Assessment of factors affecting heart rate of the limpet Patella vulgata on the natural shore

 Heart rate variations of a population of the limpet Patella vulgata were monitored in the natural environment (Lough Hyne, southern Ireland) by non-invasive, optoelectronic recording. The heart rates of 145 limpets of different sizes, living on vertical and horizontal substrata, were measured both in air and water at different environmental temperatures, while the animals were inactive on their home scars. The heart rates of emersed, inactive limpets were positively related to air temperature and negatively related to limpet size. These relationships were similar for limpets on vertical and horizontal substrates. In contrast, no significant relationship between heart rate and temperature was found in submerged limpets, probably due to the narrow thermal range of the water during the study period. During submersion, a significant negative relationship between heart rate and size was evident for limpets on vertical surfaces but not for limpets on horizontal surfaces. In general, submerged limpets had a higher heart rate, 1.16 times that of limpets exposed to air. Moreover, the heart rates of nine animals were recorded while they were moving and while inactive on their home scars. Active limpets had a faster heart rate, 1.6 times that of limpets resting on their home scars. The dependence of heart rate on environmental temperature, size, respiratory medium and activity, as observed in limpets on the shore, agrees well with laboratory data and with previous findings of the correlation of oxygen consumption with the same factors. Such in situ measurements may, therefore, prove useful in attempts to determine natural levels of energy expenditure in models on the behaviour of foraging molluscs. Received: 5 September 1999 / Accepted: 17 April 2000     

Effects of Forest Fragmentation on a Dung Beetle Community in French Guiana

Abstract:  Fragmentation is the most common disturbance induced by humans in tropical forests. Some insect groups are particularly suitable for studying the effects of fragmentation on animal communities because they are taxonomically and ecologically homogenous. We investigated the effects of forest fragmentation on a dung beetle species community in the forest archipelago created in 1994–1995 by the dam of Petit Saut, French Guiana. We set and baited an equal number of pitfall traps for dung beetles on three mainland sites and seven island sites. The sites ranged from 1.1 to 38 ha. In 250 trap days, we captured 50 species in 19 genera. Diversity indices were high (2.18–4.06). The lowest diversity was on the small islands and one mainland site. Species richness and abundance were positively related to fragment area but not to distance from mainland or distance to the larger island. The islands had lower species richness and population than mainland forest, but rarefied species richness was relatively invariant across sites. There was a marked change in species composition with decreasing fragment that was not caused by the presence of a common fauna of disturbed-area species on islands. Small islands differed from larger islands, which did not differ significantly from mainland sites. Partial correlation analyses suggested that species richness and abundance of dung beetle species were positively related to the number of species of nonflying mammals and the density index of howler monkeys ( Alouatta seniculus ), two parameters positively related to fragment area.     

Correlations between Human Collecting and Intertidal Mollusc Populations on Rocky Shores

We tested whether the population structures of intertidal molluscs varied with the activity of human collectors in northern Port Phillip Bay, Victora, Australia. We examined eight sites, two of which had been protected for around 70 years by the area being used as a rifle range with the other six being visited frequently by human. The shores were examined in two different years, and in 1991 we counted humans and recorded their activities. At heavily visited shores, almost half of the people seen were simply walking, while 25% were actively collecting, despite regulations prohibiting the collection of intertidal gastritis. The remaining people were fishing or skydiving. At the protected shores, we observed only one collector and a handful of walkers over more than 20 observation periods, including weekdays, normal weekends, and holiday weekends. In 1989 and 1991, we compared size distributions of seven intertidal species between sites in the two categories. Four mollify species are taken for food and bait; the remaining three served as “controls”. The size distributions and abundance of noncarbonated species (alicized napalm, Leaderless viola, and emization ebonies) did not differ between visited and protected sites in either year, although most species did vary between individual sites. In contrast, three of the four collected species, alicize trampoliners, Austrocochlea constricta, and Nerita atramentosa, were significantly larger at the protected sites, and N. atramentosa was markedly less abundant at heavily visited sites. The fourth collected species, Turbo undulatus, has a distribution extending into the subtidal zone, and it is possible that intertidal populations ate replenished from deeper water. The changes in mean size of molluscs are well within the range reported from South America and South Africa under subsistence fishing, and they demonstrate that at least close to major urban centers, human recreational activities can have substantial effects in developed countries.     

