Facilitation and competition between invasive and indigenous mussels over a gradient of physical stress

The interactions between invasive exotic and indigenous species can have profound harmful effects on the recipient community; however, not all such interactions are negative. Facilitation is increasingly recognised as important in shaping natural communities and is believed to vary under different conditions. Earlier studies have shown that the indigenous intertidal mussel Perna perna initially facilitates survival of the invasive Mytilus galloprovincialis in the low mussel zone by providing protection against waves, but later excludes M. galloprovincialis through interference competition for space. Here, we examined interactions between these species in the mid and upper mussel zones, moving mussels to experimental plots in different combinations of densities and species. Mussels were left on the shore for more than a year and treatment effects on mortality, shell length and condition were compared. In the high zone, treatment had no effects and P. perna showed greater mortality than M. galloprovincialis, indicating that its exclusion from the high shore is due to emersion stress. In the mid zone, treatment had no significant effects on M. galloprovincialis, but multiple comparisons among treatments involving P. perna showed that facilitation occurred. P. perna survived better at higher densities, but survived even better when mixed with the physiologically more tolerant M. galloprovincialis. Length data indicated both inter- and intraspecific competition for P. perna in the mid zone. Whereas facilitation occurs strongly in the low zone (P. perna facilitates M. galloprovincialis) and weakly in the mid zone (M. galloprovincialis facilitates P. perna), the lack of facilitation in the high zone suggests that the probability of facilitation is not linearly linked to increasing physical stress. Instead it is likely to be hump shaped: relatively unimportant under conditions that are benign for a particular species, significant under more severe conditions, and overridden by physical stress under very harsh conditions.     

Effect of resource gradient on age and size at maturity and their influence on early‐life fecundity in the predatory Asian lady beetle,Harmonia axyridis

Variation in food availability impacts the performance of insects in terms of their size and age to maturity and fecundity. Age at maturity determines how quickly individuals in a population can start to reproduce and how much they can reproduce. Results from studies on various insect species show that food availability influences the size and fecundity of adult females. It is predicted that under poor growth conditions, variation in size is low, but variation in age at maturity is considerable. This prediction was examined in a widely distributed lady beetle species, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), a predator of aphids and coccids. Using a food gradient from low to high aphid prey density, performance of females that were reared on excess food was recorded for pre‐reproductive duration, size at reproductive maturity, number of aphids consumed, and fecundity in the first 10 days of their reproductive period. Results suggested that female H. axyridis that were reared on surplus food when kept at low prey density (poor growth condition) took, on average, three times longer to attain maturity and produced, on average, 14 times fewer eggs than females that were also reared on surplus food, but kept at high prey density (good growth condition). Females performed best at a prey density of 30 aphids per female per 150 cm². Results suggested that the current food availability significantly influenced the age and size of females at maturity and their fecundity. Age and size at maturity of female lady beetles showed non‐linear responses to prey density as well as the occurrence of a minimum size of females, below which H. axyridis females fail to mature. The steep slope recorded at lower prey densities suggests relatively high variation in age at maturity but low variation in size.     

Successive changes in distribution patterns as an adaptive strategy in the bivalveMacoma balthica (L.) in the Wadden sea

During their first year in the Wadden Sea, high proportions of the tidal-flat populations of the tellinid bivalveMacoma balthica (L.) redistribute twice: immediately after their settlement in late spring, the postlarvae show a net transportation in a shoreward direction — whereas in the subsequent winter, the grown spat move in the reverse direction. As a consequence of these two periods of high mobility, distribution patterns shift twice: though initial settlement takes place mainly in the lower half of the intertidal, most spat grow to a size of ∼0.5 cm in the upper half of it, whereas most adults live in the middle and lower zones. The successive distribution patterns of spat and olderMacoma are described in detail for Balgzand, an extensive tidal-flat area in the westernmost part of the Wadden Sea. Long-term observations in this area and published evidence from other areas are used to evaluate the suitability of different tidal zones as a habitat for successive life stages ofMacoma. For spat, the upper zone is a more favourable habitat than the lower, because predation pressure (mainly from shrimp) and disturbance are less and growth is more rapid. For adults, the upper parts are no longer a favourable habitat, because the few animals that stay suffer from high parasite load, low survival, slow growth and low reproductive output. It is concluded that in their successive life stages the majority ofM. balthica live at the intertidal level most favourable to them. The strategy of timely shifts to areas more suitable to the next life stage contributes to the success of the species: it is the most widespread and common (and one of the most stable) macrozoobenthic species in the Wadden Sea.     

