Successful sexual reproduction of the scleractinian coral Porites panamensis: Evidence of planktonic larvae and recruitment

Porites panamensis is a hermatypic brooder coral endemic to, and distributed along, the Eastern Tropical Pacific, and is considered a species vulnerable to local effects because it has limited capacity for long‐distance dispersal (and low genetic diversity). Although larvae of P. panamensis have been previously shown to recruit to artificial settlement platforms, they have never been observed in the water column. The present study describes the reproductive behavior of P. panamensis, with a focus on using molecular tools to document evidence for a larval planktonic stage and for successful recruitment. Larvae collected from the water column, and recruitment on natural and artificial substrata were documented. Phylogenetic analysis of two ribosomal markers, 18s rRNA and ITS (ITS1‐5.8‐ITS2), and one mitochondrial marker, cytochrome oxidase subunit 1 (cox1), confirmed the taxonomic identity of larvae, and showed that larvae and recruits have genotypes similar to adults of P. panamensis. Lipid vacuoles and Symbiodinium sp. were present in the gastrodermis of all larvae. A total of 12 and 371 recruits settled on artificial and natural substrates, respectively, and the recruitment rate differed significantly over time. By documenting the reproductive success of the species, we show the potential for existing individuals both to maintain the population in the study area and to contribute to maintenance of the coral reef community in the coming decades.     

ROLE OF RECRUITMENT IN STRUCTURING BEDS OF SARGASSUM SPP. (PHAEOPHYTA) AT ROTTNEST ISLAND,WESTERN AUSTRALIA1

The importance of annual recruitment to the structure of adult stands of Sargassum was determined for a mixed species Sargassum bed at Rottnest Island, Western Australia. The morphologically similar species Sargassum spinuligerum Sonder, S. distichum Sonder, and S. podacanthum Sonder grew together in the shallow subtidal (6 m). Positive species determinations were only possible when thalli were reproductive, so recruits, bases, and vegetative annuals for all species were grouped together. Densities of recruits, perennial bases, vegetative annuals, and reproductive annuals were determined at monthly intervals from 20 randomly placed 0.25-m^?2 quadrats. Recruitment and mortality for recruits and adults were further determined at three monthly intervals from 6-×-1-m^?2 permanent quadrats. The density of adults varied little with season (between 32 and 58 m^?2). Growth of annuals was initiated in April, thalli became reproductive by late August–early September, and senescence occurred in December–January. Density of recruits was highly variable (1.6–210 individuals-m^?2) and peaked seasonally during late summer (January–February) and then declined rapidly. Adults showed a complete turnover of thalli in the bed over 25–27 months. Adult mortality was compensated by annual recruitment from propagules (43%) and vegetative regeneration from fragments of holdfasts left on the reef (57%). A seasonal pattern in survivorship was observed for adults that grew from recruits with higher initial numbers and lower mortalities for August and November cohorts. Little seasonally was observed in survivorship of adults that grew vegetatively from remnant crusts. Although initial cohort sizes were smaller for adults grown from recruits than from remnant crusts, mortality was lower, resulting in similar contributions to adult density from both recruits and remnant crusts. Recruitment from propagules and vegetative regeneration played an important role in buffering the adult stand from high rates of mortality and reducing seasonal variation in adult density and contributed to the persistence and seasonal structure of Sargassum beds at Rottnest Island.     