Sex change in a giant endemic limpet,Patella kermadecensis,from the Kermadec Islands

Sex-ratios of the limpetPatella kermadecensis Pilsbry, collected at three times of the year during 1984 and 1985 at Raoul Island in the Kermadecs group, varied with the size of the limpets. Small limpets were predominantly male, while amongst larger ones about half were female. The presence of hermaphroditism inP. kermadecensis was confirmed by microscopic examination of gonad material from over 50 limpets, which revealed a high percentage of individuals with both male and female gametes in their gonads. It is postulated that protandric sex change occurs in this large limpet, and that this phenomenon is related to a change in habitat from the backs of large shells (limpets less than approximately 50 mm) to bedrock (larger limpets). The pattern of sex change inP. kermadecensis is compared with that in other species of limpet, and it is suggested that a reproductive system consisting of gonochoristic males and sequential hermaphrodites may be particularly advantageous for a large, slow-growing species that has a very restricted geographical range.     

No evidence for genetic differentiation between Antarctic limpet Nacella concinna morphotypes

The extent to which genetic divergence can occur in the absence of physical barriers to gene flow is currently one of the most controversial topics in evolutionary biology, with implications for our understanding of speciation, phenotypic plasticity and adaptive potential. This is illustrated by a recent study reporting a surprising pattern of genetic differentiation between intertidal and subtidal morphotypes of the broadcast-spawning Antarctic limpet Nacella concinna. To explore this further, we collected almost 400 Antarctic limpets from four depths (intertidal, 6, 15 and 25 m) at Adelaide island, Antarctica, and conducted a combined morphometric and genetic analysis using 168 polymorphic amplified fragment length polymorphism (AFLP) loci. Morphological analysis revealed not only pronounced differences between the two morphotypes, but also a continuous cline in shell shape from the intertidal zone down to 25 m depth, suggesting that the distinction between the morphotypes may be artificial. Moreover, genetic analysis using both F _st and a Bayesian analogue found no evidence for differentiation either between the two morphotypes or by depth, and a Bayesian cluster analysis did not detect any cryptic genetic structure. Our findings lend support to the notion that limpets can be phenotypically highly plastic, although further studies are required to determine unequivocally whether there is any genetic basis to the observed variation in shell morphology.     

Individual- and population-level responses of a keystone predator to geographic variation in prey

Investigating how food supply regulates the behavior and population structure of predators remains a central focus of population and community ecology. These responses will determine the strength of bottom-up processes through the food web, which can potentially lead to coupled top-down regulation of local communities. However, characterizing the bottom-up effects of prey is difficult in the case of generalist predators and particularly with predators that have large dispersal scales, attributes that characterize most marine top predators. Here we use long-term data on mussel, barnacle, limpet, and other adult prey abundance and recruitment at sites spread over 970 km to investigate individual- and population-level responses of the keystone intertidal sunstar Heliaster helianthus on the coast of Chile. Our results show that this generalist predator responds to changes in the supply of an apparently preferred prey, the competitively dominant mussel Perumytilus purpuratus. Individual-level parameters (diet composition, per capita prey consumption, predator size) positively responded to increased mussel abundance and recruitment, whereas population-level parameters (density, biomass, size structure) did not respond to bottom-up prey variation among sites separated by a few kilometers. No other intertidal prey elicited positive individual predator responses in this species, even though a large number of other prey species was always included in the diet. Moreover, examining predator-prey correlations at approximately 80, 160, and 200 km did not change this pattern, suggesting that positive prey feedback could occur over even larger spatial scales or as a geographically unstructured process. Thus individual-level responses were not transferred to population changes over the range of spatial scales examined here, highlighting the need to examine community regulation processes over multiple spatial scales.     

Effects of a complex archipelago on genetic subdivision of the intertidal limpet Siphonaria kurracheensis

Allozyme variation in the intertidal limpet Siphonaria kurracheensis was examined in 11 populations from the Houtman Abrolhos Islands, Western Australia, and four from the adjacent mainland, to test if the effects of these islands on genetic subdivision should include species with high dispersal. Consistent with extensive planktonic dispersal, genetic subdivision was low, with an average standardized variance in allelic frequencies (F_ST) of 0.010 over 400 km along the mainland, and 0.009 over 70 km in the Abrolhos Islands. Nevertheless, subdivision was statistically significant at the smallest scale examined (about 10 km), showing a pattern of local heterogeneity and large-scale homogeneity, which is common in planktonic dispersers. Among the island populations, heterozygosity was positively correlated with a multivariate index of exposure. The level of heterozygosity at the most exposed island sites is equivalent to that at the (also exposed) mainland sites, whereas it is lower in the most sheltered island sites. Although this indicates that some island populations are less well connected to the major pool of larvae, the overall effect of the archipelago on genetic subdivision is no greater than occurs on the mainland coast. This is contrary to findings in previously studied species with less potential for dispersal, suggesting that the local impediments to dispersal are selective in their effects, and are unlikely to be broadly important for genetic divergence in widely dispersing species.     

