retention increased exponentially with fluid viscoelasticity, and this happened more abruptly and at a lower threshold for flies compared with ants. Flies were more often retained if they fell into the traps on their backs, thus wetting their wings. Insect retention and death rate increased with fluid acidity, with a lower threshold for ants than for flies, and the time-to-kill decreased with increasing acidity. The laboratory experiments showed that fewer flies escaped from acidic solutions compared with water.Conclusions In addition to viscoelasticity, the pitcher’s fluid acidity and wetting ability influence the fate of insects and hence the diet of Nepenthes. The plants might select the prey that they retain by manipulating the secretion of H⁺ ions and polysaccharides in their pitcher fluid. This in turn might participate in possible adaptive radiation of this genus with regard to nutrient sequestration strategy. These plants might even structurally influence insect fall-orientation and capture-probability, inspiring biomimetic designs for pest control.     

A Space Oddity: Geographic and Specific Modulation of Migration in Eudyptes Penguins

Post-breeding migration in land-based marine animals is thought to offset seasonal deterioration in foraging or other important environmental conditions at the breeding site. However the inter-breeding distribution of such animals may reflect not only their optimal habitat, but more subtle influences on an individual’s migration path, including such factors as the intrinsic influence of each locality’s paleoenvironment, thereby influencing animals’ wintering distribution. In this study we investigated the influence of the regional marine environment on the migration patterns of a poorly known, but important seabird group. We studied the inter-breeding migration patterns in three species of Eudyptes penguins (E. chrysolophus, E. filholi and E. moseleyi), the main marine prey consumers amongst the World’s seabirds. Using ultra-miniaturized logging devices (light-based geolocators) and satellite tags, we tracked 87 migrating individuals originating from 4 sites in the southern Indian Ocean (Marion, Crozet, Kerguelen and Amsterdam Islands) and modelled their wintering habitat using the MADIFA niche modelling technique. For each site, sympatric species followed a similar compass bearing during migration with consistent species-specific latitudinal shifts. Within each species, individuals breeding on different islands showed contrasting migration patterns but similar winter habitat preferences driven by sea-surface temperatures. Our results show that inter-breeding migration patterns in sibling penguin species depend primarily on the site of origin and secondly on the species. Such site-specific migration bearings, together with similar wintering habitat used by parapatrics, support the hypothesis that migration behaviour is affected by the intrinsic characteristics of each site. The paleo-oceanographic conditions (primarily, sea-surface temperatures) when the populations first colonized each of these sites may have been an important determinant of subsequent migration patterns. Based on previous chronological schemes of taxonomic radiation and geographical expansion of the genus Eudyptes, we propose a simple scenario to depict the chronological onset of contrasting migration patterns within this penguin group.     

Shared pollinators and pollen transfer dynamics in two hybridizing species, Rhinanthus minor and R. angustifolius

Gene flow between hybridizing plant species depends strongly on pollinator behaviour, which affects pollen transfer among floral types and reproductive isolation. We examined bumblebee behaviour and pollen transfer between two hybridizing Rhinanthus species that are very similar in ecology and floral traits. The two species, Rhinanthus minor and R. angustifolius, shared similar pollinator guilds and assemblages, but pollinator recruitment and flower visitation rates were higher in R. angustifolius sites, probably because of its higher reward levels and better visibility. When presented with Rhinanthus flowers, bumblebees that previously foraged on R. angustifolius were less prone to visit R. minor inflorescences, while R. minor foragers accepted both species in similar proportions. Although Rhinanthus has been cited as a case of mechanical isolation resulting from interactions between bee behaviour and differences in stigma and anther placement, we found no support for efficient mechanical reproductive isolation. Bumblebees that foraged on R. minor flowers carried more pollen, but pollen placement on their bodies was similar to that of bees that visited R. angustifolius, and cross-specific stigmatic pollen deposition was similar in both directions. However, the asymmetry in pollinator handling time between the two species, due to dissimilar pollen rewards, may have lowered relative heterospecific pollen receipt on R. angustifolius, suggesting that net gene flow resulting from pollen transfer dynamics is more likely towards R. minor, although this effect remains weak and will be most likely counterbalanced by context-based labile pollinator preference.     

