The feeding habits of the amphipods Lembos websteri Bate and Corophium bonnellii Milne Edwards

The gut contents of both Lembos websteri Bate and Corophium bonnellii Milne Edwards include detritus and sand grains. Lembos websteri collected from Laminaria holdfasts also contain strands of filamentous algae although Corophium bonnellii do not. In addition, macroalgal fragments and naviculoid diatoms are present, more often in Lembos websteri than in the guts of Corophium bonnellii. Occasionally, in animals collected from a sheltered algal turf, arthropod appendages may also be observed in the gut of Lembos websteri. The size of particles consumed by Lembos websteri is larger than in Corophium bonnellii. Feeding in both species is primarily by filtration but a variety of other methods may be used, including deposit-feeding, consumption of food of animal origin (including the species' own eggs), and consumption of tube material. These alternative techniques are brought to bear increasingly in poor feeding conditions. The behavioural sequences involved in these activities are described in detail and food preferenda related to the nutritional state of the animals. Defaecation in Lembos websteri (males and females) is normally followed by immediate refection of the faecal pellet but again is more noticeable when food is scarce. Gut-residence time in both species is ≈ 2 h at 13–15 °C. The proposition is made that Corophium bonnellii is a more efficient small-particle filter-feeder thanLembos websteri and that as such, it has met no evolutionary requirement to recycle its faeces (large particles) in the manner of L. websteri.Lembos, on the other hand, having a very “plastic” feeding repertoire appears to be better adapted to withstand periods of low food availability. The larger particles on which L. websteri feeds will be scarcer and will require more turbulence to keep them in suspension and even at exposed localities where L.websteri is more common, periods of calm occur, so flexibility in food-gathering technique is clearly advantageous. The more efficient small particle filter-feeding technique of Corophium bonnellii is well suited to a life in sheltered waters, where only the smallest particles remain in suspension for long periods, which is so typical of the genus.     

Tube building by the amphipods Lembos websteri Bate and Corophium bonnellii Milne Edwards

Lembos websteri Bate and Corophium bonnellii Milne Edwards are tubicolous. The morphology of tubes collected from Laminaria holdfasts is described and contrasted with that of tubes built amongst filamentous algae in a wave-sheltered environment. In both species tube length is a function of animal length. Corophium build tubes with openings raised above the substratum more commonly than Lembos, and will construct mats of tubes away from solid surfaces; Lembos tend to build against solid surfaces. Tube construction appears to depend upon the ‘knitting’ activity of the first two pairs of peraeopods, utilizing a secretion containing acid and sulphated mucopolysaccharides. The process of tube construction on substrata of known granulometry and the morphology of tubes built in the laboratory are described. The burrowing capabilities of both species are compared. On sand, Lembos tend to build on the sediment surface, whilst Corophium tend to burrow into the substratum. Corophium begin tube building after transference to a new situation much sooner than Lembos, with female Lembos starting to build sooner than male Lembos, Corophium can build a covering tube in < 15 min. In the absence of extraneous material, both species line glass tubes with a meshwork of mucoid filaments. In empty dishes, mucoid tubes are rarely built. The ability of Lembos to colonize wave-exposed sites may be related to its preference for more sheltered spaces within kelp holdfasts.     

A comparative study on the tubes and feeding behaviour of eight species of corophioid Amphipoda and their bearing on phylogenetic relationships within the Corophioidea