Substratum relationships of some tropical Pacific intertidal gastropods

Among tropical Indo-West Pacific herbivorous intertidal gastropods, high intertidal neritids, and limpets and certain lower intertidal limpets are restricted to a narrow range of substratum types. Littorinids, planaxids, low intertidal neritids, most lower intertidal limpets, and the trochid Monodonta labio tolerate a wide variety of substrata. This pattern of substratum specialization is related to the overall adaptive strategies of the various groups: those with a large area of soft tissue in contact with the substratum (neritids and limpets) often specialize to a particular substratum, especially at high shore levels, while those in which the area of contact between animal and substratum is small do not. High intertidal neritids and limpets able to live on a wide range of substrata tend to be species of limited geographic distribution. It is concluded that, even within a tropically similar group of animals (raspers in this case) relevant dimensions of niche separation depend upon the structure of the environment as well as upon the structure of the animals themselves.     

Effects of Human Exclusion on Parasitism in Intertidal Food Webs of Central Chile

Abstract:  Numerous ecological studies have demonstrated the dramatic effects that humans have on coastal marine ecosystems. Consequently, marine reserves have been established to preserve biodiversity. Recent reviews show that this strategy has paid off because inside reserves, most species have rapidly increased in size and abundance. Even though these studies focused on free-living organisms and paid little attention to parasite populations, numerous authors support the hypothesis that parasitism levels could be good indicators of ecosystem stability. We examined harvesting effects on the dynamics of a parasitic trematode ( Proctoeces lintoni ) that completes its life cycle in intertidal mussels ( Perumytilus purpuratus ), keyhole limpets ( Fissurella crassa ), and clingfish ( Sicyases sanguineus ). All of these species are directly or indirectly affected by humans. Prevalence and abundance of the trematode P. lintoni in the three host species were compared in four study sites that differed in the intensity of human harvest. Parasitism infection in limpets and mussels was significantly higher in areas protected from human harvesting than in open-access areas, which suggests a significant change in parasite dynamics inside reserves. Yet the average parasitic biomass found in the gonads of F. crassa did not differ between protected and open-access areas. These results show, then, that the parasite system responded by increasing infection rates in marine protected areas without implication for reproductive success of the intermediate host. Our findings show that the indirect effects of harvesting by humans on the embedded parasite communities of littoral ecosystems require further scientific investigation.     

Parasitic castration in slipper limpets infested by the symbiotic crab Calyptraeotheres garthi

Two ill-explored hypotheses might explain host castration by parasitic pea crabs. The ‘energy drain’ hypothesis states that castration is caused by host-derived nutrient consumption of parasites that ultimately diminishes host-energy intake. The ‘steric interference’ hypothesis states that castration occurs when parasites physically inhibit host reproduction. This study evaluated whether Calyptraeotheres garthi, a pea crab from the southwestern Atlantic, is a parasitic castrator and explored whether the two hypotheses above explain castration in the limpet Crepidula cachimilla. None of three studied limpet species brooded embryos during the reproductive season when infested by mature female pea crabs. Also, limpets of C. cachimilla infested by C. garthi did not reproduce during a 90-day experimental period while crab-free limpets did spawn embryos during this period. Limpets resumed reproduction soon after pea crabs were experimentally removed from their brooding chamber. Thus, C. garthi does castrate limpets, and castration is reversible. Pea crabs ‘steal’ food from limpets, and infested limpets did not modify their feeding behavior to counteract nutrient loss. Thus, infested limpets are expected to ingest less food which provides partial support for the ‘energy drain’ hypothesis. However, the limpet’s body condition increased or was not affected by pea crabs during the breeding season which argues against the same hypothesis. Furthermore, that limpets promptly recovered reproductive activity once pea crabs were experimentally removed, that castration was not induced by the smallest pea crabs in the population (that fill only partially the brooding chamber), and that parasitized limpets did exhibit fully mature ovaries, support the ‘steric interference’ hypothesis explaining parasitic castration.     