Seasonal migration promoting assortative mating in Littorina brevicula on a boulder shore in Japan

Littorina brevicula Philippi is one of the most common snails found in the upper intertidal zone of Japan. In Amakusa, some of the population of L. brevicula migrate to the lower zone in the winter, while the rest stay in the upper zone. Thus, during the winter, which is its reproductive season, the population of L. brevicula divides into two sub-populations. This leads to a hypothesis that the migration pattern in winter is genetically controlled and this behavioural dimorphism is maintained by reproductive isolation between the two sub-populations. In order to test this hypothesis, the following three points were investigated: (1) whether the same snails migrate in a similar way every winter, (2) whether there is a significant tidal level preference in snails, and (3) whether reproductive isolation occurs between the two sub-populations. The results showed (1) the migration behaviour of each snail was consistent over two successive winters, i.e. the same group of snails migrated downward every winter and the same group of snails stayed in the upper zone every winter, (2) transplanted snails moved toward the original zones where they were caught, suggesting that the snails actively selected their tidal zone in winter, and (3) most of the snails copulated within each sub-population. Therefore, reproductive isolation between the two sub-populations was considered to be established to some extent by the dimorphic migration behaviour. In conclusion, the migratory behaviour of L. brevicula is determined separately for each individual and might be genetically controlled, and the behavioural dimorphism may be maintained by partial reproductive isolation between the two sub-populations.     

Clinal genetic variation and the 'rare allele phenomenon' in random mating populations of <Emphasis Type="Italic">Urophora cardui</Emphasis> (Diptera: Tephritidae)

In the present study we investigate a contact zone between two population groups of the tephritid fly Urophora cardui. We investigate scenarios that may have produced the genetic differentiation of the two groups, and we describe the 'rare allele phenomenon' from the contact zone. The rare allele phenomenon refers to alleles that are found at high frequency in contact zones but are rare or lacking outside the contact zone. The phenomenon is often observed in hybrid zones between subspecies of limited reproductive compatibility, but seldom in populations with random mating. Clinal genetic variation was observed at three loci in the contact zone. Three alleles at the locus Aat showed steep clines, between 20--70 km wide. A rare Aat-A allele occurred at high frequency in the centre of the contact zone. Two further loci, Hk and Pgd, showed less steep clinal genetic variation, the transition being in and slightly south of the centre of the Aat cline. Populations showed Hardy--Weinberg proportions and there was no evidence for linkage dis-equlibrium. These findings suggest random mating and gradual introgression between the population systems, which may originate from at least two range expansions. Aat's steep clines and rare allele may indicate selection on Aat alleles, although we presently can not quantify any agents. Because U. cardui experiences random mating in the contact zone with no apparent 'hybrid' incompatibility, mating experiments offer the possibility for future enquiries about the genetic basis of the rare allele phenomenon.     