Observations are presented on mouthpart functional morphology, and on feeding, grooming and defaecatory behaviour of eight species of corophioid Amphipoda, viz. Corophium bonnellii, Lembos websteri, Aora gracilis, A. spinicornis, Gammaropsis nitida, Ericthonius punctatus, Jassa falcata and J. marmorata. These data are considered in relation to tube structure and amphipod posture in relation to the tube. All these species occupy double-ended cylindreical tubes made from ''amphipod silk'' secreted by the third and fourth preraeopods, incorporating sediment and other debris to varying degrees. Uniquely among this set of species, however, E. punctatus has a tube that is architecturally distinct. It tapers along its length and has a distinctive oblique main entrance at its widest end. This end is used preferentially. The other species studied use either opening with equal facility. Such a feature is adaptive in facilitating deployment of the antennae and shielding the head of E. punctatus.Two groupings of species are propose: group A which feed inside their tube using pleopod-induced through-tube currents, and group B which feed outside or at the entrance to their tube using external water currents. Group A includes C. bonnellii, L. websteri and the Aora species. Group B includes E. punctatus and the Jassa species. Gammaropsis nitida exhibits traits from both groups, adding weight to its perceived status as a genus representative of the stem corophioid. The ischyrocerid habit of externalizing food-gathering may be regarded as the first step along an evolutionary line leading to the rod-building podocerid types and ultimately towards the caprellids. All species examined show a degree of flexibility in their feeding habits which helps to explain the success of this taxon, which has radiated into a great diversity of aquatic biotopes.     

Tube irrigation by Lembos websteri Bate and Corophium bonnelli Milne Edwards (Crustacea: Amphipoda)

The pumping activity of the tubicolous amphipods Lembos websteri Bate and Corophium bonnelli Milne Edwards was measured by an apparatus that incorporated a sensitive pressure measuring device. The pumping rate of C. bonnelli was many times that of L. websteri when the animals were pumping in clean glass tubes. There was no significant difference in the pumping activity when animals were allowed to establish a detritus lined tube within a glass tube, although there was an indication that the pumping rate was increased in the case of L. websteri. The presence of clay in suspension had no significant effect on the pumping activity of either species but, in the case of C. bonnelli, the proportion of time spent pumping was greater when animals were in moving water than when in still water. No differences in the pumping activity were found between brooding and non-brooding mature female C. bonnelli nor between male and brooding female L. websteri.     

Turbidity and kelp holdfast amphipoda. I. Wales and S. W. England

Analyses are given of a collection of over 65,000 amphipods derived from kelp holdfasts surveyed around Wales and S.W. England. Faunal affinity is predominantly biased latitudinally, warmwater elements being confined to the extreme southwest. Total amphipod density is related to holdfast weight. There are no significant differences in amphipod density with differences in wave exposure or turbidity. Standardized density is, however, higher in west coast than in east coast holdfasts. Diversity is lower in turbid waters due to the increased dominance of a few species. The distribution, population structure, and fecundity of the previously implicated turbidity-susceptible species, Lembos websteri Bate and Corophium bonnellii Milne Edwards are examined in the light of new ethological information. Supporting and conflicting data are examined and mechanisms are suggested to show how suspended inorganic particles could be directly detrimental to both species. In addition, indirect effects involving climatic fluctuation and competitive interactions, particularly with C. sextonae Crawford emend. Hurley and Jassa falcata (Montagu) cannot be excluded.     

The effect of depth and sediment type on the spatial distribution of shallow soft-bottom amphipods along the southern Portuguese coast

The present study provides updated information on amphipod assemblages of the southern Portuguese coast, the most south-western part of continental Europe. The influence of depth and sediment type on the spatial distribution of shallow soft-bottom amphipods was tested by means of univariate and multivariate methods. Samples were collected down to 40?m depth within areas ranging from very fine sand to gravel. The ratio between density and diversity was found to increase with depth and from coarser to finer sediments. Both physical variables tested were found to play a major role in structuring amphipod assemblages. However, some dominance of depth was evident as samples were mainly separated according to this variable. Moreover, while only 15 taxa were common to all depth levels, 35 were collected from all sediment types. Both depth and sediment type may override the potential effects of geographical location, as samples from different areas along the southern Portuguese coast were biologically related. A peculiar habitat was sampled in this study: a maerl bed, which was confirmed to be a hotspot of density. Nevertheless, this habitat did not harbor a distinct amphipod assemblage but some taxa such as Caprella spp., Lembos websteri, lysianassids and Gammarella fucicola were preferentially collected there. The biogeographical analysis pointed out the occurrence of 8% of Mediterranean endemic species, confirming the influence of the Mediterranean Sea beyond the Strait of Gibraltar, and the Atlantic-Mediterranean feature of the amphipod fauna of this geographically relevant area. The present results support the need to interpret the spatial distribution patterns of marine species in relation to the combined effect of depth and sediment characteristics.     