Distribution,movement and diet of the snailSearlesia dira in the intertidal community of San Juan Island,Puget Sound,Washington

Searlesia dira Reeve is a locally abundant, carnivorous gastropod that occurs from Alaska to central California. I studied populations of this snail at a variety of sites on San Juan Island in order to provide information on their distribution and abundance and to analyze their community role, i.e., their potential impact on their prey populations and their dietary overlap with co-occurring invertebrate predators. I established permanent transects, both random and in crevices, at three areas of differing exposure to wave action. On these transects I recorded the number and size ofS. dira, activity, distance to nearest neighbor, other organisms present, and tidal height on four phases of the mixed semi-diurnal tidal cycle. The results are (1) an increase in density with increase in substrate relief, (2) an occurrence of highest densities in moderately exposed and exposed sites rather than in protected ones, (3) an increase in mean size with increase in depth along the intertidal gradient, (4) an increase in both total activity and feeding success on the high water immediately following an extreme minus low tide occurring during the daytime in the summer, (5) a relative restriction of feeding success to that high tide, and (6) a similarity of the diet to the relative abundance of the main observed prey for most areas. The main exceptions are chitons and one lower intertidal limpet, all of which are taken in excess of their apparent abundance and especially in the middle intertidal. The upward extension of some lower intertidal prey populations, such as chitons, may be limited by the increased susceptibility of individuals at the upper margin of the population to predators such asS. dira; susceptibility appears to be increased after periods with a higher probability of increased physiological stress, such as extreme low tides on summer days.     

Size-assortative mating and non-reciprocal copulation in a hermaphroditic intertidal limpet: test of the mate availability hypothesis

Mating behaviour and mating patterns are affected to a large extent by body size in both hermaphrodites and gonochorists. Detailed research on mating patterns, mate choice, pre- and post-copulatory sexual selection in marine pulmonates is wanting, thus warranting more attention as a study system in the future. The simultaneous hermaphroditic limpet Siphonaria capensis (Pulmonata: Basommatophora) shows size-dependent fecundity, and acts as a suitable organism to test the effects of body size on mating patterns, mating success, gender expression and reciprocity in hermaphroditic marine gastropods. We mainly used bootstrap resampling techniques to estimate the effects of different factors on mating patterns. In the populations studied, a strong size-assortative mating pattern was observed, where small-scale spatial distribution of potential mating partners (the mate availability hypothesis) could explain 65% of this pattern, while mate choice and mating constraints explaining the rest. No significant difference in mating success between limpets with different body size was found, even though in one population, the sperm recipients were larger than the non-copulating limpets. Interestingly, we found that intromission was non-reciprocal during copulations. This may mean that this species does not conform to the common rule of reciprocity predicted for hermaphrodites, unless there is sex-role alternation between individuals in a mating pair. The mating partners consisted of similar sized, acting males and females, thus without any indication of the body size determining the gender expression. The matings took place in early mornings only at spring tides and the animals were observed laying egg masses only during neap tides. Since siphonariid limpets possess both a spermatheca and a seminal vesicle, which may have either a sperm storing and/or digestive function, post-copulatory sexual selection (e.g., sperm competition) cannot be dismissed.     

Life-history features of the marine intertidal limpet Notoacmea scutum (Gastropoda) in central California

Growth of individually marked specimens of Notoacmea scutum (Rathke) was followed for 15 months (April, 1978 to June, 1979) at a field site in central California, USA. Growth rates were highly seasonal, being highest from late spring through early summer and lowest during winter. Growth rate also varied significantly with intertidal height: limpets lower in the intertidal zone grew faster. Both relationships may have been related to the relative abundance of food or to the amount of time available for foraging. Recently settled limpets, which were present throughout the year, grew to approximately 17 mm in length by the end of their first year; by then, they had also reached the size at which mature gonads were consistently found (16 mm). Limpets were almost 26 mm long by the end of the second year. Substantial mortality occurred throughout the year, but it was particularly severe during winter, when a large portion of the reproductive population was lost. This pronounced winter mortality was probably not caused by invertebrate predators such as seastars. Most limpets at the study site were less than 2 yr old.     