The visual fields of Common Guillemots Uria aalge and Atlantic Puffins Fratercula arctica: foraging,vigilance and collision vulnerability

Significant differences in avian visual fields are found between closely related species that differ in their foraging technique. We report marked differences in the visual fields of two auk species. In air, Common Guillemots Uria aalge have relatively narrow binocular fields typical of those found in non‐passerine predatory birds. Atlantic Puffins Fratercula arctica have much broader binocular fields similar to those that have hitherto been recorded in passerines and in a penguin. In water, visual fields narrow considerably and binocularity in the direction of the bill is probably abolished in both auk species. Although perceptual challenges associated with foraging are similar in both species during the breeding season, when they are piscivorous, Puffins (but not Guillemots) face more exacting perceptual challenges when foraging at other times, when they take a high proportion of small invertebrate prey. Capturing this prey probably requires more accurate, visually guided bill placement and we argue that this is met by the Puffin's broader binocular field, which is retained upon immersion; its upward orientation may enable prey to be seen in silhouette. These visual field configurations have potentially important consequences that render these birds vulnerable to collision with human artefacts underwater, but not in air. They also have consequences for vigilance behaviour.     

Nitrogen limitation, 15N tracer retention, and growth response in intact and Bromus tectorum-invaded Artemisia tridentata ssp. wyomingensis communities

Annual grass invasion into shrub-dominated ecosystems is associated with changes in nutrient cycling that may alter nitrogen (N) limitation and retention. Carbon (C) applications that reduce plant-available N have been suggested to give native perennial vegetation a competitive advantage over exotic annual grasses, but plant community and N retention responses to C addition remain poorly understood in these ecosystems. The main objectives of this study were to (1) evaluate the degree of N limitation of plant biomass in intact versus B. tectorum-invaded sagebrush communities, (2) determine if plant N limitation patterns are reflected in the strength of tracer ¹⁵N retention over two growing seasons, and (3) assess if the strength of plant N limitation predicts the efficacy of carbon additions intended to reduce soil N availability and plant growth. Labile C additions reduced biomass of exotic annual species; however, growth of native A. tridentata shrubs also declined. Exotic annual and native perennial plant communities had divergent responses to added N, with B. tectorum displaying greater ability to use added N to rapidly increase aboveground biomass, and native perennials increasing their tissue N concentration but showing little growth response. Few differences in N pools between the annual and native communities were detected. In contrast to expectations, however, more ¹⁵N was retained over two growing seasons in the invaded annual grass than in the native shrub community. Our data suggest that N cycling in converted exotic annual grasslands of the northern Intermountain West, USA, may retain N more strongly than previously thought.     

Decomposition and nutrient release in halophytes of a Mediterranean salt marsh

This study dealt with the decomposition and nutrient release from the halophytes Atriplex portulacoides, Arthrocnemum macrostachyum, Limoniastrum monopetalum, and Spartina densiflora, the dominant species in the Castro Marim salt marsh, Portugal. Environmental effects on decomposition were also assessed. The study was carried out for one year using the in situ litterbag technique. S. densiflora showed a lower decomposition rate (k = 0.003 day⁻¹) than the other study species (k = 0.005-0.009). Study species showed similar decomposition patterns, that is, the weight loss mostly occurred during the autumn-winter period (study beginning in November). This indicates that temperature in this period did not hamper the decomposition process. The decomposition rate was positively affected by the initial N concentration (r² = 0.87, P < 0.05) and negatively by the C:N ratio (r² = 0.86, P < 0.05) in decomposing materials. At the end of the study, S. densiflora and L. monopetalum, the species with lower initial N concentrations, retained much higher proportion of initial N (89-109%) than the others (5-14%). Also, S. densiflora with the lowest P concentration retained higher proportion of initial P (48%) than the others (5-20%). Release of K and Mg were also slower from S. densiflora and was associated with their initial low concentration in this species. The lowest Mn release was observed from A. macrostachyum and also in relation to the lowest initial concentration. Our study supports the hypothesis that decomposition patterns of marsh species are mostly associated with differences regarding their morphology and chemical composition. Given the higher resistance of S. densiflora to decomposition, its progressive spreading may result in accumulation of organic detritus overtime in invaded salt marshes.     