Tetrachyrin,a new rearranged kaurenoid lactone,and diterpene acids from Tetrachyron orizabaensis and Helianthus debilis

(?)-Kaur-16-en-19-oic acid and tetrachyrin, a new rearranged kaurenoid lactone, were isolated from Tetrachyron orizabaensis var. websteri and Helianthus debilis ssp. debilis. The latter species also afforded angeloylgrandifloric acid.     

Isolation and Characterization of Microsatellite Markers for <Emphasis Type="Italic">Viola websteri</Emphasis> (Violaceae) and Cross-Species Amplification within the Genus <Emphasis Type="Italic">Viola</Emphasis>

We isolated and characterized microsatellite loci in Viola websteri (Violaceae), an endangered species from Korea and endemic to Northeast Asia. A total of 27 microsatellite loci were developed and tested in Korean and Chinese populations. The number of alleles per locus varied from two to eight. The observed and expected heterozygosities within two populations were 0.000 to 1.000 and 0.080 to 0.816, respectively. Korean and Chinese populations were clearly distinguished by the private alleles from 16 loci. A total of 21 loci out of the 27 developed loci were successfully cross-amplified in 39 other Viola species. We believe that these microsatellite loci will be useful for future studies on genetic diversity and population structure of V. websteri, as well as other Viola species.     

Sites of self-pollen tube inhibition in Papaveraceae (sensu lato)

Papaver rhoeas (Papaveraceae) has a well-characterized gametophytic self-incompatibility system in which self-pollen tube growth ceases either just before, or just after, emergence from the copal aperture. Papaver flowers are unusual, however, in having flat stigmatic rays sitting directly on top of the hard ovary and no style. Immediate self-pollen arrest might be required with this floral architecture. There is much variation in floral architecture among Papaveraceae and self-incompatibility is widespread. However, there are no reports of the site of self-pollen tube inhibition in Papaveraceae other than P. rhoeas. We examined the site of self-pollen tube inhibition in four species (Argemone munita, Lamprocapnos spectabilis, Eschscholzia californica, and Platystemon californicus) representing a broad phylogenetic and morphological sample of Papaveraceae. Squash preparation was used for species with soft stigmas whereas woody tissue was sectioned with a cryostat and images were stitched into a mosaic to visualize pollen tubes on whole stigmas. For three species, self-pollen tube inhibition appeared similar to that described for P. rhoeas. Self-pollen tubes were arrested before any substantial penetration of female tissue and usually did not grow longer than 100?μm. In the fourth species, A. munita, self-pollen tubes grew up to 500?μm in length. However, self-pollen tubes appeared to grow along the stigmatic spines, and growth ceased once tubes contacted the stigma surface. Despite variation in floral architecture, rapid arrest of self-pollen tubes occurred before or just after penetration of female tissue in all species, consistent with the hypothesis that members of the family share the same incompatibility mechanism.     

Effect of native natural enemies on the population dynamics of the grape mealybug,Pseudococcus maritimus (Hom.: Pseudococcidae), in apple and pear orchards

The impact of native natural enemies on populations of the grape mealybug,Pseudococcus maritimus (Ehrhorn) in apple and pear orchards was assessed using a combination of techniques, including exclusion cages, limb-banding, and visual inspection of shoots and fruits. The complex of native natural enemies (which included two encyrtid parasitoids, (Pseudaphycus websteri Timberlake andMayridia species), a coccinellid beetle (Hyperaspis lateralis Mulsant), and a chamaemyiid fly (Leucopis verticalis Malloch), provided reasonably good control in orchards that had not been treated with insecticides for one to two years. However, surveys indicated that most of these species were absent from orchards regularly sprayed with pesticides.     

