The interplay between environmental conditions and allee effects during the recovery of stressed zooplankton communities

Many important ecological phenomena depend on the success or failure of small introduced populations. Several factors are thought to influence the fate of small populations, including resource and habitat availability, dispersal levels, interspecific interactions, mate limitation, and demographic stochasticity. Recent field studies suggest that Allee effects resulting from mate limitation can prevent the reestablishment of sexual zooplankton species following a disturbance. In this study, we explore the interplay between Allee effects and local environmental conditions in determining the population growth and establishment of two acid-sensitive zooplankton species that have been impacted by regional anthropogenic acidification. We conducted a factorial design field experiment to test the impact of pH and initial organism densities on the per capita population growth (r) of the sexual copepod Epischura lacustris and the seasonally parthenogenetic cladoceran Daphnia mendotae. In addition, we conducted computer simulations using r values obtained from our experiments to determine the probability of extinction for small populations of acid-sensitive colonists that are in the process of colonizing recovering lakes. The results of our field experiment demonstrated that local environmental conditions can moderate the impacts of Allee effects for E. lacustris: Populations introduced at low densities had a significantly lower r at pH 6 than at pH 7. In contrast, r did not differ between pH 6 and 7 environments when E. lacustris populations were introduced at high densities. D. mendotae was affected by pH levels, but not by initial organism densities. Results from our population growth simulations indicated that E. lacustris populations introduced at low densities to pH 6 conditions had a higher probability of extinction than those introduced at low densities to a pH 7 environment. Our study indicates that environmental conditions and mate limitation can interact to determine the fate of small populations of sexually reproducing zooplankton species. If a more rapid recovery of acid-damaged zooplankton communities is desired, augmentation of dispersal levels may be needed during the early phases of pH recovery in order to increase the probability of establishment for mate-limited zooplankton species.     

Adaptive reversals in acid tolerance in copepods from lakes recovering from historical stress.

Anthropogenic habitat disturbance can often lead to rapid evolution of environmental tolerances in taxa that are able to withstand the stressor. What we do not understand, however, is how species respond when the stressor no longer exists, especially across landscapes and over a considerable length of time. Once anthropogenic disturbance is removed and if there is an ecological trade-off associated with local adaptation to such an historical stressor, then evolutionary theory would predict evolutionary reversals. On the Boreal Shield, tens of thousands of lakes acidified as a result of SO2 emissions, but many of these lakes are undergoing chemical recovery as a consequence of reduced emissions. We investigated the adaptive consequences of disturbance and recovery to zooplankton living in these lakes by asking (1) if contemporary evolution of acid tolerance had arisen among Leptodiaptomus minutus copepod populations in multiple circum-neutral lakes with and without historical acidification, (2) if L. minutus populations were adaptively responding to reversals in selection in historically acidified lakes that had recovered to pH 6.0 for at least 6-8 years, and (3) if there was a fitness trade-off for L. minutus individuals with high acid tolerance at circum-neutral pH. L. minutus populations had higher acid tolerances in circum-neutral lakes with a history of acidification than in local and distant lakes that were never acidified. However, copepods in circum-neutral acid-recovering lakes were less acid-tolerant than were copepods in lakes with longer recovery time. This adaptive reversal in acid tolerance of L. minutus populations following lake recovery was supported by the results of a laboratory experiment that indicated a fitness trade-off in copepods with high acid tolerances at circum-neutral pH. These responses appear to have a genetic basis and suggest that L. minutus is highly adaptive to changes in environmental conditions. Therefore, restoration managers should focus on removing environmental stressors, and adaptable species will be able to reverse evolutionary responses to environmental disturbance in the years following recovery.     