The relationship between leaf nitrogen,nitrogen metabolites and herbivory in two species of Nyctaginaceae from the Brazilian Cerrado

Guapira graciliflora and Neea theifera are taxonomically related species of the tribe Pisoneae. Both species are found in the same environment, the Brazilian Cerrado, and therefore, are subjected to similar selective pressures. These species occur in oligotrophic environments, yet contain high concentrations of nitrogen in their leaves. The present study was carried out to investigate the ecological role of nitrogen in herbivory on these species. The differences in the N content, compositions of secondary N-metabolites, mechanical resistance, and water content between their leaves indicate that these species have different adaptations as defense mechanisms. In both species, their high nitrogen content seems to promote herbivory. The presence of secondary nitrogen metabolites does not prevent the species from suffering intense damage by herbivores on their early leaves. The herbivory rates observed were lower for mature leaves of both species than for young leaves. In G. graciliflora, nutritional content and leaf hardness are the most important variables correlated with reduction of herbivory rates, whereas in N. theifera, N compounds are also correlated with herbivory rates. Despite the differences in the strategies of these two species, they exhibit a similar efficiency of protection against natural enemies because their total herbivory rates are similar. The difference in their N defense allocation may imply benefits for survival under Cerrado conditions. We briefly discuss the oligotrophic habitat conditions of the studied plants and possible advantages of their strategies of N accumulation and metabolic uses.     

Isotopic niche overlap between sympatric Australian snubfin and humpback dolphins

Ecological niche theory predicts the coexistence of closely related species is promoted by resource partitioning in space and time. Australian snubfin (Orcaella heinsohni) and humpback (Sousa sahulensis) dolphins live in sympatry throughout most of their range in northern Australian waters. We compared stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) in their skin to investigate resource partitioning between these ecologically similar species. Skin samples were collected from live Australian snubfin (n = 31) and humpback dolphins (n = 23) along the east coast of Queensland in 2014–2015. Both species had similar δ¹³C and δ¹⁵N values and high (>50%) isotopic niche space overlap, suggesting that they feed at similar trophic levels, have substantial dietary overlap, and rely on similar basal food resources. Despite similarities, snubfin dolphins were more likely to have a larger δ¹⁵N value than humpback dolphins, indicating they may forage on a wider diversity of prey. Humpback dolphins were more likely to have a larger δ¹³C range suggesting they may forage on a wider range of habitats. Overall, results suggest that subtle differences in habitat use and prey selection are likely the principal resource partitioning mechanisms enabling the coexistence of Australian snubfin and humpback dolphins.     

Introduced Goldfish Affect Amphibians through Inhibition of Sexual Behaviour in Risky Habitats: an Experimental Approach

The introduction of alien species is one of the major causes of current and global biodiversity loss. The introduction of fish can be a particular threat to native amphibian populations, which are declining worldwide. One way for amphibians to persist in such altered environments is to adopt anti-predator strategies especially at the behavioural level. However, although it has been shown that avoidance behaviour may decrease the probability of being detected by a potential predator, little is known on the consequences on sexual behaviour. In this study, we tested the hypothesis that adult Alpine newts (Ichthyosaura alpestris) use shelters more often and exhibit less sexual activity in the presence of goldfish (Carassius auratus) and that they reduce sexual activity more in risky micro-habitats than in safe environments. To this end, we assessed behavioural patterns of adult newts in a replicated laboratory design. Goldfish were present in direct contact with newts in half of the tanks. Consistently throughout the study period, significantly more newts used shelter in the presence of fish than in their absence. Newts also significantly decreased their sexual activity level overall, but specially outside the shelter when they were in direct contact with fish. These results show that fish presence can affect newts in complex ways, such as through inhibition of their reproduction. Our work highlights that integrating behaviour in conservation studies is essential to understanding the patterns of coexistence and exclusion between introduced fish and amphibians.     

