The life cycles of two coexisting copepods, <Emphasis Type="Italic">Calanus chilensis</Emphasis> and <Emphasis Type="Italic">Centropages brachiatu</Emphasis>s,in the upwelling zone off northern Chile (23°S)

A time series study of the copepods Calanus chilensis and Centropages brachiatus was carried out at the coastal upwelling zone of Mejillones (23°S, northern Chile), to analyze their annual life cycles in association with upwelling variation. These species co-exist in the upwelling zone. Weekly sampling of zooplankton and oceanographic variables including Chlorophyll-a and phytoplankton composition were obtained during January–December 2002 at a fixed station (ca. 90 m depth). Stages of abundances, their proportions, changes in body length of adult females, sex ratio and egg production rate (EPR), were used as proxies to examine copepods’ demography. Upwelling, assessed by weekly Ekman transport and oceanographic conditions, was intermittent throughout the year with lack of periodicity components. Populations of both copepod species did not correlate with these non-predictable upwelling events. C. chilensis reproduced year-round and the population showed ca. 15 peaks of adults with an average time interval between peaks of 20 days. C. brachiatus showed a similar life cycle, also having 15 peaks of adults at about 22 days of time intervals. Cross-correlation functions and spectral analysis showed that both populations correlated positively through time, but not in phase, evidencing a time lag for their reproductive cycles. The lag was also evident in their population abundances. Both species differ in their development rates and this may result in non-in phase life cycles. Our findings suggest that species-dependent attributes, such as development rates, modulated by adaptations to temperature, might impose constraints in the species life cycles determining the population cycles. Such attributes must be considered when modeling and understanding population dynamics and secondary production of copepods.     

(<Emphasis Type="Italic">Z</Emphasis>)-9-Pentacosene − contact sex pheromone of the locust borer, <Emphasis Type="Italic">Megacyllene robiniae</Emphasis>

Summary. Male locust borers, Megacyllene robiniae (Förster), responded to females only after contacting them with their antennae, indicating that mate recognition was mediated by a contact sex pheromone. GC-MS analyses of whole-body extracts of males and females determined that the profiles of compounds in the extracts were qualitatively similar, but differed considerably in the ratios of compounds between sexes. Biological activities of reconstructed blends of the most abundant straight-chain (nC₂₃, nC₂₄, nC₂₅, nC₂₆), methyl-branched (3me-C₂₃, 3me-C₂₅), and unsaturated (Z9:C₂₃, Z9:C₂₅, Z9:C₂₇ compounds in extracts from females were assessed in arena bioassays, assessing four distinct steps in the mating behavior sequence of males (orientation, arrestment, body alignment, mounting and attempting to couple the genitalia). Males were unresponsive to freeze-killed, solventwashed females treated with blends of straight-chain and methyl-branched alkanes, but responded strongly to females treated with the blend of alkenes. Further trials determined that the complete sequence of mating behaviors, up to and including coupling the genitalia, was elicited by Z9:C₂₅ alone. Z9:C₂₅ comprised 16.4 ± 1.3% of the total hydrocarbons in whole-body hexane extracts of females and was co-dominant with two other hydrocarbons that were not active. In contrast, in solid phase microextraction (SPME) wipe samples from several areas of the cuticle, Z9:C₂₅ appeared as the single dominant peak, comprising 34.6 – 37.8% of the sampled hydrocarbons. Our data indicate that Z9:C₂₅ is a contact sex pheromone of M. robiniae, being the most abundant hydrocarbon on the surface of the cuticular wax layer of females where it is readily accessible to the antennae of males.     

Covering behaviour in <Emphasis Type="Italic">Paracentrotus</Emphasis><Emphasis Type="Italic">lividus</Emphasis>: is light important?