Maintenance of zonation patterns in two species of flat periwinkle,Littorina obtusata and L. mariae

The zonation patterns of Littorina obtusata (L.) and Littorina mariae Sacchi et Rastelli were shown to be quite distinct on a sheltered rocky shore. L. obtusata was found at all the heights sampled; it reached peak numbers at mid shore on the alga Ascophyllum nodosum L. (Le Jol). There was no difference in the tidal height occupied by adults or juveniles; or in the mean size of L. obtusata along the vertical gradient of the shore. In contrast L. mariae occurred exclusively low on the shore, on Fucus serratus L. Translocation of the two species within their respective levels resulted in random movement after 4 days, although initial movements after 1 and 2 days were sometimes directional. Animals transplanted to the normal level of the other species showed directional movement towards their home zone; this was most pronounced after 4 days. There was no difference in the distance moved by the two species, although the distance moved did vary with tidal height, both species moving further at mid shore than low shore. Distances moved by littorinids at replicate areas in the low shore were similar but those at mid shore did vary. There was an interaction between the species and the different tidal heights which revealed that transplanted species moved further than translocated species at the same tidal level. However, this was only significant in the case of L. mariae. It is suggested that the close relationship between the winkles and their host algae may direct the homing behaviour of displaced individuals.     

The components of life-history traits variation in Daphnia magna population

The variation of six growth and reproduction traits in the parthenogenetic clones of Daphnia magna and its relationships with the genotypes at 5 isozyme loci were studied under controlled conditions at three food concentrations. A significant genetic component of age at maturity, growth rate and fecundity variation was detected. Nested ANOVA revealed the influences of the genotypes of the isozyme loci, that are more pronounced at the high food concentration. In some cases the genotype-environment interactions were found.The variation of body length at 6 days and total number of offspring includes a significant heterozygosity — environment interaction. The most heterozygous clones show the least differences in life history traits between food conditions.Life-history traits heritability, genotype-environment interactions and different reproductive strategies of the clones are discussed.     

Population stability in relation to resource availability in an introduced population of an herbivorous lady beetle

Mechanisms responsible for population stability in relation to resource availability were studied in an introduced herbivorous lady beetle,Epilachna niponica. The introduced population was relatively constant over a seven-year study period. Egg density was related to the variation in host-plant abundance in different years, and was highly stabilized during the period from reproductive adult to egg stage. Two density-dependent processes were identified in the reproductive season: (1) density-dependent reduction in fecundity and (2) density-dependent increase in female mortality and/or emigration, all of which operated early in the season. As a result, temporal variability in cumulative egg density was greatly reduced by mid-May, by which time approximately40% of total eggs were laid. A field cage experiment demonstrated that egg-laying of individual females was largely limited by resource availability even at low levels of leaf herbivory. Since movement activity of ovipositing females increased in a density-dependent manner, inter-plant movement is more likely to cause density-dependent reduction in fecundity and female loss, due to enhanced energy expendiditure. The introduced population was less stable than the source population, probably because of decreased inter-plant movement of females and the unlikelihood of egg resorption, both of which contribute significantly to the temporal stability ofE. niponica population densities.     

醉马草水浸液对豌豆蚜触杀活性及种群增长的影响

为探讨醉马草水浸液对豌豆蚜触杀活性及种群增长的影响,采用带虫浸叶法比较不同生育期醉马草带菌（E+）和不带菌（E-）水浸液对豌豆蚜触杀活性及种群生命表,测定了豌豆蚜的死亡率及触杀后对其生殖期,平均繁殖力,繁殖率及生命表参数的影响。结果表明,不同生育期醉马草带菌（E+）水浸液触杀豌豆蚜后对其各项指标均有显著影响。在苗期时,E+水浸液触杀豌豆蚜后校正死亡率最高,繁殖力最低,内禀增长率（r_m=0.145 d^-1）和净生殖率（R₀=4.802头）均为最小值。在成熟期时,E+水浸液触杀豌豆蚜24 h、48 h和72 h后校正死亡率分别为26.15%,19.01%,9.07%;繁殖期（3.87 d）,平均繁殖力（8.80头）,繁殖率（1.40%）,内禀增长率（r_m=0.208 d^-1）和净生殖率（R₀=8.820头）。在枯黄期时,E+水浸液触杀豌豆蚜后校正死亡率最低,繁殖力最强,内禀增长率（r_m=0.247 d^-1）和净生殖率（R₀=13.647头）均为最大值。不同生育期醉马草E-水浸液触杀豌豆蚜后对其种群繁殖无显著影响,与对照差异不显著（P>0.05）。综上,苗期醉马草E+水浸液对豌豆蚜有较好的触杀效果,校正死亡率高,且触杀后当代繁殖力减弱,种群扩建时间延长,不利于其种群繁殖和增长;故苗期醉马草E+水浸液具有很好的杀虫潜力,所采用水浸液方法制备简单,成本低,可为新型植物源农药研发提供重要理论依据。     

