Ecology of a corallivorous gastropod,<Emphasis Type="Italic"> Coralliophila abbreviata</Emphasis>, on two scleractinian hosts. I: Population structure of snails and corals

Despite their potential importance in structuring reef communities, invertebrate corallivores and their population structures are poorly understood. We found distinct differences in the population structures (length-frequency distribution and sex ratio) of the corallivorous gastropod Coralliophila abbreviata residing on two coral-host taxa, Montastraea spp. and Acropora palmata, in the Florida Keys. In each of two survey years, around 50% of the Montastraea spp. colonies were infested, with a mean snail density of eight snails per infested colony (range 1–45), while around 20% of A. palmata colonies harbored three snails per infested colony (range 1–23). Variation in patterns of snail occurrence was also observed within a host taxon. A. palmata occurred in low- and high-density stands (0.4 and 1 colony m^–2, respectively, at the initial survey) at different sites. Hurricane Georges struck the area in September 1998. When resurveyed in 1999, density of colonies in low-density stands had decreased by 75% to 0.1 colonies m^–2. This decrease was accompanied by a doubling in the proportion of colonies infested with snails (from 19% to 46%) and an increase in snail density per infested colony (from 3.7±3.3 SD to 5.4±4.6 SD) as snails apparently concentrated on surviving A. palmata. In contrast, sites with high density A. palmata stands (thickets) retained colony densities of about ~1 colony m^–2 among years, while snail infestation increased only from 9% to 14% of colonies surveyed and snail density essentially remained unchanged (from 2.7±1.8 to 2.9±1.9 snails per infested colony). Snails collected from Montastraea spp. were shorter than those from A. palmata in low-density stands and were longest on A. palmata in thickets. On both host taxa, female snails were longer than males. The sex ratio of snails on Montastraea spp. hosts was even (1:1), while that of snails on A. palmata was skewed (70% males). Factors that could explain observed differences in size structure and sex ratio between Coralliophila populations on the two coral host taxa include: differential susceptibility to predators, influence of host tissue nutritional quality and/or secondary metabolite content, and genetic differences (cryptic species). The host-specific characteristics of C. abbreviata populations imply that the impact of gastropods on reef communities will vary with the coral species composition.     

Shifts in coral community structures following cyclone and red tide disturbances within the Gulf of Oman (United Arab Emirates)

This study documents the effects of two consecutive disturbances on coral community structures in the Gulf of Oman (United Arab Emirates); Cyclone Gonu in June 2007 and the Cochlodinium polykrikoides harmful algal bloom (HAB) that persisted from August 2008 until May 2009. Coral cover, colony densities, size class frequency distributions, and geometric growth rates derived from size class transition probability matrices were used to assess the post-Gonu and post-HAB recovery trajectories at four sites. The net effects of these disturbances were fourfold: (i) storm damage caused >50% losses of live branching and tabular coral cover by fragmentation and dislodgment of pocilloporid and acroporid colonies; (ii) Pocillopora damicornis colonies that survived the cyclone experienced mass mortality during the first 3?months of the HAB, resulting in localized extirpation of this species; (iii) variable Acropora mortality during the HAB indicated individual colony, rather than taxa-wide, susceptibility; and (iv) massive colony coral taxa were resistant to both disturbances.     

Fine-scale patchiness and genetic heterogeneity of recruits of the corallivorous gastropod Drupella cornus

The size-frequency distributions and genetic composition of recruits of the corallivorous snail Drupella cornus (Röding, 1798) were examined in outbreak populations collected from Ningaloo Reef, Western Australia in August 1990. The recruits are found in groups on digitate Acropora spp. corals. Among coral colonies, mean lengths of recruits in our samples ranged from <9 to 22 mm, but the snails within a group were generally similar in size. Despite the fact that D. cornus has planktonic larvae, there were marked genetic differences between groups of recruits on different coral colonies. The relatively large genetic subdivision among groups of recruits within sites over distances <80 m was measured as a value of F _ST (standardized variance in allelic frequencies) of 0.044. This was three times the value from comparisons of pooled samples of recruits from areas up to 119 km apart, and six times as great at the genetic subdivision among populations of adults over a distance of 180 km. Much of the genetic heterogeneity among groups of recruits is associated with mean size of the snails. Taken together, the size-frequency distributions and the genetic differences indicate that recruits within the same coral colony shared a common history of settlement, suggesting a cohesiveness of groups of larvae. Although the mechanisms for this patchiness are not understood, one implication is that studies of size-frequency distributions and genetic composition of cohorts of D. cornus must treat the group, not the individual snail, as the unit of replication.     