MICROBIAL COLONIZATION ON NATURAL AND ARTIFICIAL MACROPHYTES IN A PHOSPHORUS-LIMITED,HARDWATER LAKE1

Epiphyte communities in a phosphorus-limited hard-water lake were compared over a 14-week period from Potamogeton illinoensis and structurally similar artificial plants of different leaf ages. Artificial plants were serially incubated in the lake to simulate the age of natural leaves. The physiognomy of loosely attached epiphytes appeared similar on the two substrata. Algal cell number and biovolume were 15-fold and 17-fold higher, respectively, on artificial leaves early in the growing season, but total algal density gradually became similar on natural and artificial plants. In contrast, the taxonomic composition of loosely attached algae became increasingly distinct, and mean cell biovolume on natural leaves was twice that on artificial leaves. Adnate epiphytes on both substrata developed from sparse populations of bacteria on new leaves to a community of diatoms, blue-green algae and numerous bacteria on mature and senescent leaves. Adnate community succession on natural leaves in late senescence/death differed from that on artificial leaves colonized for comparable periods in having (1) a marked increase in filamentous blue-green algae, (2) a subsequent decrease in all algae, and (3) a final fungi-dominated stage. The trends in colonization indicate that macrophytes in this oligotrophic lake provided a distinct habitat from that of artificial substrata for epiphytes throughout the growing season.     

Utilization of holdfast fragments for vegetative propagation of Macrocystis integrifolia in Atacama, Northern Chile

We attempted to propagate Macrocystis integrifolia (giant kelp) by fragmentation and regeneration of holdfasts, which consist of creeping stolons with lateral haptera. Stolons from a natural kelp population in Bahia Chasco (Atacama, North Chile) were cut into fragments, each containing one or more lateral stipe and frond initials. Fragments were attached to longlines with elastic bands. We used two additional types of inoculants as controls: (1) natural recruits from the local parent population and (2) laboratory-cultured young sporophytes. Length, reproductive phenology, and mortality were determined monthly. Our results confirmed the feasibility of M. integrifolia propagation by reattachment of stolon fragments, and we obtained up to eight new individuals from one parent holdfast. Individuals from holdfast regeneration formed sporangial sori 3 months earlier than control specimens from recruits and laboratory culture, while all three types gave similar values for mortality. Holdfast morphology of regenerates differed strongly depending on their origin: stolon fragments continued growth and developed new haptera except in the cut surfaces, natural recruits formed typical M. integrifolia holdfast morphs, and sporophytes originating from laboratory culture produced minor stolons with many haptera. Implications of our results on ecology, repopulation, and aquaculture are discussed.     

Demographic costs and benefits of natural regeneration during tropical forest restoration

For tropical forest restoration to result in long‐term biodiversity gains, native trees must establish self‐sustaining populations in degraded sites. While many have asked how seedling recruitment varies between restoration treatments, the long‐term fate of these recruits remains unknown. We address this research gap by tracking natural recruits of 27 species during the first 7 years of a tropical forest restoration experiment that included both planted and naturally regenerating plots. We used an individual‐based model to estimate the probability that a seedling achieves reproductive maturity after several years of growth and survival. We found an advantage for recruits in naturally regenerating plots, with up to 40% increased probability of reproduction in this treatment, relative to planted plots. The demographic advantage of natural regeneration was highest for mid‐successional species, with relatively minor differences between treatments for early‐successional species. Our research demonstrates the consequences of restoration decision making across the life cycle of tropical tree species.     

Diurnal, horizontal and vertical dispersal of kelp-associated fauna

The kelp Laminaria hyperborea is host for a rich fauna of mobile invertebrates. Dispersal patterns of these animals may be crucial for their availability to visual predators like fish, which are known to search for food in these productive habitats. Diurnal, horizontal and vertical dispersal within and out of the kelp forest were studied by analysing colonization of artificial substrata placed among kelps. The species composition of the fauna was different on three parts of the kelp: lamina, stipes (with epiphytes) and holdfast. The artificial substrata were colonized mainly by species associated with kelp. More species and individuals colonized the artificial substrata at night than during the day. The most abundant faunal groups on the kelps and the artificial substrata were amphipods and gastropods. Both groups dispersed at a significantly higher rate at night than during the day. Rapid horizontal dispersal out of the kelp forest was found. The artificial substrata were also colonized quickly by kelp fauna at all vertical levels inside and above the kelp forest. However, species associated with the kelp holdfast tended to disperse close to the bottom, while stipe-associated fauna moved through all parts of the kelp forest and even above the canopy layer. A high dispersal rate appears common amongst the mobile species living on kelp and seems to be advantageous, despite the risks involved in emigrating from the habitat and being exposed to predators. Higher activity at night may reduce availability to predators.     