Distribution and abundance of co-occurring chthamalid barnacles Chthamalus montagui and Chthamalus stellatus (Crustacea,Cirripedia) on the southwest coast of Portugal

Two co-occurring chthamalid barnacle species occur on the rocky shores of continental Portugal: Chthamalus stellatus (Poli) and Chthamalus montaguiSouthward. In the present study, patterns of distribution and abundance (density and percentage cover), both vertically (tidal level) and horizontally (wave action), were studied for each species on headlands (more wave action) and not on headlands (less wave action) and at two spatial scales (shore and site) and three tidal levels (low, mid and upper). Chthamalus montagui was significantly more abundant than C. stellatus in both degrees of wave action and at all tidal levels. Despite this great difference in abundance and considering each species individually, C. montagui was more abundant at mid tide-level, while C. stellatus was more abundant at the low tide-level, and C. montagui was less abundant on headlands, while C. stellatus was more abundant on headlands. Small-scale variability (between sites, tens of meters) of abundance of each species was detected, while differences on a larger scale (between shores, tens of kilometres) were not significant.     

Biological Control ofBemisia argentifolii(Homoptera: Aleyrodidae) on Poinsettia with Inundative Releases ofEncarsia formosa(Hymenoptera: Aphelinidae): Are Higher Release Rates Necessarily Better?

The effectiveness of inundative releases of the parasitoidEncarsia formosafor control ofBemisia argentifoliion poinsettia was determined in replicated experimental greenhouses. We evaluated two release rates ofE. formosa:a low release rate (1 wasp/plant/week, released in two greenhouses, in spring 1995) and a high release rate (3 wasps/plant/week, released in two greenhouses, in fall 1993), each over a 14-week growing season. Each release trial had one or two control greenhouses in whichB. argentifoliideveloped on poinsettia in the absence ofE. formosa.Life-tables were constructed forB. argentifoliiin the presence and absence ofE. formosaby using a photographic technique to follow cohorts of whiteflies on poinsettia leaves. Weekly population counts of the whitefly were also made. In the absence ofE. formosa,egg to adult survivorship ofB. argentifoliion poinsettia was 75–81%. At the low release rate, egg to survivorship ofB. argentifoliiwas 5% and parasitism was 13%. At the high release rate, egg to adult survivorship forB. argentifoliiwas 8% and parasitism was 23%. The net reproductive rates (R₀) forB. argentifoliipopulations in the absence ofE. formosaranged from 18.01–26.12, indicating a rapidly increasing population. Net reproductive rates for whitefly populations subject to wasp releases were 1.54 for the low release rate greenhouses and 2.11 for the high release rate greenhouses, indicating substantially reducedB. argentifoliipopulation growth. The low release rate provided better control ofB. argentifoliithan the high release rate. This difference was attributed to higher levels of mortality of whiteflies at the low release rate in the first 5–6 weeks of the growing period. We suggest that mutual interference may also have affected observed levels of mortality and parasitism.     