A dynamic size-structured population model: does disturbance control size structure of a population of the massive coral Gardineroseris planulata in the Eastern Pacific?

Using a long-term data set of Gardineroseris planulata (Dana) on Uva Island reef, Panamá, we developed a simulation model that relates size-specific schedules of growth and partial mortality to predation by Acanthaster planci (Linnaeus) and El Niño–Southern Oscillation (ENSO)-related elevation of water temperature. We compared the model predictions to field observations for both the subpopulation of colonies that was used for model development and for the entire population. No statistically significant differences in the size-frequency distributions of the real and modeled coral populations were found for the subpopulation during any of 9 yr or for the entire population during 4 yr. These results suggested that the model relationships were reflecting field conditions. Longer-term (100 yr) simulations were conducted to assess the relative importance of predation and ENSO-related colony losses in determining the size structure of the coral population. Predation by A. planci was of overwhelming importance due to both stronger effects of predation (larger transitions) and the frequency of predation (yearly) compared to ENSO (episodically). Even the least-frequent predation scenario skewed the distribution toward smaller colonies, while simulations where populations were subjected to frequent ENSO events (≥3 yr) still maintained colonies in the largest size class. The model results suggested that sea star predators may not have been in the present abundance on this reef prior to the last 30 yr; with predators present, the model predicts that the distribution would be skewed toward smaller colonies.     

Purification of multiple vitellogenins in grey mullet (<Emphasis Type="Italic">Mugil cephalus</Emphasis>)

Coralliophila abbreviata (Lamarck) is a corallivorous gastropod that lives and feeds on several species of scleractinian coral in the Western Atlantic and Caribbean. Previous studies of C. abbreviata have revealed that snails on branching acroporid corals are larger and consume more tissue than those on massive and plating corals. To ascertain whether snail life-history and fitness are differentially affected by the coral host, an analysis of the age structure and female reproductive output of snail populations on three coral host taxa (Acropora palmata, Diploria spp., and Montastraea spp.) was conducted at four shallow (2–7 m depth) reef sites off Key Largo, Florida in June through August, 2004. Snails were, on average, almost twice as large on A. palmata than on Diploria spp. and Montastraea spp., averaging 30.3 mm shell length, compared to 17.2 and 17.6 mm, respectively. Brood size increased as a power function with female shell length. Females on A. palmata were significantly larger than females on the other two hosts and, therefore, produced more offspring per female. The number of growth striae on the inner surface of the operculum was used to estimate snail age. Estimates of growth rate were obtained by fitting the Gompertz growth function to size-at-age plots and mortality was estimated using growth parameters and size-frequency data. The data suggest that C. abbreviata inhabiting A. palmata are larger than on alternative hosts due to a combination of a faster growth rate and longer life-span. The species is believed to be a protandrous hermaphrodite. The timing of sex change varied among hosts; snails on A. palmata changed sex later at larger sizes relative to those on the other two hosts. Based on these results, it seems probable that C. abbreviata has developed reaction norms for life-history traits, allowing snails to adjust and maximize fitness in the different environments associated with various coral hosts.     

Multi-year, seasonal genotypic surveys of coral-algal symbioses reveal prevalent stability or post-bleaching reversion

This report documents the extent to which coral colonies show fluctuations in their associations with different endosymbiotic dinoflagellates. The genetic identity of Symbiodinium from six coral species [Acropora palmata (Lamarck), A. cervicornis (Lamarck), Siderastrea siderea (Ellis and Solander), Montastrea faveolata (Ellis and Solander), M. annularis (Ellis and Solander), and M. franksi (Gregory)] was examined seasonally over five years (1998 and 2000–2004) in the Bahamas and Florida Keys at shallow (1 to 4 m) fore-reef/patch reef sites and at deeper fore-reef (12–15 m) locations. Symbionts were identified genetically using denaturing gradient gel electrophoresis (DGGE) fingerprinting of the internal transcribed spacer region 2 (ITS2) of ribosomal RNA gene loci. Repetitive sampling from most labeled colonies from the Bahamas and the Florida Keys showed little to no change in their dominant symbiont. In contrast, certain colonies of M. annularis and M. franksi from the Florida Keys exhibited shifts in their associations attributed to recovery from the stresses of the 1997–1998 El Niño southern oscillation (ENSO) event. Over several years, a putatively stress-tolerant clade D type of Symbiodinium was progressively replaced in these colonies by symbionts typically found in M. annularis and M. franksi in Florida and at other Caribbean locations. Greater environmental fluctuations in Florida may explain the observed changes among some of the symbioses. Furthermore, symbiotic associations were more heterogeneous at shallow sites, relative to deep sites. The exposure to greater environmental variability near the surface may explain the higher symbiont diversity found within and between host colonies.     