Mating systems and sex ratios in the egg parasitoids, Trichogramma dendrolimi and T. papilionis (Hymenoptera: Trichogrammatidae)

Arrhenotokous parthenogenesis was confirmed in Trichogramma dendrolimi and T. papilionis. Subsequently the possible links between mating systems and biological traits of males, and sex ratios, were investigated in these two species, using Papilio xuthus eggs for their hosts. T. dendrolimi males attained 100% insemination of females in the parasitized host before egress from it. The high insemination rate was guaranteed by male precedence in emergence, lack of courtship in mating behaviour and short copulation time, combined with a long stay of emerging wasps within the host. The males were often the last to leave a host and made no mating attempts outside the host. Most but not all T. papilionis females were also inseminated in the host. The lower insemination rate of this species resulted from almost simultaneous emergence of both sexes, which prevented males from mating with inactive females. Another mating site of T. papilionis was just outside the host, which males left prior to most females. A reduced possibility of outbreeding was inferred in T. dendrolimi on the grounds that males were short-lived, frequently failed to expand their wings and died in the host. The reduced outbreeding was reflected in considerably lower sex ratios in T. dendrolimi.     

AtLURE1/PRK6-mediated signaling promotes conspecific micropylar pollen tube guidance

Reproductive isolation is a prerequisite to form and maintain a new species. Multiple prezygotic and postzygotic reproductive isolation barriers have been reported in plants. In the model plant, Arabidopsis thaliana conspecific pollen tube precedence controlled by AtLURE1/PRK6-mediated signaling has been recently reported as a major prezygotic reproductive isolation barrier. By accelerating emergence of own pollen tubes from the transmitting tract, A. thaliana ovules promote self-fertilization and thus prevent fertilization by a different species. Taking advantage of a septuple atlure1null mutant, we now report on the role of AtLURE1/PRK6-mediated signaling for micropylar pollen tube guidance. Compared with wild-type (WT) ovules, atlure1null ovules displayed remarkably reduced micropylar pollen tube attraction efficiencies in modified semi-in vivo A. thaliana ovule targeting assays. However, when prk6 mutant pollen tubes were applied, atlure1null ovules showed micropylar attraction efficiencies comparable to that of WT ovules. These findings indicate that AtLURE1/PRK6-mediated signaling regulates micropylar pollen tube attraction in addition to promoting emergence of own pollen tubes from the transmitting tract. Moreover, semi-in vivo ovule targeting competition assays with the same amount of pollen grains from both A. thaliana and Arabidopsis lyrata showed that A. thaliana WT and xiuqiu mutant ovules are mainly targeted by own pollen tubes and that atlure1null mutant ovules are also entered to a large extent by A. lyrata pollen tubes. Taken together, we report that AtLURE1/PRK6-mediated signaling promotes conspecific micropylar pollen tube attraction representing an additional prezygotic isolation barrier.

A modified ovule targeting assay revealed that AtLURE1/PRK6-mediated signaling promotes micropylar guidance of Arabidopsis thaliana pollen tubes while discriminating tubes of related Arabidopsis lyrata.     

Sex ratios and sex-biased infection behaviour in the entomopathogenic nematode genus Steinernema

In experimentally infected insects, the sex ratio of first generation nematodes of five species of Steinernema was female-biased (male proportion 0.35-0.47). There was a similar female bias when the worms developed in vitro (0.37-0.44), indicating that the bias in these species is not due to a lower rate of infection by male infective juveniles (IJs). Experimental conditions influenced the proportion of males establishing in insects, indicating that male and female IJs differ in their behaviour. However, there was no evidence that males are the colonising sex in any species, contrary to what has previously been proposed. Time of emergence from the host in which the nematodes had developed influenced sex ratios in experimental infections. In three species (Steinernema longicaudum, Steinernema glaseri and Steinernema kraussei), early emerged nematodes had a higher proportion of males than those that emerged later, with the reverse trend for Steinernema carpocapsae and Steinernema feltiae. In a more detailed in vitro study of S. longicaudum, the proportion of males was similar whether or not the nematodes passed through the developmentally arrested IJ stage, indicating that the female bias is not due to failure of males to exit this stage. The sex ratio in vitro was independent of survival rate from juvenile to adult, and was female-biased even when all juveniles developed, indicating that the bias is not explained by failure of males to develop to adults. The female-biased sex ratio characteristic of Steinernema populations appears to be present from at least the early juvenile stage. We hypothesise that the observed female bias is the population optimal sex ratio, a response to cycles of local mate competition experienced by nematodes reproducing within insect hosts interspersed with periods of outbreeding with less closely related worms following dispersal.     