Allee Effect and the Uncertainty of Population Recovery

Recovery of depleted populations is fundamentally important for conservation biology and sustainable resource harvesting. At low abundance, population growth rate, a primary determinant of population recovery, is generally assumed to be relatively fast because competition is low (i.e., negative density dependence). But population growth can be limited in small populations by an Allee effect. This is particularly relevant for collapsed populations or species that have not recovered despite large reductions in, or elimination of, threats. We investigated how an Allee effect can influence the dynamics of recovery. We used Atlantic cod (Gadus morhua) as the study organism and an empirically quantified Allee effect for the species to parameterize our simulations. We simulated recovery through an individual‐based mechanistic simulation model and then compared recovery among scenarios incorporating an Allee effect, negative density dependence, and an intermediate scenario. Although an Allee effect significantly slowed recovery, such that population increase could be negligible even after 100 years or more, it also made the time required for biomass rebuilding much less predictable. Our finding that an Allee effect greatly increased the uncertainty in recovery time frames provides an empirically based explanation for why the removal of threat does not always result in the recovery of depleted populations or species. El Efecto Allee y la Incertidumbre de la Recuperación de Poblaciones     

Assessing Changes in Amphibian Population Dynamics Following Experimental Manipulations of Introduced Fish

Abstract: Sport‐fish introductions are now recognized as an important cause of amphibian decline, but few researchers have quantified the demographic responses of amphibians to current options in fisheries management designed to minimize effects on sensitive amphibians. Demographic analyses with mark–recapture data allow researchers to assess the relative importance of survival, local recruitment, and migration to changes in population densities. I conducted a 4‐year, replicated whole‐lake experiment in the Klamath Mountains of northern California (U.S.A.) to quantify changes in population density, survival, population growth rate, and recruitment of the Cascades frog (Rana cascadae) in response to manipulations of non‐native fish populations. I compared responses of the frogs in lakes where fish were removed, in lakes in their naturally fish‐free state, and in lakes where fish remained that were either stocked annually or no longer being stocked. Within 3 years of fish removals from 3 lakes, frog densities increased by a factor of 13.6. The survival of young adult frogs increased from 59% to 94%, and realized population growth and recruitment rates at the fish‐removal lakes were more than twice as high as the rates for fish‐free reference lakes and lakes that contained fish. Population growth in the fish‐removal lakes was likely due to better on‐site recruitment of frogs to later life stages rather than increased immigration. The effects on R. cascadae of suspending stocking were ambiguous and suggested no direct benefit to amphibians. With amphibians declining worldwide, these results show that active restoration can slow or reverse the decline of species affected by fish stocking within a short time frame.     

Floater mortality within settlement areas can explain the Allee effect in breeding populations

The Allee effect (the positive relationship between population growth rate and population size) is a constraint of some animal populations at low numbers, which increases their likelihood of extinction because of a decrease in reproduction and/or survival. We were able to demonstrate that the Allee effect can be the result of a mortality increase affecting floaters (i.e. dispersing individuals able to enter as breeders in the reproductive population when a breeding territory or a potential mate – owner of a suitable breeding territory – becomes available). Previously, potential mechanisms underlying Allee effects were always related to the breeding portion of a population only. In contrast, our understanding of or solutions to population declines due to the Allee effects can reside elsewhere, away from breeding territories.     

Impact of predation-disturbance by large epifauna on sediment-dwelling harpacticoid copepods: Field experiments in a subtidal seagrass bed

The hypothesis was tested that predation-disturbance by epibenthic macrofauna affects the abundance of sediment-dwelling harpacticoid copepods. The copepods inhabited a subtidal seagrass (Zostera marina L.) bed in British Columbia, Canada. The response of the harpacticoid community was observed in controlled field experiments in which epibenthic predators-disturbers were excluded from portions of the seagrass bed. Controlled, exclusion-cage (0.8 m² area, 7-mm mesh) experiments were conducted within the seagrass bed from late March/early April to mid-June in both 1986 and 1987. Sampling was conducted biweekly. Exclusion of large epibenthic predators-disturbers had little effect on sediment-dwelling harpacticoid copepod density. Total harpacticoid numbers and abundances of dominant species generally did not increase in the exclusion treatment relative to the control. Shifts in species composition of the harpacticoid community did not occur. The treatment control was adequate in simulating the exclusion cage structure. It appears that large epibenthic predators-disturbers have little impact on the abundance of harpacticoid copepod populations at this study site.     