Unidirectional hybridization and reproductive barriers between two heterostylous primrose species in north-west Yunnan,China

Background and Aims

Heteromorphy in flowers has a profound effect on breeding patterns within a species, but little is known about how it affects reproductive barriers between species. The heterostylous genus Primula is very diverse in the Himalaya region, but hybrids there have been little researched. This study examines in detail a natural hybrid zone between P. beesiana and P. bulleyana.

Methods

Chloroplast sequencing, AFLP (amplified fragment length polymorphism) markers and morphological comparisons were employed to characterize putative hybrids in the field, using synthetic F₁s from hand pollination as controls. Pollinator visits to parent species and hybrids were observed in the field. Hand pollinations were conducted to compare pollen tube growth, seed production and seed viability for crosses involving different morphs, species and directions of crossing.

Key Results

Molecular data revealed all hybrid derivatives examined to be backcrosses of first or later generations towards P. bulleyana: all had the chloroplast DNA (cpDNA) of this species. Some individuals had morphological traits suggesting they were hybrids, but they were genetically similar to P. bulleyana; they might have been advanced generation backcrosses. Viable F₁s could not be produced with P. bulleyana pollen on P. beesiana females, irrespective of the flower morphs used. Within-morph crosses for each species had very low (<10 %) seed viability, whereas crosses between pin P. bulleyana (female) and pin P. beesiana had a higher seed viability of 30 %. Thus genetic incompatibility mechanisms back up mechanical barriers to within-morph crosses in each species, but are not the same between the two species. The two species share their main pollinators, and pollinators were observed to fly between P. bulleyana and hybrids, suggesting that pollinator behaviour may not be an important isolating factor.

Conclusions

Hybridization is strongly asymmetric, with P. bulleyana the only possible mother and all detected hybrids being backcrosses in this direction. Partial ecological isolation and inhibition of heterospecific pollen, and possibly complete barriers to F₁ formation on P. beesiana, may be enough to make F₁ formation very rare in these species. Therefore, with no F₁ detected, this hybrid zone may have a finite life span as successive generations become more similar to P. bulleyana.     

Functional Analysis of the Brassica napus L. Phytoene Synthase (PSY) Gene Family

Phytoene synthase (PSY) has been shown to catalyze the first committed and rate-limiting step of carotenogenesis in several crop species, including Brassica napus L. Due to its pivotal role, PSY has been a prime target for breeding and metabolic engineering the carotenoid content of seeds, tubers, fruits and flowers. In Arabidopsis thaliana, PSY is encoded by a single copy gene but small PSY gene families have been described in monocot and dicotyledonous species. We have recently shown that PSY genes have been retained in a triplicated state in the A- and C-Brassica genomes, with each paralogue mapping to syntenic locations in each of the three “Arabidopsis-like” subgenomes. Most importantly, we have shown that in B. napus all six members are expressed, exhibiting overlapping redundancy and signs of subfunctionalization among photosynthetic and non photosynthetic tissues. The question of whether this large PSY family actually encodes six functional enzymes remained to be answered. Therefore, the objectives of this study were to: (i) isolate, characterize and compare the complete protein coding sequences (CDS) of the six B. napus PSY genes; (ii) model their predicted tridimensional enzyme structures; (iii) test their phytoene synthase activity in a heterologous complementation system and (iv) evaluate their individual expression patterns during seed development. This study further confirmed that the six B. napus PSY genes encode proteins with high sequence identity, which have evolved under functional constraint. Structural modeling demonstrated that they share similar tridimensional protein structures with a putative PSY active site. Significantly, all six B. napus PSY enzymes were found to be functional. Taking into account the specific patterns of expression exhibited by these PSY genes during seed development and recent knowledge of PSY suborganellar localization, the selection of transgene candidates for metabolic engineering the carotenoid content of oilseeds is discussed.     