The effects of partial mortality on the fecundity of three common Caribbean corals

The recent intensification of human disturbances in the Caribbean has increased the prevalence of partial mortality on coral colonies. Partial mortality can change colony size by directly shrinking colonies or by splitting colonies into fragments. A reduction in colony size can also adversely affect fecundity and fitness as internal resources shift away from reproduction toward colony maintenance. This study aimed to determine whether three Caribbean coral species, Siderastrea siderea, Montastraea faveolata, and Diploria strigosa, along the reef tract in Puerto Morelos, Mexico (20^o52′N, 86^o51′W), continued to dedicate resources to reproduction when colonies were fragmented to pre-maturation size. Contrary to expectations, eggs were found in colonies that were smaller than the maturation size and had been subjected to partial mortality. The continued dedication of resources toward reproduction, even in the smallest colonies, suggests that resource trade-offs away from reproduction are not as rigid as previously suggested in stressed corals.     

Survey of deep-dwelling red coral (<Emphasis Type="Italic">Corallium rubrum</Emphasis>) populations at Cap de Creus (NW Mediterranean)

The distribution and population structure of the eurybathic gorgonian Corallium rubrum were studied off Cap de Creus (Costa Brava, Northwestern Mediterranean Sea). Red coral is endemic to the Mediterranean Sea and the adjacent NE Atlantic coast, where it has been over exploited for centuries. This study presents, the first quantitative data on the spatial distribution and structure of a population extending between 50 (common SCUBA limits) and 230 m depth, and compared it with shallow populations previously studied in the same area. Different remotely operated vehicles (ROV) and two methodological approaches were employed during four cruises between 2002 and 2006: 1-Extensive surveys: sea to coast transects in which red coral density and patch frequency were recorded; 2-Intensive surveys, in which parameters describing colony morphology were recorded. Most of the hard substrate between 50 and 85 m depth was inhabited by red coral colonies, showing a patch frequency of 8.3 ± 7.9 SD patches per 100 m-transect (total transect area: 34 m²), and within-patch colony densities of 16–376 colonies m⁻² (mean of 43 ± 53 colonies m⁻²). Below 120 m depth red coral was less abundant, and rather than forming dense patches as in shallow water, isolated colonies were more common. The population structure differed between sites that are easily accessible to red coral fishermen, and remote ones (both at similar depth, 60–80 m), as colonies in easily accessible locations were smaller in height and diameter, and showed a less developed branching pattern. At shallower locations (10–50 m depth) the population structure was significantly different from those at deeper locations, due to the heavy harvesting pressure they are exposed to in the shallows. Twenty-five to forty-six percentage of the deeper colonies were taller than 6 cm, while only 7–16% of the shallow water colonies exceeded 6 cm colony height. Forty-six to seventy-nine percentage of the colonies in deeper waters were large enough to be legally harvested, while only 9–20% of the shallow water colonies met the 7 mm legal basal diameter to be collected. The branching pattern was also better developed in deeper colonies, as up to 16% of the colonies showed fourth order branches, compared to less than 1% of the shallow water colonies (of which 96% consisted of only one single branch). The results thus confirm that C. rubrum populations above 50 m depth are exposed to a higher harvesting intensity than deeper populations in the same area.     