"Covering" or "heaping" behaviour is common to a number of regular echinoids living in a variety of different habitats. Many theories have been proposed to explain this behaviour, among which are several that link covering to light, including ultraviolet (UV) light. However, previous investigations of this light theory have been largely qualitative. In the present study, we used a systematic laboratory protocol to examine quantitatively the covering behaviour of the shallow water echinoid, Paracentrotus lividus, under four different light regimes: white light (400-700 nm), UV-A+B (315-400 nm), UV-A (320-400 nm), and darkness. These experiments demonstrated that light, in particular UV light, influences the covering behaviour of P. lividus. Under the UV regimes, significantly more individuals were found to display covering behaviour, and individuals spent more time at the base of aquaria, farthest from the light source. Moreover, covering items were retained for the longest period of time under the UV-A+B regime. We propose that protection against the harmful effects of UV radiation may be one of the functions of covering behaviour in P. lividus.     

Laboratory investigation of <Emphasis Type="Italic">Fallopia</Emphasis> × <Emphasis Type="Italic">bohemica</Emphasis> fruits dispersal by watercourses

Seed and fruit dispersal along watercourses favours the long-distance migration of invasive species, not only for aquatic or wetland species, but also for terrestrial wind-dispersed plants, like the Japanese knotweed. The present paper aims at investigating the role of watercourses in the dispersal of the knotweed due to its frequent occurrence on riverbanks and production of fertile achenes (type of fruit of the Japanese knotweed). This dispersal occurs along two steps after the fruits deposit on the water surface: floatation first and then sinking towards the bottom of the watercourse. Regarding the first step, the effects of agitation of the water, temperature, surface tension and luminosity on the achenes floatability are experimentally studied. While no influence of luminosity is observed, an increase of temperature greatly decreases the floating time. Floating time also decreases as the contact between water and the fruit is enhanced (through submersion of achenes, agitation of the water or lower surface tension). Regarding the second step, the fall velocity of the fruits in water at rest is measured and appears to be independent of the seed history (floating time). 3D helical motions are systematically observed with constant tangential velocity with respect to the falling velocity. The trajectory of the fruits in a shear flow is then measured and the evolution of their velocity components along the sinking process is discussed. Finally, the contribution of both steps to the long-distance migration of the seeds is estimated.     

Sibling species or poecilogony in the polychaete<Emphasis Type="Italic"> Scoloplos armiger</Emphasis>?

In marine invertebrates multiple modes of development, or poecilogony, may occur in a single species. However, after close examination, many of such putative cases turned out to be sibling species. A case in point may be the cosmopolitan orbiniid polychaete Scoloplos armiger, which inhabits marine shallow sediments. In addition to the well-known direct, holobenthic development from egg cocoons, pelagic larvae have also been described. Our culture experiments revealed a spatially segregated source of the two developmental modes. All females of an intertidal population produced egg cocoons and no pelagic larvae. All but 2 out of 15 females of an adjacent subtidal population produced pelagic larvae and no egg cocoons. Based on these results we performed a molecular genetic analysis (RAPD-PCR) on three intertidal and four subtidal populations in the North Sea. Selected samples from all sites were analysed also by the AFLP method. We found significantly higher genetic diversity within subtidal than within intertidal populations. This is consistent with a wider dispersal by pelagic larvae and a smaller effective population size when development is holobenthic. Total genetic divergence is not related to distance but to the intertidal/subtidal division. We suggest that S. armiger actually represents two sibling species.     

Sibling competition in guinea pigs (<Emphasis Type="Italic">Cavia aperea</Emphasis> f. <Emphasis Type="Italic">porcellus</Emphasis>): scrambling for mother’s teats is stressful