SPORE SUPPLY AND HABITAT AVAILABILITY AS SOURCES OF RECRUITMENT LIMITATION IN THE GIANT KELP MACROCYSTIS PYRIFERA (PHAEOPHYCEAE)1

The causes of spatial variation in the recruitment of benthic marine algae are frequently misunderstood because of difficulties in distinguishing among the many factors that influence the supply and establishment of microscopic propagules. We used the recently constructed San Clemente Artificial Reef (SCAR) experiment to examine the roles of dispersal distance, size of spore source, and habitat availability as sources of variation in the recruitment of the giant kelp Macrocystis pyrifera (L.) C. Ag., a species whose recruitment has often been considered to be dispersal limited. Sparse colonization on SCAR by adult Macrocystis occurred within 6 months after reef construction via drifters (i.e. individuals from neighboring kelp beds that became dislodged and set adrift). The abundance of drifters on SCAR declined exponentially with distance from the nearest source population (San Mateo), suggesting that San Mateo was the likely source of drifters. Dense recruitment of small Macrocystis sporophytes was observed within 8 months of reef construction. The density of recruits on SCAR showed an initial increase with distance from San Mateo before declining exponentially. Nonetheless, substantial recruitment was observed at the most distant locations on SCAR located 3.5 km from San Mateo. In contrast to drifters, the density of recruits was positively correlated to the bottom cover of artificial reef substrate. Importantly, no correlation was found between the local density or fecundity of drifters and the local density of kelp recruits suggesting that recruitment on SCAR resulted from widespread spore dispersal rather than from the local dispersal of spores from drifters.     

Impact of clonal growth on effective population size in Hymenoxys herbacea (Asteraceae)

By influencing the proliferation of different genotypes, clonal growth can affect the maintenance of genetic variability and magnitude of genetic drift within plant populations. However, estimates of effective population size rarely incorporate the contribution of both asexual and sexual reproduction. We estimated effective size (Ne) for two populations of the clonal, self-incompatible plant, Hymenoxys herbacea, using a stage-structured demographic model for organisms with asexual and sexual recruitment and then examined the impact of reproductive strategy using an elasticity analysis. Plant rosettes monitored in two successive years had high survival rates in both populations (mean 0.94). The mean number of sexually derived recruits per initial ramet was 0.041 (SE 0.039), whereas the mean number of clonal recruits was 0.61 (SE 0.90). Effective size was 1642 and 5769 in the two populations and the Ne/N ratio averaged 0.34, comparable to values for other clonal species. Elasticity analysis indicated that increases in both clonal and sexual recruitment cause an increase in Ne while increasing the variance reduced Ne. However, Ne was more sensitive to changes in the mean and variance of asexual recruitment than sexual recruitment. These results highlight the importance of considering asexual modes of reproduction when examining the role of genetic stochasticity in populations.     

Recovery of Triatoma infestans populations after insecticide application: an experimental field study

The capacity of populations of Triatoma infestans (Klug) (Hemiptera: Reduviidae) to survive and recover was assessed after application of insecticide (gamma-HCH at a rate of 0.5 g a.i./m2) at different seasons. T. infestans populations were maintained in experimental chicken houses under natural climatic conditions in a region of Argentina endemic for Chagas disease transmitted by these bugs. Based on previous studies of T. infestans populations in these habitats, each experimental group was set up with a total of 626 T. infestans, comprising 390 eggs, 204 nymphs of particular stages, fourteen male and eighteen female adults. The chicken houses were dismantled and rebuilt at monthly intervals to study the vector population changes over a period of 33 months. When the insecticide was applied during winter, spring or summer, populations of T. infestans recovered to untreated or precontrol levels during the next reproductive season (i.e. during the hot season, October-March). In contrast, populations treated during autumn (March) remained at very low densities for 2 years and then increased rapidly to surpass the untreated populations. All populations of the bugs fell to very low numbers (sometimes less than twenty individuals) after gamma-HCH applications, but none was driven to extinction. In all cases, the density of surviving populations was independent of their density before treatment. The fact that all treated populations recovered within 1-3 years, to at least the density of untreated populations, shows the high reproductive potential of T. infestans to recover from very low population densities. Moreover, the additive effect of climatic-induced mortality and insecticide-induced mortality is only apparent when insecticides are applied just before the onset of the cold winter months during which reproductive rates are at their lowest.     