Assessing the effects of disease and bleaching on Florida Keys corals by fitting population models to data

Coral diseases have increased in frequency over the past few decades and have important influences on the structure and composition of coral reef communities. However, there is limited information on the etiologies of many coral diseases, and pathways through which coral diseases are acquired and transmitted are still in question. Furthermore, it is difficult to assess the impacts of disease on coral populations because outbreaks often co-occur with temperature-induced bleaching and anthropogenic stressors. We developed spatially explicit population models of coral disease and bleaching dynamics to quantify the impact of six common diseases on Florida Keys corals, including aspergillosis, dark spots, white band, white plague, white patch, and Caribbean yellow band. Models were fit to an 8-year data set of coral abundance, disease prevalence, and bleaching prevalence. Model selection was used to assess alternative pathways for disease transmission, and the influence of environmental stressors, including sea temperature and human population density, on disease prevalence and coral mortality. Classic disease transmission from contagious to susceptible colonies provided the best-fit model only for aspergillosis. For other diseases, external disease forcing, such as through a vector or directly from pathogens in the environment, provided the best fit to observed data. Estimates of disease reproductive ratio values (R₀) were less than one for each disease, indicating coral colonies were below densities required for diseases to become established through contagious spread alone. Incidences of white band and white patch disease were associated with greater susceptibility or slower recovery of bleached colonies, and no disease outbreaks were associated with periods of elevated sea temperatures alone. Projections of best-fit models indicated that, atleast during the period of this study, disease and bleaching did not have substantial impacts on populations and impaired rates of population growth appeared to be attributable to other stressors. By applying epidemiological models to field data, our study gives qualitative insights into the dynamics of coral diseases, relative stressor impacts, and directions for future research.     

Polymorphism in soft coral larvae revealed by amplified fragment-length polymorphism (AFLP) markers

The dioecious Red Sea soft coral Parerythropodium fulvum fulvum breeds its nonsymbiotic planula larvae on the surface of female colonies for less than a week. After completing their development, larvae crawl and settle near maternal colonies. Here we study the genetic polymorphism of developing larvae by the use of amplified fragment-length polymorphism markers. Four reproductive colonies from shallow water populations (two from a dense population and two from a less densely populated area 100 m away) were chosen, and ten larvae were randomly collected from each colony. DNA was analyzed by using three different primer combinations producing 61, 63, 63 polymorphic markers, respectively. All larvae exhibited different banding patterns from one another, illustrating the prominent role of sexual reproduction for the production of larvae. Nei's mean genetic distances for all 12 possible pair-wise combinations for larval origins revealed, in most cases, that sister larvae are genetically closer than larvae from different colonies and that larvae may be grouped into three statistical clusters in accordance with colony origin and population studied. The usefulness of molecular methodologies in coral population genetics is discussed. Received: 26 March 1999 / Accepted: 19 October 1999     

Influences of habitat and natural disturbances on contributions of massive Porites corals to reef communities

We compared densities, distributions and size frequencies of massive corals in the genus Porites on five relatively exposed, mid-shelf reefs (50 km offshore) in the central Great Barrier Reef with those on a sheltered inshore reef (10 km offshore). Data included various transect and mapping studies between 1984 and 1990, estimates of size-dependent damage from the crown-of-thorns starfish Acanthaster planci, estimated densities of herbivorous sea urchins (potential predators of juveniles), and observations of size-specific effects of tropical cyclones. Assemblages of Porites spp. on mid-shelf reefs were dominated by small colonies (2 to 10 cm diam) established either from planula larvae or from small tissue remnants that had survived A. planci predation in the early to mid-1980s. Large colonies (up to 10 m diam) were scarce, except for localized aggregations on terraces at the base of reef slopes (6 to 12 m deep). Extensive space suitable for settlement by coral larvae can be attributed to recurrent cyclones and A. planci outbreaks. Despite low sea urchin predation, the slowly growing Porites juveniles are likely to die from overgrowth by numerous, much faster growing corals. On the sheltered inshore reef, the coral community was dominated by very large (>5 m diam) Porites colonies, several centuries old; recruitment was mainly by fragmentation of large colonies; there was little space available for settlement, and probabilities of juvenile mortality from grazing urchins were high. Differences in settlement and early survival of Porites spp. are exacerbated by different regimes of storm damage. A model is proposed that links wave climate with the size and age reached by corals before dislodgement by storm waves, and which is consistent with observed densities and size-frequency distributions of Porites in sheltered and exposed areas.     