Lactation is the most energy-intense period in the life of a female mammal. This can cause severe conflict between mother and offspring over the duration of lactation but also between siblings over the amount of milk each pup gets from its mother. Thus, competitive interactions between siblings are expected, and competition is likely to increase with litter size, particularly in species where the number of offspring exceeds the number of teats. We studied sibling competition in the domestic guinea pig (Cavia aperea f. porcellus), which has two teats, but frequently bears litters of up to five pups. By cross-fostering we created non-competition (control) litters with two pups and competition litters with four pups and observed nursing behaviour on days 5, 10, 15 and 20 postpartum. Pups of larger litters had lower growth rates, indicating increased competition among siblings in these litters. Pups of larger litters had to wait longer for access to a teat and spent less time suckling than pups of smaller litters but ate more solid food instead. Additionally, we manipulated the individual short-term need of pups by separating half of the pups of each litter for 2 h from their mothers before observation. Within a litter, hungry pups achieved access to milk faster and spent more time suckling than non-hungry pups. Pups competed mostly by scramble competition. Aggressive interactions occurred only in large litters. Pups of large litters had higher cortisol levels than pups in small litters. These effects decreased with age as pups became increasingly independent of maternal milk. Pup behaviour appears to fit better with models of scramble competition than with those of honest signalling. This contribution is part of the special issue “Sibling competition and cooperation in mammals” (guest editors: Robyn Hudson and Fritz Trillmich).     

Genetic variation among life-history stages of mussels in a<Emphasis Type="Italic"> Mytilus edulis–M</Emphasis>.<Emphasis Type="Italic"> trossulus</Emphasis> hybrid zone

The mussel species Mytilus edulis L. and M. trossulus Gould coexist and hybridize throughout a large area that includes the north coast of Maine and Atlantic Canada. Previous studies provided genetic evidence for limited hybridization between the two species for mussels >15 mm. The present study used two genetic markers (ITS, Glu-5) to examine the genetic composition of early life-history stages by sampling veliger and pediveliger larvae, juveniles (<2.0 to 15.0 mm shell length) and adults (>15 mm shell length) in Trinity Bay, Newfoundland, during three consecutive years (1995–1997) to determine if differential mortality limits the relative abundance of hybrids. The relative frequency of the two species and the different hybrid genotypes was similar among the larvae, juveniles and small adult mussels. The double hybrid genotype (F₁-like) was the rarest genotype observed. There was no evidence for differential mortality during the early life-history stages, and factors limiting production of hybrids appear to operate before the late larval stage. The observed frequency of hybrids is probably due to a combination of pre- and postzygotic reproductive isolating mechanisms operating early in the mussels life history. M. trossulus dominated the early life-history stages, possibly due to a higher population density and a greater reproductive output than M. edulis. Differential mortality may explain the observed decrease in frequency of M. trossulus and increase in frequency of M. edulis with increasing shell length. A similar frequency of hybrid mussels from larvae to the size class of 55 mm shell length may indicate a rate of mortality intermediate between the two parental species. The M. edulis–M. trossulus hybrid zone appears to be maintained by reproductive isolating mechanisms limiting the production of hybrids and life-history differences that allow the two species to coexist.Communicated by J.P. Grassle, New Brunswick     

Life cycle of <Emphasis Type="Italic">Oithona</Emphasis> <Emphasis Type="Italic">similis</Emphasis> (Copepoda: Cyclopoida) in Kola Bay (Barents Sea)

The annual population dynamics (nauplii, old copepodites CIV–CV and adults) and seasonal variations in reproductive parameters of the cyclopoid copepod Oithona similis were investigated on the basis of the data 1999–2006 in Kola Bay, a large subarctic fjord in the Barents Sea. Population density of O. similis ranged from 110 to 9,630 ind m⁻³ and averaged 1,020 ± 336 ind m⁻³. The relative abundance of adults was high during winter (~60%). At the end of winter (mid-March), the population included a large percentage of later-stage copepodites (stage CIV 23% and stage CV 57%). There were two periods of mass spawning, in late June and September. Autumn and summer generations strongly differed in abundance, average prosome length (PL), clutch size (CS), egg diameter (D), egg production rates (EPR and SEPR) and female secondary production. Average PL decreased with increasing water temperature, while D and CS were strongly correlated with PL but unaffected by temperature. Annual average EPR and SEPR were 0.55 ± 0.18 eggs female⁻¹ day⁻¹ and 0.0011 ± 0.003 day⁻¹, respectively. Female secondary production averaged 0.8 ± 0.3 μg C m⁻³ day⁻¹ (range 0.001–3.58). There were positive relationships between abundance, EPR, SEPR, production and water temperatures. Reproductive parameters appeared to be controlled by hydrological factors and food conditions.     