Observations on the diurnal vertical migrations of an oceanic animal community

Diurnal changes in abundance caused by vertical migrations have been examined in populations of copepods, ostracods, euphausiids, amphipods, decapods, chaetognaths, siphonophores and fish. The animals were taken in a series of hauls made over a 24 h period with an opening-closing midwater trawl system (RMT 1+8), consisting of a net of 1 m² mouth area combined in the same frame as one of 8 m² mouth area. The samples were taken at 250 m depth in a position 30°N; 23°W on 7/8 April 1972. The specific composition of the community and the numbers of individuals changed continuously with time. The numbers of fish, decapods and chaetognaths increased at night, but those of copepods, ostracods and euphausiids decreased. More species of fish, decapods and copepods were present by night than by day, whereas the numbers of species per haul for other groups remained fairly constant. The relative abundances of groups caught by the RMT 1 have been analysed, but similar treatment of the RMT 8 samples was impossible as only 3 groups were taken from this net. Non-migrants were a minority in every group except chaetognaths. Migrant species have been put into one of 6 transitory categories according to their patterns of abundance and hence migrations. Within each category, migratory behaviour varied both inter- and intraspecifically. The patterns of abundance of many species were smooth and continuous, suggesting slow migratory cycles of small amplitude. Conversely, extensive migrants had discontinuous patterns and presumably more rapid movements. Few migrants had a steady numerical plateau between their upward and downward migrations, and most apparently moved up or down continuously. The presence of migratory species in the sampled layer depended upon the time of day or night. It is concluded that, in a vertical series of hauls, the depths of occurrence of migrants will vary with the sampling time. Further-more, a vertical series will show a species minimum migration range but not necessarily its maximum. Individuals of some species were out of phase with the migrations of their main populations. There is evidence that the distributions and migrations of some species of decapods, euphausiids, copepods and fish could be related to the distribution of underwater light. Three pairs of congeneric copepod species were both spatially and temporally segregated for at least part of their diurnal cycles. Such an orderly arrangement could provide a means of reducing competition between species. Some species, however, overtook others on their migrations and the pattern of underwater light cannot, therefore, regulate the distribution of all species in the same way.     

Invading parasitoids suffer no Allee effect: a manipulative field experiment

One frequent explanation for the failure of biological invasions is the Allee effect: due to positive density dependence, initially small invading populations may fail to establish and spread. Populations released for biological control are similar to fortuitous invading populations and may therefore suffer from Allee effects. However, unlike fortuitous invasions, biological control allows the experimental manipulation of initial population size and, thus, offers a unique opportunity to test for the occurrence of Allee effects. We manipulated the initial size of 45 populations of a parasitoid wasp introduced for the biological control of a phytophagous insect and followed the population dynamics of both parasitoids and hosts during three years. Our results suggest an absence of Allee effects but clear negative density dependence instead: (1) the probability of establishment after three years was not affected by initial population size; (2) net reproductive rate was highest at low parasitoid density and high host density; (3) the sex ratio, reflecting the proportion of virgin females, did not increase at low density, suggesting that low densities did not impede mate-finding; (4) the depression of host populations did not depend upon the number of parasitoids introduced. This is, to our knowledge, the first experimental test of the Allee effect in an invading parasitoid. It leads us to propose that a number of behavioral and life-history features of many parasitoids could protect them from Allee effects.     

Etude de la respiration chez des copépodes pélagiques méditerranéens (bassin occidental et Mer Ionienne) et de ses variations en fonction de la bathymétrie des espèces et de leur origine géographique

