Search

In order to investigate the serendipitous find of a gastropod encrusted by the symbiont-bearing colonial coral Oculina patagonica, we examined several specimens of cnidarian-encrusted gastropods, ranging in age from the Pliocene to the Recent, and characterized in detail their sclerobiont cover. The results of our analysis suggest that gastropod shells can be encrusted by symbiont-bearing colonial corals at various times: (1) when the gastropod is alive; (2) when the shell is being used by a secondary inhabitant (e.g., hermit crabs or sipunculid worms); (3) when the shell is discarded but yet to be buried. The relationship between the symbiont-bearing coral and the inhabitant(s) of the encrusted shell is an example of facultative mutualism, i.e., it is non-obligate yet beneficial for both ends as the former obtains the capability to move, and the latter improves the resistance and resilience of its armor, thus obtaining extra protection from predators. Being able to move could prove particularly useful for a symbiont-bearing coral because, in addition to removing the risk of being smothered by sediment, it would also favor the photosynthetic activity of its algal endosymbionts by allowing the coral to be always clean of sedimentary particles. Although the resulting epibiotic association would be limited in size by the ability of either the gastropod or the secondary inhabitant of the shell to move at the seafloor, these small and easy-to-miss benthic islands might become the seeds that allow sessile carbonate producers such as hermatypic colonial corals to colonize unconsolidated substrates.

Despite the great abundance and diversity of molluscs, only a few have attained "model research organism" status. One of those species is the slipper snail Crepidula fornicata. Its embryos were first used for classical lineage tracing studies in the late 19th century, and over a 100 years later they were "re-discovered" by our labs and used for modern fate mapping, gene perturbation, in vivo imaging, transcriptomics, and the first application of CRISPR/Cas9-mediated genome editing among the Spiralia/Lophotrochozoa. Simultaneously, other labs made extensive examinations of taxonomy, phylogeny, ecology, life-history, mode of development, larval feeding behavior, and responses to the environment in members of the family Calyptraeidae, which includes the genus Crepidula. Recently, we developed tools, resources, and husbandry protocols for another, direct-developing species, Crepidula atrasolea. This species is an ideal "lab rat" among molluscs. Together these species will be valuable for probing the cellular and molecular mechanisms underlying molluscan biology and evolution.

Het muiltje Crepidula fornicata (L., 1758) is een in Europa geïmporteerde slak. Rond 1880 werd hij voor het eerst gesignaleerd in Essex, Engeland, vermoedelijk uit de VS meegekomen met Amerikaanse oesters Crassostrea virginica (Gmelin, 1791), die de overbeviste Europese oesters moesten aanvullen (Korringa, 1942). Dit artikel behandelt het spoor dat de schelp achterlaat op een andere schelp van eigen of andere soort.

Marine larvae that experience some sub-lethal stresses can show effects from those stresses after metamorphosis, even when they seem to recover from those stresses before metamorphosis. In this study we investigated the short and long-term effects of exposing the larvae of three calyptraeid gastropods (Crepidula fornicata, Crepidula onyx, and Crepipatella fecunda) to temporary reductions in salinity. Larvae of all three species showed slower larval growth rates, longer time to metamorphic competence, and substantial mortality after being stressed in seawater at salinities of 10, 15, and 20 for less than 48 h. larval tolerance to low salinities varied widely within and among species, but longer stresses at lower salinities were generally more harmful to larvae. However, larvae in nearly all experiments that were able to metamorphose survived and grew normally as juveniles; there were no documented "latent effects." For all three species, starving larvae in full-strength seawater was not as harmful as exposing larvae to low salinity stress, indicating that detrimental effects on larvae were caused by the salinity stress per se, rather than by an indirect effect of salinity stress on feeding. C. fornicata that were stressed with low salinity as juveniles were more tolerant of the stress than larvae: all stressed juveniles lived and showed reduced growth rates for no more than 3 days. Our data suggest that even though reduced salinity is clearly stressful to the larvae of these 3 gastropod species, metamorphosis seems to generally provide individuals with a fresh start. (C) 2010 Elsevier B.V. All rights reserved.

Marine larvae that experience some sub-lethal stresses can show effects from those stresses after metamorphosis, even when they seem to recover from those stresses before metamorphosis. In this study we investigated the short and long-term effects of exposing the larvae of three calyptraeid gastropods (Crepidula fornicata, Crepidula onyx, and Crepipatella fecunda) to temporary reductions in salinity. Larvae of all three species showed slower larval growth rates, longer time to metamorphic competence, and substantial mortality after being stressed in seawater at salinities of 10, 15, and 20 for less than 48 h. larval tolerance to low salinities varied widely within and among species, but longer stresses at lower salinities were generally more harmful to larvae. However, larvae in nearly all experiments that were able to metamorphose survived and grew normally as juveniles; there were no documented "latent effects." For all three species, starving larvae in full-strength seawater was not as harmful as exposing larvae to low salinity stress, indicating that detrimental effects on larvae were caused by the salinity stress per se, rather than by an indirect effect of salinity stress on feeding. C. fornicata that were stressed with low salinity as juveniles were more tolerant of the stress than larvae: all stressed juveniles lived and showed reduced growth rates for no more than 3 days. Our data suggest that even though reduced salinity is clearly stressful to the larvae of these 3 gastropod species, metamorphosis seems to generally provide individuals with a fresh start. (C) 2010 Elsevier B.V. All rights reserved.