Concentration of ascorbic acid and innate immune effectors in <Emphasis Type="Italic">Engraulis ringens</Emphasis> and <Emphasis Type="Italic">Strangomera bentincki</Emphasis> during their main spawning period (2007–2008) in the Humboldt current system off Chile

In this study, we determined variations in ascorbic acid (AA) concentrations in the gonads and liver, and of the innate immune response (lysozyme activity and antiprotease) in the plasma of Engraulis ringens (anchoveta) and Strangomera bentincki (common sardine) during their reproductive seasons of 2007 and 2008 in the central area of the Humboldt Current. During the main spawning season in 2007, colder environmental conditions, higher phytoplankton biomasses, low dinoflagellate abundance and the dominance of large copepods in winter-early spring and of small copepods in late spring contributed to explain higher AA concentrations in anchoveta tissues. During the warmer year of 2008, lower phytoplankton biomasses, a notable increase of dinoflagellate abundance and the dominance of small-size copepods in winter and of larger copepods in spring occurred along with a drastic drop in AA concentrations in anchoveta tissues. These results contrasted with those in common sardine, a species in which AA did not vary as much as in anchoveta. An inverse trend between AA concentration in the liver and the gonadosomatic index suggested the AA utilization during gonadic maturation. Innate immune parameters were determined for the first time in small pelagic fish and constitute a new baseline data to evaluate their natural response to environmental changes. Changes in the distribution and abundance of edible taxa containing AA and edible for fish along with variations in AA in fish tissues during their reproductive season could become good indicators of the overall fish physiological condition resulting from variations in the trophic web structure.     

Simulation of exchange flow between open water and floating vegetation using a modified RNG <Emphasis Type="Italic">k</Emphasis>-<Emphasis Type="Italic">ε</Emphasis> turbulence model

The paper focuses on the numerical simulation of the exchange flow between open water and floating vegetation, which plays an important role in maintaining the ecological balance by transporting nutrient matter. The simulation was conducted using a new solver developed upon OpenFOAM. A modified RNG k-ε turbulence model, which is expected to model both the high- and low-Reynolds number flows correctly, was used to determine the eddy viscosity. Several particular terms were added into the momentum equations and turbulence model equations to model the effects of vegetation and buoyancy. Among these terms, the term for the effect of vegetation in the ε-equation was re-modelled. The model was validated by properly predicting the profiles of mean velocity and turbulent kinetic energy for flows through suspended canopies. The density flow between open and vegetated water was simulated with the same conditions as those of the experiment conducted by Zhang and Nepf. The predicted results agreed well with the experimental data and provided more detailed information of such exchange flow. The convection between the root layer and the layer beneath the roots, which was not observed in the experiment, was observed in the numerical simulation.     

Variation in colony geometry modulates internal light levels in branching corals, <Emphasis Type="Italic">Acropora humilis</Emphasis> and <Emphasis Type="Italic">Stylophora pistillata</Emphasis>