Mating Behavior of Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae): Potential for Outbreeding in a Predominantly Inbreeding Species

The mating behavior of the quasi-gregarious egg parasitoid Trissolcus basalis (Wollaston) was investigated under field conditions. Trissolcus basalis has female-biased sex ratios and is a protandrous species, with males emerging 1–2 days before females. Males competed aggressively for control of the egg mass, with one male assuming dominance and control of the egg mass, although changes in dominance occurred at least once on each egg mass observed. Typical mating behavior involved the dominant male mating his sisters immediately upon their emergence from the egg mass. These behaviors are characteristic of an inbreeding species that manifests local mate competition. However, several aspects of the mating behavior of T. basalis are inconsistent with that of an inbreeding species. Over 18% of emerging females were not mated by the dominant male upon emergence, 13% of females were not observed to be mated at all and may have left their natal site as virgins, 25% of females were mated multiple times and sometimes by multiple males, females remained near the natal site for up to several hours after emergence before emigrating, and males dispersed away from the natal site during female emergence. Trissolcus basalis may be a predominantly inbreeding species but its emergence and mating behavior suggest that low-frequency outbreeding is also likely to occur.     

Biodiversity of benthic invertebrates and organic matter processing in shallow marine sediments: an experimental study

The main objective of this study was to measure the impact of benthic invertebrate diversity on processes occurring at the water-sediment interface. We analyzed the effects of interactions between three shallow water species (Cerastoderma edule, Corophium volutator, and Nereis diversicolor). The impacts of different species richness treatments were measured on sediment reworking, bacterial characteristics, and biogeochemical processes (bromide fluxes, O₂ uptake, nutrient fluxes, and porewater chemistry) in sediment cores. The results showed that the three species exhibited different bioturbation activities in the experimental system: C. edule acted as a biodiffusor, mixing particles in the top 2 cm of the sediments; C. volutator produced and irrigated U-shaped tubes in the top 2 cm of the sediments; and N. diversicolor produced and irrigated burrow galleries in the whole sediment cores. C. edule had minor effects on biogeochemical processes, whereas the other species, through their irrigation of the burrows, increased the solute exchange between the water column and the sediment two-fold. These impacts on sediment structure and solute transport increased the O₂ consumption and the release of nutrients from sediments. As N. diversicolor burrowed deeper in the sediment than C. volutator, it irrigated a greater volume of sediments, with great impact on the sediment cores.Most treatments with a mixture of species indicated that observed values were often lower than predicted values from the addition of the individual effects of each species, demonstrating a negative interaction among species. This type of negative interaction measured between species on ecosystem processes certainly resulted from an overlap of bioturbation activities among the three species which lived and foraged in the same habitat (water-sediment interface). All treatments with N. diversicolor (in isolation and in mixture) produced similar effect on sediment reworking, water fluxes, nutrient releases, porewater chemistry, and bacterial characteristics. Whichever species associated with N. diversicolor, the bioturbation activities of the worm hid the effect of the other species. The results suggest that, in the presence of several species that use and modify the same sediment space, impact of invertebrates on ecosystem processes was essentially due to the most efficient bioturbator of the community (N. diversicolor). In consequence, the functional traits (mode of bioturbation, depth of burrowing, feeding behaviour) of an individual species in a community could be more important than species richness for some ecosystem processes.     