Overestimations of food abundance: predator responses to prey aggregation

Understanding and predicting the consequences of trophic interactions for community processes requires knowledge of the role of food availability, which is often wrongly conflated with prey abundance. For prey animals in groups, this is not fully understood. Previous work has shown that oystercatchers more frequently attack solitary rather than aggregated limpets and are more successful in predation attempts on singletons. It has also been demonstrated that an attack on one limpet in a group alerts the entire group, all of which then clamp down and become unavailable. I show that Eurasian Oystercatchers (Haematopus ostralegus L.) attack only one limpet in a group and then move on to attack another individual limpet, and I also demonstrate that the distance they move is greater than the distance at which groups of limpets have been known to detect attacks. Thus in the oystercatcher-limpet predator-prey system on rocky shores, groups of limpets are actually one prey item independently of the number of limpets in the group. This has implications for assessment of food supply for avian predators on rocky shores, with consequences for our understanding of previously documented trophic cascades.     

Analyses of limpet defense and predator offense in the field

Between November 1982 and August 1984 diving observations of two intertidal limpets' defensive behaviours in response to two predatory species were made near the Seto Marine Biological Laboratory of Kyoto University, in Wakayama Prefecture, Japan, for over 800 h. The focus of the observations was a simple isolated rock, 0.9 m² in area with an inclination of 75°. The pulmonate homing limpet Siphonaria sirius responded by fleeing from its home in response to the whelk Thais clavigera and by calmping tightly to its home scar in response to the starfish Coscinasterias acutispina. Both behaviours were successful for avoiding predation. After a whelk moved far away, limpets returned to their homes. The patellid limpet Cellana toreuma showed mantle folding behaviour or fled for an average distance of 8 cm vertically upwards in response to the whelk and for an average distance of 21 cm vertically upwards in response to the starfish. The amount of time predators spent foraging increased in the low intertidal zone relative to the mid and high zones, and most predator attacks were observed during the limpets' resting periods (submersion in late afternoon or nighttime). The upward directional flight displayed by C. toreuma was related to the upward orientation of the limpets' heads when they are in a resting position. Half of the limpets observed fleeing downward away from a starfish, a movement elicited by a downward attack, received a second attack. These limpets were preyed upon at the second encounter 67% of the time. The upward fleeing behaviour of C. toreuma resulted in the shifting of its resting site higher up the study site where fewer attacks occurred and where the mortality rate was at its lowest. It is suggested that the homing behaviour of S. sirius and the upward fleeing behaviour of C. toreuma are mechanisms by means of which interspecific competition between limpets is reduced and coexistence on a small rock surface is maintained.     

Ecological studies on a tropical limpet,Cellana radiata. Structural variations in the shell in relation to distribution

Structural variations in the shell of the tropical limpet Cellana radiata (Born) inhabiting the high and low water zones and the exposed and protected surfaces at Palm Beach, Waltair Coast, India, were investigated. Attempts have also been made to study which environmental factors have a marked influence upon the characters of the shell. The results are discussed and are compared with those on temperate climate limpets.     

Population differentiation and taxonomic status of the exploited limpet Patella candei in the Macaronesian islands (Azores,Madeira, Canaries)

There has been considerable confusion in the taxonomy of limpets of the North East Atlantic Ocean and Mediterranean Sea, particularly those from the Macaronesian islands. The present study compared populations of the intertidal limpet Patella candei d'Orbigny from the Azores, Madeira and the Canaries with those of P. caerulea Linnaeus and P. depressa Pennant from the European and African continental coasts. No major differences in radular morphology were detected between the three species. However, electrophoretic analysis of 15 enzyme loci gave overall genetic identity (I) values of 0.5 between the three species, indicating that they cannot be regarded as conspecific as previously thought, and suggesting that P. candei is endemic to the Macaronesian islands. Comparisons of P. candei within these islands showed that, although populations did not differ with respect to radular morphology and soft-body parts, populations from the Azores were distinct from those in Madeira and the Canaries in shell shape and gene frequencies. Individuals from the Azores had, no average, taller shells and longer radulae, while those in Madeira and the Canaries had a shallow, depressed and stellate shell form. This was interpreted as being due to the wider habitat distribution of the species in the Azores compared to Madeira and the Canaries. Electrophoretic results showed that P. candei from the Azores differed from P. candei in Madeira and the Canaries by almost 40% of the loci investigated (I=0.660), suggesting that the former is a separate endemic species. An I value of 0.969 between populations in Madeira and the Canaries was typical of conspecific populations.     