Marine larvae that experience some sub-lethal stresses can show effects from those stresses after metamorphosis, even when they seem to recover from those stresses before metamorphosis. In this study we investigated the short and long-term effects of exposing the larvae of three calyptraeid gastropods (Crepidula fornicata, Crepidula onyx, and Crepipatella fecunda) to temporary reductions in salinity. Larvae of all three species showed slower larval growth rates, longer time to metamorphic competence, and substantial mortality after being stressed in seawater at salinities of 10, 15, and 20 for less than 48 h. larval tolerance to low salinities varied widely within and among species, but longer stresses at lower salinities were generally more harmful to larvae. However, larvae in nearly all experiments that were able to metamorphose survived and grew normally as juveniles; there were no documented "latent effects." For all three species, starving larvae in full-strength seawater was not as harmful as exposing larvae to low salinity stress, indicating that detrimental effects on larvae were caused by the salinity stress per se, rather than by an indirect effect of salinity stress on feeding. C. fornicata that were stressed with low salinity as juveniles were more tolerant of the stress than larvae: all stressed juveniles lived and showed reduced growth rates for no more than 3 days. Our data suggest that even though reduced salinity is clearly stressful to the larvae of these 3 gastropod species, metamorphosis seems to generally provide individuals with a fresh start. (C) 2010 Elsevier B.V. All rights reserved.

Calyptraeid gastropods are common fouling organisms in some areas, and have been introduced frequently into bays and ports in the northern hemisphere. In some cases, introduced calyptraeids have spread extensively, while several other calyptraeid introductions have persisted but failed to expand geographically. Calyptraeid species are often difficult to identify to species making it difficult to identify the origin of many introductions. Here we use DNA sequence data to confirm that a Mediterranean population of the spiny slipper snail Bostrycapulus, previously referred to as B. aculeatus and B. calyptraeformis, is in fact B. odites from the South Atlantic, and that this population probably originated in South America.

Calyptraeid gastropods are common fouling organisms in some areas, and have been introduced frequently into bays and ports in the northern hemisphere. In some cases, introduced calyptraeids have spread extensively, while several other calyptraeid introductions have persisted but failed to expand geographically. Calyptraeid species are often difficult to identify to species making it difficult to identify the origin of many introductions. Here we use DNA sequence data to confirm that a Mediterranean population of the spiny slipper snail Bostrycapulus, previously referred to as B. aculeatus and B. calyptraeformis, is in fact B. odites from the South Atlantic, and that this population probably originated in South America.

L'étude réalise un premier inventaire du peuplement halieutique et du rôle écologique d'un habitat original, : les vasières consolidées à Haploops. Haploops est un petit crustacé amphipode qui élabore des tubes souples et la concentration de ces tubes dans ces zones est telle que la pratique du chalutage y est très difficile. Situé dans la partie externe de la baie de la Vilaine, ce faciès couvre plus de 100 km2. Deux types de chaluts ont été adaptés à l'échantillonnage de ces fonds encombrés (chalut à perche String et chalut à panneaux avec grille ventrale) et utilisés au cours de trois campagnes du navire de recherche Gwen Drez. Une marée expérimentale sur le Marika, navire de pêche professionnel, a été réalisée pour apprécier l'intérêt commercial du secteur. Les principaux résultats montrent que, au sein de la même communauté des fonds côtiers vaso-sableux, des différences d'occurrence et d'abondance opposent le faciès à Haploops de son voisinage immédiat. Le benthos y est plus diversifié et marqué par l'abondance relative en crustacés décapodes, alors que les mollusques gastéropodes et céphalopodes y sont moins bien représentés. Quelques espèces commerciales y trouvent un habitat préférentiel, en particulier les roussettes, le tacaud, la dorade grise et l'étrille, ainsi que la coquille Saint-Jacques et, en faible abondance, la baudroie et le Saint-Pierre. Certaines espèces, au contraire, ne sont bien représentées qu'à l'extérieur du faciès, notamment les poissons plats (sole, plie, céteau), le merlu, le merlan et le bar. L'analyse de la fraction juvénile des principaux poissons montre que les vasières à Haploops jouent le rôle de nourricerie pour la petite roussette, la dorade grise, le Saint-Pierre, mais que la nourricerie de la baie de Vilaine ne se prolonge pas dans le faciès ; les juvéniles de merlu, merlan, bar, plie, sole et céteau demeurent en zone externe. Les fonds à Haploops constituent un habitat de moindre intérêt que les fonds de la baie de Vilaine