Variability in the reproductive effort of the limpet,Cellana tramoserica

Summary The reproductive effort of the limpet, Cellana tramoserica was investigated in high, mid, and low intertidal areas of the rocky shore as well as in a subtidal population. The reproductive effort of individuals within a population was independent of age. There was, however, a noticeable difference among populations with respect to the relative amount of gonad material spawned per year. The individuals from the subtidal population had the greatest reproductive effort, spawning the equivalent of 101% of their somatic weight per year, while the low, mid and high intertidal populations spawned the equivalent of 66%, 20%, 51%, respectively.Experimental transplants showed that individuals from all three intertidal populations increased their reproductive effort to match that of subtidal individuals. It was unlikely, therefore, that the observed patterns were the result of genetic differences among the populations. The reproductive effort of the populations was also not correlated with the rate of mortality of the adults in the four populations. The observed differences could be explained by a combination of the availability of food and intraspecific competition. The flexibility in the reproductive pattern of Cellana is hypothesised to be a result of this species having a dispersive larval stage, which may result in selection for plasticity of reproductive allocation, depending upon the environmental conditions at the site of settlement and growth to adult size.     

HABITAT DIFFERENCES IN THE TIMING OF REPRODUCTION OF THE INVASIVE ALGA SARGASSUM MUTICUM (PHAEOPHYTA,SARGASSACEAE) OVER TIDAL AND LUNAR CYCLES1

Sargassum muticum (Yendo) Fensholt is an invasive species that is firmly established on intertidal and subtidal rocky shores of Europe and the Pacific coast of North America. Local success and spread of S. muticum is thought to rely on its reproductive potential that seems dependent on exogenous factors like tidal and lunar cycles. This study is the first to compare the reproductive patterns (periodicity of egg expulsion and embryo settlement) of this invader in two different habitats: the middle and low intertidal. The combination of monthly, daily, and tidal samples at triplicate sites within each habitat showed a semilunar periodicity of egg expulsion and embryo settlement coincident with increasing tidal amplitude just before full and new moons. In both habitats, duration of each egg expulsion event was ～1 week, and embryo settlement occurred during the first daily low tide and with the incoming high tide during spring tides. However, both expulsion and settlement started 1–2 d earlier, expulsion saturation was faster, and settlement was higher in the mid‐ compared to the low intertidal. Our results suggest that the exact timing of gamete expulsion and embryo release of S. muticum responds to local factors, including tidal cues, which result in differences between mid‐ and low‐intertidal habitats.     

Diseases and reproductive success in a wild mammal: example in the alpine chamois

Density-dependent and climatic factors affect reproduction and dynamics of wild ungulates. Parasites can also decrease reproductive success through either a direct abortive effect or a negative impact on host growth and body condition. However, few studies have investigated the effect of parasitism on fecundity of ungulates in natural conditions. We studied three bacterial infections caused by Salmonella enterica serovar Abortusovis, Chlamydophila abortus and Coxiella burnetii. These bacteria are leading causes of reproductive failure in sheep, goat and cattle, which raises the question of their influence on population dynamics of wild ungulates. A long-term study of demography and epidemiology of an alpine chamois (Rupicapra rupicapra, L.) population (Les Bauges Reserve, France) and a generalized linear modeling approach were used to analyze the reproductive success of chamois according to population density, weather conditions and the prevalence of antibodies against the three bacteria in females. This approach enabled us to identify the confounding effect of weather and parasitism on fecundity in a natural population. After accounting for density, the prevalence of antibodies against the three bacteria explained 36% of the annual variation in reproductive success, and weather conditions explained an additional 31%. This study was, to our knowledge, the first to compare the decrease in fecundity due to bacterial infections and weather conditions in a population of wild mountain ungulates.     

胶州湾西北部潮滩湿地大型底栖动物功能群

2009年2、5、8和11月进行了7个断面35个站位的大型底栖动物调查,选取高潮区(A)、中潮区(B、C、D)和低潮区(E)研究了胶州湾西北部潮滩湿地大型底栖动物功能群组成及其时空变化.调查共发现大型底栖动物71种,主要种类为软体动物(31种)、环节动物(20种)和节肢动物(14种).潮区A、B、C、D、E物种数分别为26、33、35、38、31.依据食性将主要底栖动物划分为肉食者、浮游生物食者、碎屑食者和杂食者4个功能群.各功能群物种数占总物种数的百分比由高到低依次是肉食者、浮游生物食者、碎屑食者和杂食者.各功能群中肉食者的多样性指数最高,杂食者最低.各功能群的丰度、均匀度指数、多样性指数一般都是中潮区较高,高潮区和低潮区较低.大型底栖动物功能群的分布随潮区环境的改变而变化,是对生境状况的综合反映.     