The timing of breeding and offspring size in Great Tits Parus major

The relationship between the timing of breeding, offspring weight and offspring recruitment was investigated in two populations of Great Tits Parus major. In two of six seasons a significant negative correlation was found between laying date and fledgling weight, in three seasons the opposite was true and in one season no significant correlation existed between these variables. Offspring recruitment was quantified in five seasons, and late-nesting birds were found to produce significantly fewer recruiting offspring in two of these, the same two seasons when late fledglings had significantly lower weights. No significant correlation existed between the number of recruits and laying date in the three seasons when late fledglings were as heavy as or heavier than early fledglings. The potential detriment to reproductive success of lateness thus was offset when late-nesting parents managed to produce heavy young. Between-year variations in the seasonal pattern of reproductive success has previously been suggested to contribute to the maintenance of variability in the timing of breeding in Great Tits. In this paper we suggest that producing heavy offspring late in the season might be a proximate factor influencing the seasonal patterns of reproductive success. Producing heavy offspring late in the season is probably not a "strategy" but rather an effect of chance differences between years in the seasonal availability of food or the seasonality of other factors important for the nesting birds.     

Density- and size-dependent mortality of a settling coral-reef damselfish (<Emphasis Type="Italic">Pomacentrus moluccensis</Emphasis> Bleeker)

Density-dependent mortality may regulate many populations, but due to an offshore larval phase in benthic marine organisms, it is often difficult to quantify the effects of mortality of arriving individuals. We added approximately 600 recruit-sized individuals onto coral reef that parallels patterns in naturally settling fish. Strong, positive density-dependent mortality occurred 3 days, 1–2 weeks, and 4 months after release. Since our study species was patchily distributed, we estimated both mean group size and overall density in a transect. Mortality was more strongly related to mean group size than overall density in a transect, indicating that recruit patchiness was important. Cohesive groups may suffer higher mortality than those spread over larger areas, even if overall density of the latter is greater. Aggregative responses of predators may occur in response to larger groups, so may have contributed to positive density-dependent mortality. Increased conspicuousness to predators and congeneric aggression are additional factors that may vary positively with group size. Tagging of recruits showed migration within but not between transects, so persistence was tantamount to survival. Standard otolith back-calculation techniques employed to reconstruct original size of tagged recruits that persisted 4 months after additions indicated that mortality was also size-dependent. Size-dependent mortality was apparent at the site with the highest mortality but not at the site with the lowest mortality, resulting in different mean body sizes of recruits between sites. Size-dependent mortality may influence estimation of growth and lead to onset of size-based maturity in these fish. Strong, positive density-dependent mortality may regulate recruitment, and if coupled with size-dependent mortality, may increase maturity rate, adding to reproductive schedules of those that survive.     

Effect of the substratum in the recruitment and survival of the introduced barnacle Balanus glandula (Darwin 1854) in Patagonia, Argentina

The barnacle Balanus glandula was introduced in Argentina in the 1970s, and today it dominates the high intertidal level in most Argentinean rocky shores. The aim of this work was to evaluate the effect of the type of substrata and intertidal height on a population of Balanus glandula by conducting field surveys and one-year field experiments in which we combined different substrata (hardness: hard and soft, and texture: smooth and rough) at two intertidal heights (mid and high). In natural populations, the highest density of adults and recruits occurred on soft-rough substratum and in the high intertidal. The different textures were important only on the soft substrata and high intertidal, and the density of barnacles of the soft-rough substrata was higher than soft and smooth ones. The most suitable experimental substratum was the soft-rough of the high intertidal, which had the highest recruitment, survival and final density of barnacles at the end of the experiment. In contrast, the hard and smooth of the high and middle intertidal were the least suitable in all cases. Although the recruitment of B. glandula occurred throughout the year, it was higher in the high intertidal, and it showed a recruitment peak in the winter and a second in the summer. While most studies on this barnacle investigated the effects of granite or other volcanic hard substrata, our study also focused on soft substrata. The effects of soft substrata are particularly important because soft sedimentary rocks characterise the southern Atlantic coast of South America and the presence of soft rocks appears to optimize the success of Balanus glandula.     