Colonial photosynthetic marine organisms often exhibit morphological phenotypic plasticity. Where such plasticity leads to an improved balance between rates of photosynthesis and maintenance costs, it is likely to have adaptive significance. To explore whether such phenotypic plasticity leads to more favourable within-colony irradiance for reef-building branching corals, this relationship was investigated for two coral species Acropora humilis and Stylophora pistillata, along a depth gradient representing light habitats ranging from 500 to 25 μmol photons m⁻² s⁻¹, during 2006 at Heron Island, Great Barrier Reef (23.44°S, 151.91°E). In the present study changes in flow-modulated mass transfer co-varied with light as a function of depth. In low-light (deep) habitats, branch spacing (colony openness) in A. humilis and S. pistillata was 40–50% greater than for conspecifics in high-light environments. Also, branches of A. humilis in deep water were 40–60% shorter than in shallow water. Phenotypic changes in these two variables lead to steeper within-colony light attenuation resulting in 38% higher mean internal irradiance (at the tissue surface) in deep colonies compared to shallow colonies. The pattern of branch spacing was similar for S. pistillata, but this species displayed an alternate strategy with respect to branch length: shade adapted deep and cave colonies developed longer and thinner branches, allowing access to higher mass transfer and irradiance. Corals in cave habitats allowed 20% more irradiance compared to colonies found in the deep, and had a 47% greater proportion of irradiance compared to colonies in the shallow high-light environment. Such phenotypic regulation of internal light levels on branch surfaces partly explains the broad light niches of many branching coral species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The hydrozoan coral<Emphasis Type="Italic"> Millepora dichotoma</Emphasis>: speciation or phenotypic plasticity?

This article describes ecological and biological differences between two morphs of the Red Sea fire coral Millepora dichotoma. The species is divided into two main morphs: branching and encrusting, which were found to differ both in color and morphology. Each morph has two or more sub-morphs. A total of 372 M. dichotoma colonies were examined in a census at two study sites in the Gulf of Elat. Colony size and abundance of the two morphs were found to differ significantly between sites. Experimental examination of each morph's morphological plasticity revealed different growth rates and difference in growth plasticity between the branching and the encrusting morph. Most of the fragments from the branching colonies (94%) attached to experimentally placed Plexiglas substrate, compared with much less attachment by the encrusting fragments (11%). The growth form of the branching morph on the Plexiglas switched to encrusting, spreading over and covering the substrate. When the new encrusting colony reached the edge of this substrate, it started to produce tips, and returned to growth in the classic branching form. The encrusting morph did not change its growth form. Following attachment of the original fragments of the branching morph to the substrate, 8.1% of them produced new tips. When the original branches were removed, after converting to encrusting growth form, 19% of the fragments produced new tips. The capsule size of nematocysts of the two morphs was also significantly different (t-test, P<0.05). Molecular data (ITS region) clearly demonstrate that these two M. dichotoma morphs differ considerably. Molecular evidence (srRNA) from the symbiotic zooxanthellae also shows a different pattern of clades in the hosts. The ecological, biological and molecular data thus attest to the two morphs being distinguishable. Contrary to previous reports, we consequently suggest that the two morphs of M. dichotoma found in the Gulf of Elat are actually two distinct species.Communicated by O. Kinne, Oldendorf/Luhe     

Is <Emphasis Type="Italic">Hippolyte williamsi</Emphasis> gonochoric or hermaphroditic? A multi-approach study and a review of sexual systems in <Emphasis Type="Italic">Hippolyte</Emphasis> shrimps

Sexual systems vary considerably among caridean shrimps and while most species are gonochoric, others are described as sequential protandric hermaphrodites or simultaneous hermaphrodites with an early male phase. At present, there is confusion about the sexual system exhibited by several species mostly because those studies attempting to reveal their sexual system draw inferences solely from the distribution of the sexes across size classes. Here we investigated the sexual system of the shrimp Hippolyte williamsi from Chile to determine if the species is protandric or gonochoric with sexual dimorphism (males smaller than females). Morphological identification and size frequency distributions indicated that the population comprised small males, small immature females, and large mature females, which was confirmed by dissections. No transitional individuals were found. Males maintained in the laboratory molted 1–8 times, and many grew up to reach sizes observed in only a small fraction of males in the field. No indication of sex change was recorded. Our results indicate that H. williamsi is a sexually dimorphic gonochoric species and emphasizes the importance of using several kinds of evidence (size measurements, growth experiments, morphological dissections, and histological studies) to reveal the sexual system of Hippolyte species. Whether the observed size dimorphism between males and females in many species of Hippolyte is expression of contrasting sexual and natural selection, and whether divergent sexual fitness functions can contribute to the evolution of hermaphroditism remains to be revealed in future studies.     