Natural Conditions Override Differences in Emergence Rhythm among Closely Related Drosophilids

Previous studies on adult emergence rhythm of Drosophila melanogaster (DM) done under semi-natural conditions have shown that emergence is correlated to daily changes in temperature, humidity and light at dawn. Recently we showed that under laboratory conditions D. ananassae (DA), a closely related species of DM exhibits patterns in its activity/rest rhythm distinct from the latter. Here, we report the results of a study aimed at examining whether this difference in activity/rest rhythm among species extends to other circadian behaviours such as the adult emergence rhythm under a more natural environment with multiple cyclic time cues. We monitored the adult emergence rhythm of recently wild-caught DM and DA populations in parallel with those of a related species D. malerkotliana (DK), both in the laboratory and under semi-natural conditions. We find that although DM, DK and DA showed marked difference from one another under laboratory conditions, such differences were not detectable in the emergence behaviour of these three species under semi-natural conditions, and that they respond very similarly to seasonal changes in the environment. The results suggest that seasonal changes in temperature and humidity contribute largely to the variation in adult emergence waveform in terms of gate width, phase and amplitude of the peak and day-to-day variance in the timing of the emergence peak. In all three species, seasons with cooler and wetter conditions make the rhythm less tightly gated, with low amplitude peak and high day-to-day variation in timing of the peak of emergence. We show that in nature the emergence rhythm of DM, DK and DA is strongly influenced by environmental factors such that in a given season all of them exhibit similar time course and waveform and that with the changing season, they all modify their emergence patterns in a similar manner.     

Modest enhancement of nitrogen conservation via retranslocation in response to gradients in N supply and leaf N status

Plant nutrient resorption, a ubiquitous mechanism of nutrient conservation, has often been proposed to be more pronounced in infertile than fertile habitats, and in species common to infertile compared to fertile habitats, because of the presumed advantage when nutrients are scarce. However, previous studies provide weak and inconsistent empirical support for these hypotheses, although few have examined intraspecific variation across well-quantified resource gradients. This study addresses intraspecific patterns of nutrient resorption for eight species across two N availability gradients on similar soils in an N-limited oak savanna ecosystem: a long-term fire frequency gradient with a negatively correlated N fertility gradient and a long-term N fertilization gradient. We hypothesized that both resorption proficiency (the minimum nutrient level retained in a senesced leaf) and efficiency (the proportional change in leaf nutrient concentration) would decrease with increasing soil N availability and plant N status. For the seven non-N fixers, either resorption proficiency or efficiency decreased modestly in treatments with higher N availability. In contrast, the legume Amorpha canescens Pursh had higher N levels in green and senesced leaves, and resorbed N much more weakly than the non-fixers, and did not respond in terms of proficiency or efficiency to soil N availability. Across all species and sites in each N fertility gradient, a scaling analysis showed greater resorption efficiency in plants with lower N concentrations. Our data suggest that species can have modest resorption responses reflective of soil nutrient availability and differences in resorption related to their N economy that represent mechanisms of nutrient conservation in nutrient-limited soils.     

Dominant plant species shift their nitrogen uptake patterns in response to nutrient enrichment caused by a fungal fairy in an alpine meadow

Niche partitioning by time, space and chemical forms has been suggested as an important mechanism to maintain species coexistence. Climate warming is assumed to increase soil nutrient availability through enhancing mineralization of soil organic matter in a variety of terrestrial ecosystems. However, few studies have yet examined how dominant plant species contribute to species coexistence when nutrient enrichment occurs in native ecosystems. We studied a single fairy ring (5 m diameter) in a Kobresia meadow in the Tibetan Plateau. This kind of rings is caused by a basidiomycete fungus Agaricus campestris, and is evidenced by dark-green vegetation boundaries. Nutrient enrichment occurs due to enhanced decomposition of soil organic matter (SOM) in the fungus growth zone of these rings. We conducted a short-term ¹⁵N labelling experiment and found that dominant plant species shifted their N uptake patterns and preferred N form (NO ₃ ^? , NH ₄ ⁺ , and amino acid N) in response to nutrient enrichment in an N-limited alpine meadow. The legume Gueldenstaedtia diversifolia had the lowest aboveground biomass among the five plant species studied at low available N level, although it mainly utilized ammonium (the most abundant N form). The two graminoids (Elymus nutans and Stipa aliena) demonstrated similar aboveground biomass at low and high available N levels, showing a similar pattern switching from NH ₄ ⁺ /NO ₃ ^? uptake outside the ring to glycine uptake in the annulus zone of the ring. The biomass of the forb Gentiana straminea differed significantly at low and high available N levels, but its N uptake pattern almost remained unchanged. Species therefore differed in their response to nutrient enrichment, most species showing chemical niche shifts instead of niche conservatism. This finding has important implications with regard to understanding the mechanisms responsible for species coexistence when natural nutrient enrichment is induced by climate warming in terrestrial ecosystems.     