Food availability and diet composition of three coexisting Mediterranean limpets (Patella spp.)

The gut contents of three intertidal patellid limpets were analysed by collecting foraging specimens on a breakwater on the Tyrrhenian coast (central Italy) between May 1988 and October 1989. The three species coexist there showing a different, but partially overlapping zonation: Patella aspera dominates the infralittoral fringe; the majority of P. caerulea inhabits the lower midlittoral, while P. rustica is most abundant in the upper midlittoral. The algae present on slivers of substratum over which each limpet collected was moving were identified. Moreover, floristic surveys were made along the shore in order to characterize the algal cover of the different zonal belts. The floristic study revealed that the basic elements of algal communities typical of western Mediterranean rocky shores are present in the study area. The algae found on the slivers under the foraging limpets were generally representative of the algal community typical of the same zone. There was a marked difference between the diets of P. rustica and P. aspera due to the fact that the first species forages on a few low lying epilithic and endolithic Cyanophyceae, while P. aspera feeds on a large number of species belonging to all the main algal classes and life forms considered, including frondose epilithics and epiphytics. The diet of P. caerulea resembles that of P. aspera in algal heterogeneity, but is dominated by Cyanophyceae as in P. rustica. A detailed analysis of the differences between gut contents of each limpet species and the relative slivers showed an obvious general correspondence, but revealed also that the diets of the three species do not completely reflect the availability of algae. These findings suggest that the basic diet segregation mechanism between the three populations is their zonal separation. However, the difference in gut contents of heterospecific limpets foraging in the same zone suggests the existence of supplementary morphy-functional or behavioural mechanisms for diet segregation between the three species.     

Effects of Artisanal Fishing on Marine Communities in the Galápagos Islands

Abstract: The Galápagos Islands harbor some of the least impacted marine ecosystems in the tropics, but there are indications that local artisanal fishing is affecting exploited marine communities. To quantify these effects, I sampled communities of fishes and sea urchins at a number of heavily fished and lightly fished sites throughout the central islands of the archipelago. Sites were selected based on information collected as part of a local fisheries monitoring study and standardized across a number of abiotic factors. Abundance and biomass of the primary target species were significantly lower in the heavily fished sites than in the lightly fished sites. Community structure also differed between heavily and lightly fished sites. Cluster analyses of the full community of fishes and a subset of nontarget fishes revealed that sites within a treatment were more similar to one another than sites between treatments. Herbivorous fishes tended to be lower in abundance and sea urchins tended to be higher in abundance in heavily fished sites, but these differences were not significant. My results are encouraging in that the direct effects of artisanal fishing are limited to the primary target species, which probably results from a high specificity of fishing gear. The differences in community structure, however, suggest that artisanal fishing also has cascading effects on noncommercial species throughout the community. An improved understanding of important ecological interactions, increased ecological and fishery monitoring, and effective precautionary management are needed to ensure that human effects in these waters remain minimal.     

Genetic Effects of Habitat Fragmentation on Common Species of Swiss Fen Meadows

Abstract:  The area of Caricion davallianae alliance in Switzerland has been considerably reduced and fragmented during the last 150 years. We assessed the genetic variability, inbreeding level, and among-population differentiation of two common habitat-specific plant species, Carex davalliana SM. and Succisa pratensis Moench, in 18 Caricion davallianae fen meadows subjected to fragmentation. We used a spatial field design of fen systems (six systems total), each consisting of one large habitat island and two small habitat islands. We used allozyme electrophoresis to derive standard genetic parameters ( A, P, H_O, H_E, F_IS, F_ST ). In Carex we identified a consistently lower A in isolated habitat islands; furthermore, H_E was lower in small habitat islands than in large habitat islands. In Succisa we identified a lower H_O in small habitat islands than in larger ones. Small habitat islands were marginally significantly differentiated (  F_ST ) from large islands for Succisa . For both species, no effects were evident for F_IS ; therefore, we argue that genetic drift rather than inbreeding is the main cause of the observed differences. The genetic structure of Carex and Succisa in small habitat islands differed from that in large habitat islands, but differences were small. It appears that the observed differences in genetic variability among fen meadows correspond to observed differences in fitness and demographic traits. We show that habitat fragmentation affects not only the rare species in an ecosystem but also reduces the survival probabilities of common species. One of the main goals of conservation should be to mitigate fragmentation of natural habitats in order to increase population sizes and connectivity.