Neoplasia,regeneration and growth in the reef-building coral Acropora palmata

Abnormal processes of calcification, such as regenerating lesions and neoplasia, situated near the tips(<25 cm) of colonies of Acropora palmata (Lamarck) suppressed normal linear growth. Branches having neoplasia at a larger distance from the tip do not grow significantly differently from controls. This indicates a functional minimal area in terms of energy supply. Neoplasia are pure aragonite and have the same coenosteal structure as regenerative skeletal material. Regeneration of tissue as well as tissue+skeleton lesions involves the simultaneous formation of tissue and regenerative skeleton, trapping foreign material under the regenerated surface. Recovery of a damaged surface slows down with time and this may, in other coral species, result in permanent lesions. A. palmata recovered from all lesions (n=32) within 80 d and appears to be a superior regenerator among Caribbean corals. This is consistent with other life-history characteristics of this highly specialized coral species.     

Ecology of a corallivorous gastropod,<Emphasis Type="Italic"> Coralliophila abbreviata</Emphasis>, on two scleractinian hosts. II. Feeding,respiration and growth

Coralliophila abbreviata is a corallivorous gastropod that has been observed to cause large feeding scars on reef-building corals on Floridian and Caribbean reefs. We detected differences in the population structure (length-frequency distribution and sex ratios) of C. abbreviata populations living on two coral host taxa, Acropora palmata and Montastraea spp., in the Florida Keys in a previous study. We hypothesize that diet (host) has a major influence on snail population structure and, thus, we characterize metabolism, feeding and growth for snails residing on these coral taxa. Here, we present results of a reciprocal transplant experiment demonstrating that the taxon of the host influences snail growth rates, as indicated by changes in shell and body tissue weight. Regardless of the host from which they were drawn, snails resident on A. palmata grew faster than those resident on Montastraea spp. Thus, diet influences snail population structure. However, the tissue of Montastraea spp. provides more N and C per area of tissue than that of A. palmata. Respiration rates and tissue composition of snails collected from the two host taxa did not differ. Therefore, snails feeding on Montastraea spp. should have to consume less tissue per day to satisfy their metabolic requirements compared to snails feeding on A. palmata. Feeding rates for snails on A. palmata were measured in the laboratory over 48 h (1–9 cm² coral tissue snail⁻¹ day⁻¹) and estimated from feeding scars observed in the field (weekly mean rate of 1.07 cm² coral tissue snail⁻¹ day⁻¹). The lack of definition of snail feeding scars on Montastraea spp. required the calculation of coral tissue consumption rates based on estimated minimum carbon requirements. Calculated feeding rates for C. abbreviata were 0.13–0.88 cm² coral tissue day⁻¹ snail⁻¹, when feeding on Montastraea spp., and 0.44–3.28 cm² coral tissue day⁻¹ snail⁻¹, when feeding on A. palmata. The calculated range for the latter is consistent with measured rates. Thus, C. abbreviata exhibits high variation in growth parameters in response to environmental variability and/or food source. At mean levels of snail density on reefs off Key Largo, Fla., 20% of A. palmata colonies lose between 1.32 and 9.84 cm² tissue day⁻¹, while 50% of Montastraea spp. colonies lose between 1.04 and 7.04 cm² tissue day⁻¹. Together with published coral tissue regeneration rates, these results suggest that if sustained, such rates of predation could have a serious effect on the viability of these coral populations on Florida's reefs.     

Assessing the effectiveness of marine reserves on unsustainably harvested long-lived sessile invertebrates

Although the rapid recovery of fishes after establishment of a marine reserve is well known, much less is known about the response of long-lived, sessile, benthic organisms to establishment of such reserves. Since antiquity, Mediterranean red coral (Corallium rubrum) has been harvested intensively for use in jewelry, and its distribution is currently smaller than its historical size throughout the Mediterranean Sea. To assess whether establishment of marine reserves is associated with a change in the size and number of red coral colonies that historically were not harvested sustainably, we analyzed temporal changes in mean colony diameter and density from 1992 to 2005 within red coral populations at different study sites in the Medes Islands Marine Reserve (established in 1992) and in adjacent unprotected areas. Moreover, we compared colony size in the Medes Islands Marine Reserve, where recreational diving is allowed and poaching has been observed after reserve establishment, with colony size in three other marine protected areas (Banyuls, Carry-le-Rouet, and Scandola) with the enforced prohibition of fishing and diving. At the end of the study, the size of red coral colonies at all sampling sites in the Medes Islands was significantly smaller than predicted by growth models and smaller than those in marine protected areas without fishing and diving. The annual number of recreational dives and the percent change in the basal diameter of red coral colonies were negatively correlated, which suggests that abrasion by divers may increase the mortality rates of the largest red coral colonies within this reserve . Our study is the first quantitative assessment of a poaching event, which was detected during our monitoring in 2002, inside the marine reserve. Poaching was associated with a loss of approximately 60% of the biomass of red coral colonies.     