Contemporary distributions of saltmarsh diatoms in the Seymour–Belize Inlet Complex, British Columbia, Canada: Implications for studies of sea-level change

Surface diatom samples were collected from two saltmarshes in the Seymour–Belize Inlet Complex on the mainland coast of British Columbia to examine the controls on diatom distribution and provide training sets for regional studies of sea-level change. Cluster analysis and detrended correspondence analysis (DCA) were used to examine diatom distributions within and between the two marshes whilst canonical correspondence analysis (CCA) and partial canonical correspondence analysis (pCCA) were used to analyse species–environment relationships. The diatom assemblages were shown to be quite different between the two marshes, Waump and Wawwat'l. Q-mode cluster analysis separated the diatom assemblages from Waump into three zones corresponding to recognized vegetation zones; diatom zone C corresponds to the high marsh and is dominated by Pinnularia obscura, Caloneis bacillum and Diploneis ovalis, zone B spans the high- and mid marsh and is characterised by Fragilaria exigua, Nitzchia palea and D. ovalis, whilst zone A from the low marsh/tidal flat includes high frequencies of Diploneis smithii, Ctenophora pulchella and F. exigua. Three different diatom zones were recognised at Wawwat'l; two upper zones, E and F, corresponding to the mid-marsh which are dominated by mixed abundances of N. palea and Fragilaria construens var. venter, Navicula peregrina and Navicula clementis and zone D from the low marsh/tidal flat with increased frequencies of F. exigua and Achnanthes delicatula. At Waump, the major controls on diatom distribution were found to be elevation and to a lesser extent pH, whilst at Wawwat'l, the primary controls were grain size/organic content, conductivity and elevation. The results confirm that saltmarsh diatoms may have potential for future sea-level studies in this region. However, caution must be exercised as the significant difference in the diatom assemblages between the two marshes suggests that the diatoms are responding to a number of environmental variables on a local scale. Some of this variation may reflect highly seasonal changes in sedimentation and salinity resulting from the proximity of the marshes to adjacent streams which are subject to spring freshet. Local differences in elevation between the collected samples may also account for some of the assemblage variability. The results emphasize the great diversity and high sensitivity of intertidal diatom species to environmental controls.     

Age structure,growth and reproduction ofRutilus lemmingii in an intermittent stream of the Guadalquivir river basin,southern Spain

The age, growth and reproduction ofRutilus lemmingii (Steindachner, 1866), an endemic cyprinid from the Iberian Peninsula, was studied for over a period of two years in a small seasonal tributary of the Guadalquivir river basin. Approximately 65 % of the total growth in length occured in the first year of life. Males reached a maximum age of 3+ yr (Fork Length, F.L. = 114 mm) and females 4+ yr (F.L. = 144 mm). Both sexes matured during their second year of life (1 +). The overall sex ratio (334 males to 389 females) differed significantly from unity. Somatic condition decreased markedly during the reproductive period of March to May.R. lemmingii is a multiple spawner and releases two batches of eggs per female each year. Mean egg diameter of the first batch was larger than the second one. The regression between fecundity and Fork Length (mm) was: Fec = 0.014 F.L.Z.^2.858 Compared with available information, thisR. lemmingii population, located at a lower latitude, is characterized by fast growth, early maturity, high level of reproductive effort, and a short life-span. These life-history characteristices are typical of species in unstable environments, where adult mortality is high, variable or unpredictable.     