A shipboard experimental study was conducted on the respiratory rate of pelagic copepods from two Mediterranean stations (Ionian Sea and Occidental Basin). In each sampling area, copepods were collected from different depths (0 to 2500 m) by horizontal net hauls. Selected copepod species included bathypelagic, lower mesopelagic, upper mesopelagic and epipelagic species. Respiration rates were determined at 4 different temperatures (10°, 14°, 18°, 22°C). The metabolism-temperature curves revealed varying degrees of thermal sensibility of copepod respiration related to their respective habitats. During experiments employing a large range of temperatures, bathypelagic species proved most sensitive. By contrast, vertically migrating species appeared to be more tolerant to temperature variation. The relationship between respiratory rate and copepod dry weight was investigated for the 4 temperatures used. A strong correlation emerged between log of respiration and log of weight. The equation of regression lines varied with experimental temperature and with geographic origin of the copepods studied. Comparison of the relation respiration-weight of specimens from the two Mediterranean stations and from a previously studied Atlantic station (Canarian and south Moroccan region), indicated that observed differences are related to the average temperature of the mesopelagic waters in each sampling area. The pressure effect on metabolic rate is discussed. It is suggested that hydrostatic pressure does not affect the characteristics of temperature sensibility of copopods' respiration under the temperature conditions prevailing in their natural habitat.     

Human Impacts on Drainages of the Mesa Central, Mexico, and Its Genetic Effects on an Endangered Fish, Zoogoneticus quitzeoensis

Abstract:  The Mesa Central of Mexico is of special conservation interest due to its high richness of freshwater fish species, of which the goodeines are one of the most representative groups. Through an integrated approach, we determined conservation priorities for goodeine populations. We based our recommendations on the genetic diversity (variation in five microsatellite DNA loci) in 10 populations of Zoogoneticus quitzeoensis and on an analysis of ecological (e.g., presence of exotic species), social (e.g., political situation), and environmental (e.g., pollution) information for 52 historical occurrence points for species in the genus Zoogoneticus . Patterns of genetic erosion and genetic diversity indices were closely associated with human impact. Recent bottleneck events were most evident in the populations from remnants of the lakes drained at the beginning of the twentieth century. We identified seven operational conservation units (OCUs), all of which should be conserved because they contain unique portions of the total variation of the species. Special attention needs to be given to increase genetic variability, recover population sizes, and reestablish contact among populations within OCUs. It is imperative to create an integrative and effective approach for the recovery and conservation of the freshwater fish diversity of Central Mexico that is based on social and natural sciences.     

Comparison of the food of Triphoturus mexicanus and T. nigrescens,two lanternfishes of the Pacific Ocean

Prey (chiefly euphausiids and copepods) eaten by two myctophids (lanternfishes) are compared from incidence in fish stomachs and from abundance in the environment. One lanternfish species, Triphoturus mexicanus, lives in the California Current, and the other, T. nigrescens, lives in the central Pacific Ocean. Although these two environments are very different physically and biologically, the feeding habits of the two lanternfishes are surprisingly similar. Prey biomass is 94% euphausiids, 3% copepods, and 3% other organisms for T. mexicanus and 88% euphausiids, 4.5% copepods, and 7.5% other organisms for T. nigrescens; the difference between the fish species is not significant when tested statistically. The two fishes resemble one another in frequency distributions of ingested copepod individuals, copepod species, euphausiid individuals, and euphausiid species. During a single diurnal feeding period, both fishes eat a variety of copepod species but tend to eat only a single species of euphausiid. T. mexicanus grows to twice the length of T. nigrescens and eats proportionally larger euphausiids; however, both fishes eat copepods having the same median size. The frequencies of euphausiid species in the diets of both fishes differ from the frequencies in the environment. The chief differences between the feeding habits of the two lanternfishes are that T. nigrescens, in comparison to its congener, eats a greater variety of organisms during one diurnal feeding period and captures smaller euphausiids. The feeding patterns for each lanternfish species are consistent over distances of hundreds of kilometers and over many years of sampling.     

Using survival analysis to study translocation success in the Gila topminnow (Poeciliopsis occidentalis).