Unprecedented bleaching-induced mortality in Porites spp. at Rangiroa Atoll, French Polynesia

In April-May 1998, mass coral bleaching was observed in the lagoon of Rangiroa Atoll, Tuamotu Archipelago, French Polynesia. Six months later, the extent of bleaching-induced coral mortality was assessed at three sites. Corals in the fast-growing genus Pocillopora had experienced >99% mortality. Many large colonies of the slow-growing genus Porites (mean horizontal cross-sectional area 5.8 m²) had also died - a phenomenon not previously observed in French Polynesia and virtually unprecedented world-wide. At one site, 25% of colonies, or 44% of the pre-bleaching cover of living Porites, experienced whole-colony mortality. At the two other sites, recently dead Porites accounted for 41% and 82% of the pre-bleaching live cover. Mortality in Porites was negatively correlated with depth between 1.5 and 5 m. Using a 50-year dataset of mean monthly sea surface temperature (SST), derived from ship- and satellite-borne instruments, we show that bleaching occurred during a period of exceptionally high summer SST. 1998 was the first year in which mean monthly SSTs exceeded the 1961-1990 upper 95% confidence limit (29.4°C) for a period of three consecutive months. We suggest that the sustained 3-month anomaly in local summer SST was a major cause of coral mortality, but do not discount the synergistic effect of solar radiation. Recovery of the size-frequency distribution of Porites colonies to pre-bleaching levels may take at least 100 years.     

Predators selectively graze reproductive structures in a clonal marine organism

Although the fitness consequences of herbivory on terrestrial plants have been extensively studied, considerably less is known about how partial predation impacts the fitness of clonal marine organisms. The trophic role of Caribbean parrotfish on coral reefs is complex: while these fish are important herbivores, as corallivores (consumers of live coral tissue), they selectively graze specific species and colonies of reef-building corals. Though the benefits of parrotfish herbivory for reef resilience and conservation are well documented, the negative consequences of parrotfish grazing for coral reproductive fitness have not been previously determined. We examined recently grazed colonies of Montastraea annularis corals to determine whether grazing was positively associated with coral reproductive effort. We measured gonad number, egg number and size, and proportional reproductive allocation for grazed and intact coral colonies 2–5 days prior to their annual spawning time. We found that parrotfish selectively grazed coral polyps with high total reproductive effort (number of gonads), providing the first evidence that parrotfish selectively target specific tissue areas within a single coral colony. The removal of polyps with high reproductive effort has direct adverse affects on coral fitness, with additional indirect implications for colony growth and survival. We conclude that chronic grazing by parrotfishes has negative fitness consequences for reef-building corals, and by extension, reef ecosystems.     

Bi-directional sex change in a coral-dwelling goby

Bi-directional sex change has recently been reported among obligate coral-dwelling gobies of the genus Gobiodon. However, neither the functional role of this pattern of sex change nor the frequency of sex change in either direction in natural populations is known. We investigated the social structure and pattern of sex change of Gobiodon histrio at Lizard Island on the Great Barrier Reef. The social structure of G. histrio within coral colonies usually consisted of a single juvenile or a heterosexual adult pair. The size of adult social groups was not constrained by coral colony size. In contrast to expectations for pair-forming species, G.␣histrio was primarily a protogynous hermaphrodite. All immature G. histrio were females and sex change from female to male occurred readily when two mature females were placed in a coral colony. In addition, male G. histrio were able to change back to females when two mature males were placed in a coral. Sex change from female to male, however, occurred with over twice the frequency of sex change from male to female. Where two males were placed in a coral colony, heterosexual pairs were most frequently re-established by immigration of females from outside the treatment population. This pattern might be predicted if sex change from male to female is more expensive than sex change from female to male for G. histrio. Where sex change is expensive, movement may be favoured over sex change, particularly where coral densities are high and movement among corals incurs little mortality risk. Received: 10 November 1997 / Accepted after revision: 16 May 1998     