THE COST OF MERISTEM LIMITATION IN POLYGONUM ARENASTRUM: NEGATIVE GENETIC CORRELATIONS BETWEEN FECUNDITY AND GROWTH

Growth and reproduction in higher plants depend on meristems, which have three developmental fates. A meristem can become reproductive, but doing so terminates its activity, it can differentiate vegetatively, or it can remain quiescent for extended periods. The first two fates are mutually exclusive, and only the second leads to the production of additional meristems for subsequent growth and reproduction. In Polygonum arenastrum (frequently referred to as P. aviculare in North American Floras), an annual species lacking quiescent meristems, a quantitative genetic analysis of inbred full-sibling families revealed genetic variation in the developmental pattern of axillary meristem commitment to vegetative growth versus reproduction. Developmental variation resulted in family differences in the age of first reproduction, in age-specific fecundity and growth, and in final plant size and reproductive output. Furthermore, there were strong negative genetic correlations between age-specific growth and fecundity. Early commitment of meristems to reproduction favors high early fecundity, but reduces the number of meristems available for vegetative differentiation, and leads to lowered growth rates and fecundity later in life, when meristems are limiting. Conversely, meristem commitment to vegetative growth early in life results in low early fecundity but high late fecundity and growth. Meristem limitation, like resource limitation, is a proximate mechanism that generates trade-offs between life history traits. Differences between meristem limitation and resource limitation are discussed. Meristem limitation leads automatically to a senescent life history because of the determinate fate of reproductive meristems. Developmental characters were also found to be genetically correlated with metamer characters (leaf size, internode length) and seed size in this selfing species. The pattern of correlation is suggestive of selection for particular suites of life history and morphological characters.     

Intraspecific variation in the population dynamics and growth of the limpet,Cellana tramoserica

Summary Populations of the limpet Cellana tramoserica (Sowerby) from high, mid, and low intertidal regions, and from a subtidal zone, were studied at Cape Banks (N.S.W.), Australia. Individuals from the subtidal population had the largest mean and maximum shell size, the low and high shore populations were intermediate, while the midshore population had both the smallest mean and maximum size. The density of adults showed the reverse trend: the midshore region had the greatest adult population density while the subtidal population had the smallest density. The density of juveniles and recruits was negatively correlated with tidal height in the intertidal areas, but the density of both age-classes was smallest in the subtidal region. The rate of growth of individuals was negatively correlated to the adult density of a site; individuals from the subtidal population grew the fastest, while those from the midshore region grew the slowest. The level of adult mortality of the four populations was similar, with an annual rate of between 50–60%. Juvenile mortality did, however, differ among populations; reduced percentages of juveniles reached adult size in the mid and low shore populations than in the highshore and subtidal populations.The expected lifetime fecundity of individual females differed among the four populations. Subtidal individuals were expected to spawn 40 times the amount of gonad material during their life than were individuals from the midshore population. Individuals from the high and low populations probably spawn 10 times more than those from the midshore region. Differences in the fecundities of individuals were not likely to be the result of genetic differences because experimental transplants and manipulations of density showed that individuals from all the intertidal populations could increase their growth rate to match that of subtidal individuals. The implications of the difference in lifetime fecundity among populations are discussed.     

The conservation ecology ofIris rossii Baker (Iridaceae), a threatened plant in rural Japan

Spatial distribution, size structure and reproductive activities of a population ofIris rossii were examined in managed secondary grassland with scattered pines and hardwoods. Size structure and fecundity patterns among individuals were different between the three sites, which were an open area, under pine canopy, and under hardwood canopy. Growth and reproductive parameters of the species were significantly different at each site. In the open area, mean shoot number of individuals was 9.17, and it was 6.37 under the pine canopy and 5.63 under the hardwood canopy. Fruit set ratio was 26.8% in the open area, 21.1% under the pine canopy and 12.1% under the hardwood canopy. Six seedlings were found in the open area and one under the pine canopy, while no seedlings occurred under hardwood canopy. Most of the individuals distributed in the sites where the height of herbaceous layer was low. These results suggest thatI. rossii can not grow in the closed, especially hardwood, canopies or tall herbaceous layer. Therefore, human interventions such as annual mowing for the restriction of the growth of dominant grasses and tree saplings are essential for the persistence of the population of the species.