Translocation, the intentional release of captive-propagated and/or wild-caught animals into the wild in an attempt to establish, reestablish, or augment a population, is a commonly used approach to species conservation. Despite the frequent mention of translocation as an aid in threatened or endangered species recovery plans, translocations have resulted in the establishment of few sustainable populations. To improve the effectiveness of translocation efforts, it is essential to identify and adopt features that contribute to successful translocations. This study analyzed 148 translocations of the endangered Gila topminnow (Poeciliopsis occidentalis) to identify various factors that have significantly influenced translocation success. We quantified success as the "persistence time" of translocated populations and used survival analysis to interpret the role of several factors. The following factors affected persistence times of translocated populations: season in which the fish were translocated, habitat type of the translocation site, and genetic origin of the fish stocked. In general, factors associated with stocking, the population stocked, and the site of translocation can significantly affect the persistence of translocated populations and thus increase the probability of translocation success. For Gila topminnow, future translocations should be undertaken in late summer or fall (not early summer), should occur into ponds (not streams, wells, or tanks), and should generally utilize individuals from genetic lineages other than Monkey Spring. For other species, a key lesson emerging from this work is that life history attributes for each translocated species need to be considered carefully.     

Impact of copepod grazing on the red-tide flagellate Chattonella antiqua

Although planktonic copepods are major suspension feeders in the sea, the impact of their grazing pressure upon red-tide flagellates has not been fully investigated. In the present study, the grazing of adult females of several copepod species is examined using three food types: viz. natural suspended particles, natural suspended particles mixed with cultured Chattonella antiqua, and cultured C. antiqua. The functional response on C. antiqua was investigated for five species of copepods (Acartia erythraea, Calanus sinicus, Centropages yamadai, Paracalanus parvus and Pseudodiaptomus marinus). Ingestion rates increased linearly with increasing cell concentrations until a maximum level was reached, beyond which the rates were constant. This cell concentration was higher for larger copepods. The weight-specific maximum ingestion rates were higher in the small species. In general, copepods tended to feed selectively on larger particles when feeding on natural particles. This tendency was strongest in a simulated red-tide environment. Thus, it can be surmised that copepods may selectively graze on C. antiqua during the outbreak of a red tide. Grazing pressure by the natural copepod community in Harima Nada, the Inland Sea of Japan, was calculated by integration of the laboratory determined feeding rates and field measurements of zooplankton biomass. The daily removal rate was 3.4 to 30.8% (mean: 12.3%) of C. antiqua biomass at 20 cells ml^-1 and decreased to 0.6–4.3% (mean: 1.8%) at 500 cells ml^-1. Therefore, the grazing pressure by the copepod community is important at the initial stage of the red tide.     

Effects of Widespread Fish Introductions on Paedomorphic Newts in Europe

Abstract:  As a result of factors such as global warming, habitat destruction, and species introduction, amphibians are declining worldwide. No one, however, has analyzed the status of polymorphic amphibian species at a national or continental scale, although some local reports exist. Our aim was to report on the loss of intraspecific heterochrony as a loss to diversity in determining the consequences of fish stocking on European populations of paedomorphic newts. Paedomorphosis is a polymorphism in which larval traits are retained in the adult stage. We surveyed 39 paedomorphic populations of the alpine (  Triturus alpestris ) and palmate ( T. helveticus ) newts, all but one of which initially occupied fishless ponds and lakes in France, Italy, Slovenia, Bosnia, Montenegro, and Greece. Exotic fishes were found in 44% of the studied aquatic habitats, with a 100% presence in Montenegro. At all sites paedomorphs disappeared and metamorphs declined. Only fish explained these population changes because alternative factors such as drying were not significant. More catastrophically, fish introductions occurred in habitats known to support the largest populations of newts and even some endemic subspecies. If management and legislative measures are not taken to stop fish stocking, protect paedomorphs as conservation units at national and international levels, and restore natural habitats, all the largest paedomorphic populations may disappear in the near future. Their disappearance would represent a loss of one of the rare, fascinating examples of intraspecific heterochrony.     