Dispersal behaviour and colony structure in a colonial spider

Group living in spiders is characterised by two principle modes, the cooperative social mode and the colonial non-cooperative mode. Kin-relationships due to reduced dispersal determine population genetic structure in social spiders, but the dispersal mechanisms underlying group structure remain poorly understood in colonial spiders. Assuming similar ecological benefits of group living, we address the question whether reduced dispersal shapes population structure in a colonial spider, Cyrtophora citricola (Araneidae). We analysed dispersal by studying settling decisions under semi-natural conditions in experimental trees with and without colonies, and in natural populations, we estimated dispersal and colony structure using population genetic analyses. The propensity to disperse decreased with increasing age in experimental colonies. Adult females did not disperse in the experiment. Sub-adult female spiders preferred trees with a colony to trees without a colony. Dispersal in third instar juveniles was influenced significantly by wind but not by the presence of a colony. Thus, we showed that being in a colony did not inhibit juvenile dispersal, but pre-mating females were philopatric. Genetic differentiation among colonies in natural populations was heterogeneous, colonies being either little or highly differentiated. The heterogeneous structure is likely caused by colony founding by one or a few females followed by dispersal among perennial colonies. Gene flow, however, was slightly male-biased. The experimental and indirect, genetic approaches combined showed that dispersal and the breeding system of C. citricola resemble that of solitary spiders, with juvenile dispersal occurring in both sexes, while the colonial distribution is maintained by female philopatry.     

Seasonal decline in intracolony genetic relatedness in eastern tent caterpillars: implications for social evolution

Summary Genetic relatedness in social insect colonies may vary spatially or temporally as a result of changes in colony membership due to immigration or to variation in patterns of maternity and paternity. We estimated relatedness for eastern tent caterpillars (Malacosoma americanum) in laboratory colonies derived from egg masses using multilocus genotypic data derived from electrophoresis. This estimate is compared with estimates obtained from colony samples taken in the field at four intervals spanning the larval developmental season. We found that average intracolony relatedness is close to 0.5 initially but declines through the developmental season due to colony merging, showing that caterpillars do not discriminate between siblings and nonsiblings in order to preserve colony family structure. Using the intracolony values together with relatedness values for higher levels of population structure, we estimated the effective mean number of simple families represented in single colonies through the season. The overall effective number of families per tent increased from one at the time of eclosion to 1.3 by the end of the season. Average intracolony relatedness remained relatively high despite the occurrence of colony merging, apparently as a result of the low density of tents on most trees, combined with high relatedness within the original colonies. Thus, high intracolony relatedness is maintained in M. americanum populations through the effects of adult dispersal, mating, and oviposition patterns, rather than through behavioral discrimination mechanisms of the larvae. These findings underscore the importance of considering the causes of temporal variation in genetic relatedness as well as the consequences for the indirect component of inclusive fitness. Correspondence to: J.T. Costa     

Estimating the Density of Honeybee Colonies across Their Natural Range to Fill the Gap in Pollinator Decline Censuses

Abstract: Although pollinator declines are a global biodiversity threat, the demography of the western honeybee (Apis mellifera) has not been considered by conservationists because it is biased by the activity of beekeepers. To fill this gap in pollinator decline censuses and to provide a broad picture of the current status of honeybees across their natural range, we used microsatellite genetic markers to estimate colony densities and genetic diversity at different locations in Europe, Africa, and central Asia that had different patterns of land use. Genetic diversity and colony densities were highest in South Africa and lowest in Northern Europe and were correlated with mean annual temperature. Confounding factors not related to climate, however, are also likely to influence genetic diversity and colony densities in honeybee populations. Land use showed a significantly negative influence over genetic diversity and the density of honeybee colonies over all sampling locations. In Europe honeybees sampled in nature reserves had genetic diversity and colony densities similar to those sampled in agricultural landscapes, which suggests that the former are not wild but may have come from managed hives. Other results also support this idea: putative wild bees were rare in our European samples, and the mean estimated density of honeybee colonies on the continent closely resembled the reported mean number of managed hives. Current densities of European honeybee populations are in the same range as those found in the adverse climatic conditions of the Kalahari and Saharan deserts, which suggests that beekeeping activities do not compensate for the loss of wild colonies. Our findings highlight the importance of reconsidering the conservation status of honeybees in Europe and of regarding beekeeping not only as a profitable business for producing honey, but also as an essential component of biodiversity conservation.