Embryonic development and maternal–embryo relationships of the Portuguese dogfish <Emphasis Type="Italic">Centroscymnus coelolepis</Emphasis>

The embryonic development and the level of dependency of the embryos from the maternal organism were investigated for Centroscymnus coelolepis. During the development, there is a 22–32% loss of organic matter and gains of 95–122% and 29–46% on moisture and inorganic matter, respectively. Only 28–48% of the initial yolk energy is conserved in embryos, being the remaining used in catabolism. In late pregnancy, the epithelium thickness of uterine villi decreases and vascularity increases. Results suggest that only water and minerals are transferred from the mother to the embryos. Monounsaturated fatty acids are the main type of fatty acids representing 48% of the total yolk fatty acid content, followed by polyunsaturated (30%) and saturated fatty acids (18–21%).     

Sex pheromone of <Emphasis Type="Italic">Ectinus aterrimus</Emphasis> (L<Emphasis Type="SmallCaps">inné</Emphasis>, 1761) (Coleoptera: Elateridae)

Summary.   Ectinus aterrimus (L.) is a fairly common European click beetle species which develops mainly in forests. In pheromone gland extracts of female E. aterrimus, examined using gas chromatography – mass spectrometry (GC-MS), one single compound was present. This was identified as 7-methyloctyl 9-methyldecanoate by comparison with a synthetic sample. Field trapping trials revealed a highly significant attraction of male E. aterrimus towards this ester. The structure of the compound differs remarkably from the pheromones of the closely related Agriotes spp., which exclusively use terpene esters.     

Anthranilic acid: a free amino‐acid pheromone in the black chafer, <Emphasis Type="Italic">Holotrichia loochooana loochooana</Emphasis>

Summary. Ether extract of the abdominal gland of female black chafers Holotrichia loochooana loochooana (Sawada) (Coleoptera: Scarabaeidae) induced a series of pre-mating behaviors that included short-distance orientation, mounting and abdominal bending toward females in the laboratory. When the extract was chromatographed on a silica gel column, the activity was eluted with 50% ether in hexane and ether fractions. GC-MS analyses revealed that both the active fractions contained anthranilic acid (2-aminobenzoic acid) as a major compound. When partitioned to basic, acidic and amphoteric fractions, pheromonal activity was observed with the latter two fractions and anthranilic acid was detected in these fractions by HPLC analyses. The amount of anthranilic acid in the female extract was estimated to be ca. 1.3 μg/female. Authentic anthranilic acid induced pre-mating behavior in males of at levels equal to that elicited by the compound in the extract. It also attracted male beetles to the traps in the field. Received 29 October 2002; accepted 9 January 2003. R1D=" Correspondence to:Hiroe Yasui, e-mail:yasui@affrc.go.jp     

Saving by freezing? Metabolic rates of<Emphasis Type="Italic"> Adamussium colbecki</Emphasis> in a latitudinal context

Standard metabolic rates of the endemic Antarctic scallop, Adamussium colbecki (Smith, 1902), were measured in austral summer and under simulated winter conditions. Average mass-specific metabolic rates were significantly different between "summer" (151.17±45.06 µl O ₂ g ^-1 h ^-1) and "winter" (106.52±39.65 µl O ₂ g ^-1 h ^-1) animals. The overall metabolic rates of A. colbecki are comparable to those of other Antarctic bivalve species, but well below those of temperate scallop species. Data for 24 scallop populations (13 species) from different latitudes give no evidence for elevated metabolic rates in A. colbecki as suggested by the concept of "metabolic cold adaptation". A world-wide comparison of metabolic rate and overall growth performance of scallops indicates that in the Antarctic scallop the energetic advantage of low basal metabolism does not counterbalance the disadvantage of the prolonged seasonal period of food shortage.     