Temporal patterns in diversity and species composition of deep-sea benthic copepods in bathyal Sagami Bay,central Japan

Little is known about temporal changes in the diversity and species composition of deep-sea metazoan meiofauna and their relationships with changes in the food supply. Those changes were studied for benthic copepod assemblages based on 2-year time-series data at a bathyal site in Sagami Bay (1430 m depth), central Japan, where annual fluctuation in the abundance of benthic foraminiferans was previously observed. Species diversity of benthic copepods at the site was as high as, or slightly higher than, that observed at other deep-sea sites, but did not fluctuate temporally through the study period. Multivariate analyses did not reveal any clear seasonal or directional change occurring over the longer term in their species composition, although there was some consistent pattern. These results indicate a lack of, or only weak, seasonality in the diversity and species structure of the deep-sea benthic copepod assemblages, even though the fresh organic food supply fluctuates seasonally. They also suggest that there are differences between copepods and foraminiferans in the response to changes in environmental factors, and that spatial differences in the composition of copepod communities are greater than temporal ones at this deep-sea site.Communicated by T. Ikeda, Hakodate     

An Evaluation of Restoration Efforts in Fishless Lakes Stocked with Exotic Trout

Abstract: Detrimental effects of introduced fishes on native amphibian populations have prompted removal of introduced cutthroat ( Oncorhynchus clarki ), rainbow ( Oncorhynchus mykiss ), and brook trout (  Salvelinus fontinalis ) from naturally fishless lakes at Mt. Rainier National Park, Washington ( U.S.A.). Using paleolimnological indicators (diatoms, invertebrates, and sediment characteristics) in eight 480-year-old sediment cores from eight lakes, we (1) derived estimates of baseline environmental conditions and natural variation, (2) assessed the effects of stocking naturally fishless lakes, and (3) determined whether lakes returned to predisturbance conditions after fish removal (restoration). Diatom floras were relatively stable between 315 and 90 years before present in all lakes; we used this time period to define lake-specific "baseline" conditions. Dissimilarity analyses of diatoms revealed sustained, dramatic changes in diatom floras that occurred approximately 80 years ago (when fish were introduced) in four of five stocked lakes, whereas the diatom floras in two unstocked lakes had not changed significantly in the last 315 years. Diatoms were not preserved in an eighth lake. State changes also occurred in two lakes over 200 years before European settlement of the Pacific Northwest. Preserved invertebrate densities fluctuated dramatically over time in all cores, providing a poor reference for assessing the effects of fishes. Nevertheless, fish-invertebrate interactions have been demonstrated in other paleolimnological studies and may be useful for lower-elevation or more productive lakes. Because diatom communities have not returned to predisturbance assemblages in restored lakes, even 20–30 years after fish removal, we conclude that Mt. Rainier lakes were not successfully restored by the removal of fishes.     

Experimental Demonstration of an Allee Effect in American Ginseng

Abstract: Harvesting of wild American ginseng (   Panax quinquefolius ) for the herbal trade has lowered natural population sizes. We tested for reproductive limitation due to small population size (a form of the Allee effect) by experimentally planting "natural" populations numbering 4, 16, and 64 using 4-year-old cultivated plants. Plant size traits and reproductive traits ( bud, flower, green fruit, and mature fruit) were recorded through the ensuing summer. Fruit production per flower and per plant increased in proportion to flowering population size (  p = 0.0063 and p = 0.0017, respectively), strongly suggesting that an Allee effect occurs in very small populations. The increase in fruit production was not explained by either plant or inflorescence size differences. Although population size-dependent pollination, through insufficient pollinator visitation rate or pollen transfer rate, seems the most likely cause of the observed effects, our limited observations of pollinators were not sufficient to demonstrate a change in pollination rates as a function of population size. Knowledge of the presence as well as the mechanism underlying this Allee effect may be especially useful for management and determination of minimum viable population size of the species in the wild.     