Effect of population density and presence of a potential competitor on the growth rate of the mud snail Hydrobia ulvae (Pennant)

The mud snail Hydrobia ulvae (Pennant) and the amphipod Corophium arenarium Crawford were used in a series of laboratory and field experiments to investigate the hypothesis that natural populations of such deposit-feeders live under conditions of resource limitation. Growth rates of Hydrobia confined at several different densities within cages in the field and in laboratory microcosms indicated that density-dependent reduction of growth rate and increased mortality occurred at densities found in natural populations. Similar experiments failed to provide any evidence that Corophium mortality or behaviour was modified by the presence of Hydrobia or members of its own species; nor did the presence of Corophium cause a reduction in growth rate of Hydrobia.     

Effect of female group size on harem male roosting behavior of the Indian short-nosed fruit bat <Emphasis Type="Italic">Cynopterus sphinx</Emphasis>

Mate guarding has been known to incur costs and cause constraints for harem males in many polygynous species. However, the effect of female group size on the harem male’s time budget in bats has received very limited attention. The Indian short-nosed fruit bat, Cynopterus sphinx, exhibits resource defense polygyny, in which tent roosting males construct tents and defend multiple female bats. We studied the effect of female group size on three aspects of harem male behavior: social grooming by reciprocal licking, tent maintenance, and tent guarding in the mast tree Polyalthia longifolia. In the process of reciprocal licking, all the bats in the harem were drenched in saliva before emergence, and this activity was positively and significantly correlated with female group size. Once females departed for foraging, harem males remained in their respective tents at night-time between intermittent foraging bouts and engaged in tent maintenance and tent guarding. Time invested by harem male bats in tent maintenance and tent guarding were positively and significantly correlated with female group size. Harem males extended their presence in tent by utilizing tents as feeding roosts. Female group size also influenced the emergence time of harem male bats, where males with largest group emerged later than did the smallest group. Likewise, harem male with the smallest group had more time available for foraging than the male with the largest group. Findings of this study suggest that having a larger harem may indeed be costly for the males by reducing their foraging time.     

The fate of sheep-dispersed seeds: Plant species emergence and spatial patterns

Sheep epizoochory has often been proposed as an important vector which can help to overcome the dispersal limitation of plants in fragmented landscapes. In order to evaluate the contribution of herbivores to recruitment especially of target species, the dispersal and post-dispersal fate of such seeds must be known. In a field experiment sheep with seeds of mainly target species (experimentally attached to their coats) were present at three sand plots for 24 h. Natural epizoochorous dispersal was already shown for most of the species in our study area. Seed detachment, trampling intensity and seed shadow were measured; seedling emergence and survival were recorded over an 8-month period. In addition, the effect of sheep trampling on seedling emergence and survival of two threatened species, Jurinea cyanoides and Koeleria glauca, were studied.     

Evidence for the circadian gene period as a proximate mechanism of protandry in a pollinating fig wasp

Protandry in insects is the tendency for adult males to emerge before females and usually results from intra-sexual selection. However, the genetic basis of this common phenomenon is poorly understood. Pollinating fig wasp (Agaonidae) larvae develop in galled flowers within the enclosed inflorescences (‘figs’) of fig trees. Upon emergence, males locate and mate with the still galled females. After mating, males release females from their galls to enable dispersal. Females cannot exit galls or disperse from a fig without male assistance. We sampled male and female Ceratosolen solmsi (the pollinator of Ficus hispida) every 3 h over a 24 h emergence period, and then measured the expression of five circadian genes: period (per), clock (clk), cycle (cyc), pigment-dispersing factor (pdf) and clockwork orange (cwo). We found significant male-biased sexual dimorphism in the expression of all five genes. per showed the greatest divergence between the sexes and was the only gene rhythmically expressed. Expression of per correlated closely with emergence rates at specific time intervals in both male and female wasps. We suggest that this rhythmical expression of per may be a proximate mechanism of protandry in this species.