Enhancement of Grazing Gastropod Populations as a Coral Reef Restoration Tool: Predation Effects and Related Applied Implications

We aimed to evaluate the efficacy of the gastropod grazer Trochus niloticus in controlling epilithic algae and enhancing coral recruitment on artificial substrata on coral reefs where the biomass of herbivorous fishes was low due to heavy fishing pressure. Hatchery‐reared, subadult trochus were stocked onto pallet balls (small artificial reefs composed of concrete and limestone aggregate) at a density of approximately four individuals per square meter (external surface area). This density was re‐established with releases of new trochus each month for 6 months. At the end of the experiment, there were no significant differences in algal biomass, cover and community composition, or the density of coral recruits on substrata with and without trochus. High monthly attrition of stocked trochus on the pallet balls, apparently due mainly to predation by octopus, did not allow the evaluation of the efficiency of the trochus enhancement, at the desired density, as a restoration tool. However, at the lower trochus densities (circa 1 m^?2), which occurred as a result of predation in this study, no apparent enhancement of algal grazing or coral recruitment were observed. The surprisingly high predation of stocked trochus in a heavily fished and gleaned reef site stresses the importance of understanding all the factors affecting the survival of stocked animals. To help mitigate predation of trochus, artificial habitat with refuge spaces that allow the grazers to escape predation could be provided and individuals of a larger size could be released.     

Variation in barnacle recruitment over small scales: larval predation by adults and maintenance of community pattern

Over small spatial scales, variation in the density of settlers of benthic sessile species is the result of interactions among larval behavior, local hydrodynamic conditions, and the physical, chemical and biological characteristics of the benthic habitat. It has been shown repeatedly that adult benthic filter-feeders can consume larvae of their own and other species, but their effects on the distribution and abundance of recruits have rarely been demonstrated under natural conditions in the field, particularly on hard substrata. Here we experimentally quantified the effect of the large intertidal barnacle, Semibalanus cariosus (Pallas), on the density of recruits of three common barnacle species. The experiments were conducted at the peak of the barnacle recruitment season over three successive years, on the west coast of San Juan Island, Washington. A persistent and well documented community pattern in the mid intertidal zone of the study site is a sparse bed of adult S. cariosus with bare rock spaces essentially devoid of small barnacles among the large individuals. Field experiments consisted of small areas from which either all adult S. cariosus were killed leaving the shells attached to the rock, or live adult barnacles were left intact. Our results showed that over small spatial scales of a few to tens of centimeters, the large barnacle S. cariosus can interfere and significantly reduce net settlement and recruitment of conspecific as well as other barnacle species. Between 65 and 100% reduction in settlement could be attributed to larval predation by adults, as implied by barnacle settlement patterns on different treatments and by the presence of nauplius larvae in cirri and stomach contents of S. cariosus. The negative effect on barnacle settlement was consistent between years of relatively low barnacle recruitment, which appears to be the most common situation at the study site, but it disappeared on a year of unusually high recruitment, when settling larvae seem to have swamped the filtration ability of adult S. cariosus. The different barnacle species displayed contrasting settlement patterns on bare rock and on the lateral shells of the large barnacles, which appear to be a result of differences in larval behavior. Comparisons against the relative availability of these substrata in the experimental plots suggested that larvae of different species sample the benthic microhabitat in very different ways.     