Optimal maternal investment in the dung beetle<Emphasis Type="Italic"> Onthophagus taurus</Emphasis>?

Optimal investment theory is based on the assumption that the proximate constraint acting on parental investment is resource based. A trade-off between per offspring investment and total investment seems intuitive. Consequently, a parents investment strategy is expected to represent a trade-off between the benefits of investment for current offspring and the costs to future reproduction for parents. In this study, we provide clear evidence that the costs and benefits of maternal provisioning in the dung beetle Onthophagus taurus influence the amount of provisions provided by the mother. Horse dung is typically of a higher nutritional value than cow dung and females were shown to provide 20% less dung to offspring when provisioning with horse dung. By reducing their investment per offspring and exhibiting a clear preference to provision offspring with horse dung, females were able to produce significantly more offspring. Females provisioning with horse dung received greater fitness returns per unit of investment and experienced lower provisioning costs, in terms of the minimum amount of dung required to produce a surviving offspring, than females provisioning with cow dung. Females provisioning in soil of low moisture content were found to have higher tunneling costs than those provisioning in soil of high moisture content, while the fitness returns per unit of investment did not differ. We adopted a marginal value theorem (MVT) approach to calculate the theoretical optimal level of investment for each dung type and for each soil moisture. Predicted levels of provisioning were lower for horse dung than for cow dung and for moist soil than for dry soil. Therefore, the results of this study are in qualitative agreement with MVT predictions and provide empirical support for the proposal that females can adaptively adjust their level of investment in response to resource and/or habitat quality. However, the theoretically predicted optimal investment yielded a poor quantitative fit with our observed levels of investment, with females providing over twice the investment predicted by the MVT approach. We suggest that this difference may reflect either our inability in directly quantifying all the necessary costs and benefits of investment in O. taurus and/or the applicability of the underlying assumptions of MVT.Communicated by D. Gwynne     

<Emphasis Type="Italic">Hippophae rhamnoides</Emphasis> on a coastal dune system: a thorny issue?

The study mapped the spread of the invasive non-native shrub, Hippophae rhamnoides, on a coastal dune system in South Wales. H. rhamnoides colonies spread across the system, covering around 60.9 ha in 1996 compared to 2.4 ha in 1957. Clearance activities have since decreased the total to around 23 ha. The effects of this expansion on ground flora were assessed through comparison of species assemblages in colonised, never colonised and cleared areas. Dichotomisation presented a number of markedly different communities which correlated to their status of H. rhamnoides invasion. Colonised quadrats were overwhelmingly composed of H. rhamnoides and a few shade-tolerant species such as Galium aperine and Urtica dioica. Cleared quadrats did not support desirable dune grassland species present in never colonised sites, although those which had been cleared for the longest period may be regaining some characteristic species. Clearance work in future needs to concentrate on the complete removal of smaller H. rhamnoides patches whilst also preventing the establishment of Chamerion angustifolium and other ruderal species which in turn inhibit the development of typical dune grassland communities.     

Temporal and vertical distribution of two ecologically different calanoid copepods (<Emphasis Type="Italic">Calanoides carinatu</Emphasis>s Krøyer 1849 and <Emphasis Type="Italic">Lucicutia grandis</Emphasis> Giesbrecht 1895) in the deep waters of the central Arabian Sea

The full-depth vertical distributions of males, females and juveniles of Lucicutia grandis and Calanoides carinatus are presented from the surface down to depths of 3,900 m in the central Arabian Sea at the end of the NE monsoon and the February and October inter-monsoonal periods. L. grandis is a steady component and C. carinatus is a temporal resident of the central Arabian Sea oxygen minimum zone. Maximum abundances of C. carinatus and L. grandis juveniles were found at depths around 600 m, whereas the center of the distribution of L. grandis adults was somewhat deeper at 950 m. Both copepods are indicator species which can help to understand the functioning of the mesopelagic Arabian Sea ecosystem.