Environmental stability and seasonality of a harpacticoid copepod community

The southwest monsoon on the west coast of India brings about dynamic changes in estuaries and coastal waters. The response of the meiofauna to monsoonal rain is obvious, but the impact of such environmental changes on the community structure of harpacticoid copepod species and their seasonality in the estuaries influenced by the tropical monsoon is poorly understood. In this study the spatial and temporal variability in abundance and community structure of meiobenthic copepods was investigated over an annual cycle (June 1983 to June 1984), in an estuary influenced by the tropical monsoon. Total meiofaunal abundance showed wide variations in space and time. Minimum and maximum densities were observed in the monsoon and pre-monsoon seasons, respectively. Quick recovery of harpacticoid populations in the early post-monsoon season indicated their recuperative power under adverse conditions. Of the 25 species recorded from lower, middle and upper reaches, eight comprised over 70% of the total copepod population. The peak and low occurrence of dominant species displayed striking correlations with the summer and rainy seasons, respectively. While other species were restricted in distribution, Stenhelia longifurca was recorded from all salinity regimes. The harpacticoid community was greatly influenced by the onset of the monsoon period, and their spatial and temporal variabilities were related with physico-chemical parameters and the variability of these parameters in the estuary.     

A field colonization experiment with meiofauna and seagrass mimics: effect of time,distance and leaf surface area

From a conservation point of view, it is essential to know how fast an ecosystem can recover after physical disturbance. Meiofauna and especially harpacticoid copepods are abundant in seagrass beds and are therefore useful to study ecosystem recovery after disturbance. In the western Caribbean coast, a fragmented Thalassia testudinum seagrass bed was selected to conduct a colonization field experiment by means of plastic seagrass mimics. Meiofauna colonization, with special emphasis on harpacticoid copepods, was followed in relation to: (1) colonization time (2, 4, 6, 10, 14 and 21 days); (2) distance to source of colonizers (close and far series) and (3) leaf surface area to colonize (small, medium, large). Colonization was recorded after 2 days with average meiofauna densities of 480 ind/100 cm² (close) and 1350 ind/100 cm² (far) of leaf surface area, while on average 400 ind/100 cm² were collected from the natural seagrass plants. In this early phase, the meiofauna diversity was high, with on average 8 taxa. A longer period of colonization (21 days) showed an increased meiofaunal density and diversity (average density: 3220 ind/100 cm², 13 taxa). Increasing meiofauna colonization with time is probably related to the development of a biofilm making the leaf more attractive for meiofauna. The effect of distance was not so pronounced as that of time. Total absolute densities were highest in the far series (5 m away from natural seagrass patch), mainly because of nematode densities. Meiofauna diversity was lower in the far series than in the close series (at the border of the natural seagrass patch). A larger individual leaf surface area did not affect the overall meiofauna densities but had a significant positive effect on copepod densities. Larger surface areas promoted the presence of epiphytic copepod families such as Tegastidae and Dactylopusiidae. Overall, we found a rapid recovery of meiofauna in fragmented seagrass beds with primary colonizers (both nematodes and benthic opportunistic copepods) originating from the sediment and later colonizers as epiphytic copepods and their nauplii from the local seagrass regeneration pool.     

Meiofauna on the seagrass Thalassia testudinum: population characteristics of harpacticoid copepods and associations with algal epiphytes

The composition and abundance of bladedwelling meiofauna was determined over a 15 mo period (1983–1984) from a Thalassia testudinum Banks ex König meadow near Egmont Key, Florida, USA. Harpacticoid copepods, copepod nauplii, and nematodes were the most abundant meiofaunal taxa on T. testudinum blades. Temporal patterns in species composition and population life-history stages were determined for harpacticoid copepods, the numerically predominant taxon. Sixteen species or species complexes of harpacticoid copepods were identified. Harpacticus sp., the most abundant harpacticoid, comprised 47.8% of the total copepods collected, and was present throughout the study. Copepodites dominated the population structures of the blade-dwelling harpacticoid species on most collection dates. Ovigerous females and/or copepodites were always present, indicating continuous reproductive activity. Results suggest that epiphytic algae influence meiofaunal abundance on seagrass blades, as densities of most meiofaunal taxa at Egmont Key were positively associated with percent cover of epiphytic algae throughout the study. The majority of significant correlations between meiofaunal density and cover of epiphytic algae involved filamentous algae, although encrusting algae dominated the epiphytic community. It appears that resources provided by epiphytic algae to seagrass meiofauna (additional food, habitat, and/or shelter from predation) may be associated with algal morphology.