Natural settlement dynamics of a young population of Turbinaria ornata and phenological comparisons with older populations

Our study was done during 16 months on two sites at Moruroa to (i) compare Moruroan populations (young) with Tahitian populations (old) of Turbinaria ornata and (ii) follow the natural dynamic of a young population at Moruroa. Moruroan and Tahitian populations have maximal density and maximal reproductive characteristics during the cool and hot seasons, respectively. Recruits are present throughout the year at all sites. Within the reference field site, individuals were initially distributed regularly and then became clumped through time, recruits appeared around adults. Recruits mortality followed a similar pattern, whereas mortality of adults and juveniles was even throughout the monitoring period. Populations initially had similar proportions of recruits, juveniles and adults (28, 40 and 32%, respectively), but at the end, recruits represent 92% of the population. A linear relationship between the number of individuals present within the station and the colonized substrate area was determined (R² = 0.96), suggesting a limiting effect of the substrate against the settlement of T. ornata with time. Moreover, the growth of recruits in the close vicinity of adults was arrested in the young populations: recruitment may therefore, significantly buffer high rates of adult mortality. It may contribute to the invasion and, more interesting to the persistence of T. ornata in new colonized area.     

Relative contributions of sexual and asexual regeneration strategies in Populus nigra and Salix alba during the first years of establishment on a braided gravel bed river

Populus nigra L. var betulifolia and Salix alba L. var alba are early successional riparian tree species threatened throughout Continental Europe by significant changes to the natural physical processes governing their natural habitat – geomorphologically active floodplains. River management activities have dramatically altered natural patterns of river flow and rates of sediment delivery along rivers, with possible consequences for the balance between sexual and asexual regeneration strategies in these species. Conservation strategies will benefit from a greater understanding of the ways in which dynamic physical processes on the floodplain influence sexual and asexual recruitment. This paper describes a field survey investigating the relative abundance and spatial distributions of P. nigra and S. alba sexual and asexual recruits during the first years of establishment along a braided gravel bed river. Sexual and asexual recruits were identified by excavation along transects in a wet and a dry field season and distributional differences were described in terms of elevation on the floodplain, local sediment type and exposure to floodwaters. Regeneration was overwhelmingly from seed in the first 2–3 years following recruitment, but poor survival rates among sexual recruits saw a shift in the relative abundance of regeneration strategies over time. In relating hydrological data to recruitment, unseasonal flood disturbances had a negative effect on recruitment from seed and a positive effect on vegetative regeneration. Seedlings were associated with fine sediment deposits and were restricted primarily to low elevations on the flood plain, while asexual recruits had a wider spatial distribution. Certain microsite types were unique to either regeneration strategy.     

Recruitment density can determine adult morphology and fecundity in the barnacle, Semibalanus balanoides

Although consequences of the settlement preference of larvae have been well documented, the consequence of these settlement choices on subsequent mortality, morphology and fecundity has been little studied. The aim of this study was to determine the relationship between recruit and adult density and to determine the effect of recruitment on adult morphology and egg tissue mass. This study follows 48,718 barnacles (Semibalanus balanoides) from recruitment at the end of the settling season to reproductively mature adults at a field site in the Clyde Sea (UK). Overall survivorship of the recruits to adulthood was 8.5%, although survivorship was up to 42% on low density settlement panels. In low density colonies (< 10 recruits cm-2), recruitment density was related to adult density (P < 0.001), whereas no relationship was found for higher density colonies. A shell morphology index measured at adulthood was related to recruitment density for low density recruited colonies (P < 0.001) but not high density colonies. Using ANCOVA, variations between the colonies in shell and egg tissue mass were not explained by mass of somatic tissue. However, egg mass was explained by recruitment density (P < 0.01). These results show that adult density is not a reliable indicator of the previous population density of the colony. Moreover, there are marked differences in population development between colonies with high and low recruit densities in terms of impact upon shell morphology and egg production. The dynamics that operate between recruits at the end of the settlement season and sexually mature adults to create the patterns elucidated in this paper, and other literature, remain unclear.     