Co-Flocculation of Yeast Species,a New Mechanism to Govern Population Dynamics in Microbial Ecosystems

Flocculation has primarily been studied as an important technological property of Saccharomyces cerevisiae yeast strains in fermentation processes such as brewing and winemaking. These studies have led to the identification of a group of closely related genes, referred to as the FLO gene family, which controls the flocculation phenotype. All naturally occurring S. cerevisiae strains assessed thus far possess at least four independent copies of structurally similar FLO genes, namely FLO1, FLO5, FLO9 and FLO10. The genes appear to differ primarily by the degree of flocculation induced by their expression. However, the reason for the existence of a large family of very similar genes, all involved in the same phenotype, has remained unclear. In natural ecosystems, and in wine production, S. cerevisiae growth together and competes with a large number of other Saccharomyces and many more non-Saccharomyces yeast species. Our data show that many strains of such wine-related non-Saccharomyces species, some of which have recently attracted significant biotechnological interest as they contribute positively to fermentation and wine character, were able to flocculate efficiently. The data also show that both flocculent and non-flocculent S. cerevisiae strains formed mixed species flocs (a process hereafter referred to as co-flocculation) with some of these non-Saccharomyces yeasts. This ability of yeast strains to impact flocculation behaviour of other species in mixed inocula has not been described previously. Further investigation into the genetic regulation of co-flocculation revealed that different FLO genes impact differently on such adhesion phenotypes, favouring adhesion with some species while excluding other species from such mixed flocs. The data therefore strongly suggest that FLO genes govern the selective association of S. cerevisiae with specific species of non-Saccharomyces yeasts, and may therefore be drivers of ecosystem organisational patterns. Our data provide, for the first time, insights into the role of the FLO gene family beyond intraspecies cellular association, and suggest a wider evolutionary role for the FLO genes. Such a role would explain the evolutionary persistence of a large multigene family of genes with apparently similar function.     

Social Behaviour of Individual Oval Squids (Cephalopoda,Teuthoidea, Loliginidae,Sepioteuthis lessoniana) within a Captive School

The social behaviour of individually identified squids was observed over a period of 4 mo. Squids were reared in captivity and maintained in a shoal of up to 18 animals. Distances between animals averaged 1.6 body lengths, similar to the 1.2 body lengths of wild teleost fish schools. Smaller animals schooled less frequently and were found at the periphery, and familiar and unfamiliar animals shoaled together. Eighty-six apparent reproductive events were observed. This species of squid, Sepioteuthis lessoniana, is not sexually dimorphic to the human eye and male squids copulated with both males and females but they directed their attention disproportionately towards females. Most copulations occurred more than a month before spawning, lasted only a few seconds and, unlike field observations on spawning grounds, were not followed by mate-guarding. Two body patterns are described that were used consistently in social interactions by both males and females; these patterns were both associated with reproductive behaviour but their exact function was unclear. A sister species, Sepioteuthis sepioidea, has been described as showing complex social behaviour; in S. lessoniana, no evidence for such behaviour was seen.     

Aggressive behaviour of male Microtus breweri and its demographic implications

Patterns of laboratory-determined aggressive behaviour and exploratory activity were examined in wild-caught male beach voles (Microtus breweri), an island-endemic species. This species is ecologically interesting because it has been found not to exhibit the super-annual population cycles typical of other rodents in the subfamily Microtinae. We found that body weight and reproductive condition were good predictors of aggressive behaviour, with heavier and reproductively active voles showing higher incidences of several stereotyped aggressive acts. Aggression also increased as the percentage of males and females who were reproductively active in the control population increased. Aggression was negatively correlated with population density, and more or less independent of dispersal rate. Dispersers and residents exhibited similar aggressive behaviour patterns. These observations are consistent with models which have proposed that population cycles in other vole species are controlled through the social interactions of individuals.