COMPARISON OF GROWTH AND REPRODUCTION BETWEEN TWO POPULATIONS OF SARGASSUM SILIQUASTRUM IN PING CHAU, HONG KONG

In Ping Chau Island of Hong Kong, two populations of Sargassum siliquastrum are present. One is in the shallow water of about 1 to 3 m Chart Datum (CD), and the other one in deeper water of 5 to 10 m CD. These two populations are separated by an extensive sand patch. Individuals of the "shallow" water population increased their size from a mean length of 8.3 ± 3.6 (SD) cm in Aug 1998 to a maximum of 48.2 ± 29.9 cm in early Jan 1999 before they started to die back. In the following year, they attained a minimum of 6.1 ± 3.8 cm in May 1999 and a maximum of 56.1 ± 23.6 cm in Dec 1999. Their reproductive period lasted for two to three months from Jan to Feb 1999, and again from Nov 1999 to Jan 2000. The "deep" water individuals increased their size to a maximum of 123 ± 50.8 cm at the end of Jan 1999 and started to die back in Feb, 1999. They again reached their maximum mean length in Jan 2000. Their reproductive period lasted for five to six months from Sept 1998 to Feb 1999 and again from Sept 1999 to Jan 2000. The "deep" water individuals tended to be longer in size and they attained their maximum growth a month later than the "shallow" water individuals. Their reproductive season tended to start earlier and lasted longer than those in the "shallow" water. These differences in the phenology of the two populations may be related to the temperature differences (up to 5° C difference in summer) between the two depths. Sargassum siliquastrum is likely to be a cold adapted species such that warmer temperature in the shallow water has compressed and shortened their rapid growth and reproductive period to within the few colder months in fall and winter.     

Spawning aggregations of squid (Sepioteuthis australis) populations: a continuum of ‘microcohorts’

The aim of this study was to determine how size, age, somatic and reproductive condition, abundance and egg production of southern calamary spawning aggregations changed during the spawning season in each of 2 years. During the spawning period in at least one of the years there was a decline as much as 20% in average size, 50% in somatic condition, 28–34% in size-at-age, 26–29% in reproductive status, as well as abundance and reproductive output of the stock declining during the spawning season. However, this change was not a function of the population becoming reproductively exhausted, as the aggregation was composed of different individuals with different biological characteristics. In each month the average age of individuals was ca. 6 mo, indicating that squid that had hatched at different times had entered the spawning aggregations, suggesting that the aggregation was made-up of a succession of microcohorts. Currently, management of many squid populations assumes that there is a single cohort in the aggregation. Therefore, estimating stock biomass at the start of the spawning season cannot be used as the population is constantly changing as micro-cohorts move into the aggregation. An instantaneous estimate of the spawning biomass, independent of fishing activity may be obtained by quantifying the density of deposited eggs. The strategy of individuals with a diversity of life history characteristics coming together in a single spawning aggregation may ensure the phenotypic and genetic diversity required to guarantee successful recruitment of this short-lived species. Therefore, temporally structured protection from harvest throughout the spawning season will ensure maintenance of this population diversity.     

Pollination success in monochromic yellow populations of the rewardless orchid Dactylorhiza sambucina

Dactylorhiza sambucina is a European terrestrial orchid that lacks a pollinator reward. Throughout most of its range, populations contain yellow- and purple-flowering individuals, but in western Germany, monomorphic yellow populations predominate. As elsewhere, bumblebee queens are the most important pollinators in these populations, and mean fruit set over two years was 19%, well within the range reported from dimorphic populations. Multivariate analyses of plant and population traits, including plant height, leaf number, flower number and density on the spikes, flowering population density, and nearest neighbor distance, showed that only individual plant height and population density had a unique positive effect on pollen export; female function was unrelated to height or population density. The positive effects of dense spacing of flowering conspecifics and tall size appear due to greater visual attractiveness. Good visual exposure may also explain that flowers higher up on the spikes, in spite of opening late in the season, had higher male reproductive success than early flowers.