L'étude réalise un premier inventaire du peuplement halieutique et du rôle écologique d'un habitat original, : les vasières consolidées à Haploops. Haploops est un petit crustacé amphipode qui élabore des tubes souples et la concentration de ces tubes dans ces zones est telle que la pratique du chalutage y est très difficile. Situé dans la partie externe de la baie de la Vilaine, ce faciès couvre plus de 100 km2. Deux types de chaluts ont été adaptés à l'échantillonnage de ces fonds encombrés (chalut à perche String et chalut à panneaux avec grille ventrale) et utilisés au cours de trois campagnes du navire de recherche Gwen Drez. Une marée expérimentale sur le Marika, navire de pêche professionnel, a été réalisée pour apprécier l'intérêt commercial du secteur. Les principaux résultats montrent que, au sein de la même communauté des fonds côtiers vaso-sableux, des différences d'occurrence et d'abondance opposent le faciès à Haploops de son voisinage immédiat. Le benthos y est plus diversifié et marqué par l'abondance relative en crustacés décapodes, alors que les mollusques gastéropodes et céphalopodes y sont moins bien représentés. Quelques espèces commerciales y trouvent un habitat préférentiel, en particulier les roussettes, le tacaud, la dorade grise et l'étrille, ainsi que la coquille Saint-Jacques et, en faible abondance, la baudroie et le Saint-Pierre. Certaines espèces, au contraire, ne sont bien représentées qu'à l'extérieur du faciès, notamment les poissons plats (sole, plie, céteau), le merlu, le merlan et le bar. L'analyse de la fraction juvénile des principaux poissons montre que les vasières à Haploops jouent le rôle de nourricerie pour la petite roussette, la dorade grise, le Saint-Pierre, mais que la nourricerie de la baie de Vilaine ne se prolonge pas dans le faciès ; les juvéniles de merlu, merlan, bar, plie, sole et céteau demeurent en zone externe. Les fonds à Haploops constituent un habitat de moindre intérêt que les fonds de la baie de Vilaine

Reprinted from Journal of Morphology, vol. XIII, no. 1., From the introduction: The purpose of the following work from its inception has been to make as careful a study as possible of the cleavage of the ovum, the formation of the germinal layers and definitive organs, and the axial relations of the ovum to the larval and adult axes. At the time when this work was begun, several years ago, scarcely any attempts had been made to trace the history of individual blastomeres through the entire development to the formation of definitive organs. The early stages of cleavage had received a great deal of attention, but the later stages had been largely neglected; and although the origin and homology of the germ layers was perhaps the most frequently discussed subject in embryology, yet the relation of these layers to the individual blastomeres of the cleaving ovum had been determined in comparatively few cases. Since that time a number of very valuable papers have appeared on this subject of “cell lineage,” as Wilson (‘92) has aptly termed it. The results of such work are no longer as novel as they were four or five years ago, and yet the general interest in the subject has greatly increased, and that, too, in spite of the fact that there is a growing school of biologists who believe that individual blastomeres have no necessary relation to future organs. The subject of germ layers is no longer so important as it was once considered; in fact, the theory of the homology of the germinal layers has met with so many difficulties of late that it is now generally maintained only in a greatly modified form. However, the fundamental idea which was prominent in germ-layer discussions is of vital interest to-day. In the whole history of the germ-layer theories I see an attempt to trace homologies back to their earliest beginnings. This problem is as important to-day as it ever was, and whether one find these earliest homologies in layers or regions or blastomeres or the unsegmented ovum itself, the quest is essentially the same.

Reprinted from Journal of Morphology, vol. XIII, no. 1., From the introduction: The purpose of the following work from its inception has been to make as careful a study as possible of the cleavage of the ovum, the formation of the germinal layers and definitive organs, and the axial relations of the ovum to the larval and adult axes. At the time when this work was begun, several years ago, scarcely any attempts had been made to trace the history of individual blastomeres through the entire development to the formation of definitive organs. The early stages of cleavage had received a great deal of attention, but the later stages had been largely neglected; and although the origin and homology of the germ layers was perhaps the most frequently discussed subject in embryology, yet the relation of these layers to the individual blastomeres of the cleaving ovum had been determined in comparatively few cases. Since that time a number of very valuable papers have appeared on this subject of “cell lineage,” as Wilson (‘92) has aptly termed it. The results of such work are no longer as novel as they were four or five years ago, and yet the general interest in the subject has greatly increased, and that, too, in spite of the fact that there is a growing school of biologists who believe that individual blastomeres have no necessary relation to future organs. The subject of germ layers is no longer so important as it was once considered; in fact, the theory of the homology of the germinal layers has met with so many difficulties of late that it is now generally maintained only in a greatly modified form. However, the fundamental idea which was prominent in germ-layer discussions is of vital interest to-day. In the whole history of the germ-layer theories I see an attempt to trace homologies back to their earliest beginnings. This problem is as important to-day as it ever was, and whether one find these earliest homologies in layers or regions or blastomeres or the unsegmented ovum itself, the quest is essentially the same.

L'étude réalise un premier inventaire du peuplement halieutique et du rôle écologique d'un habitat original, : les vasières consolidées à Haploops. Haploops est un petit crustacé amphipode qui élabore des tubes souples et la concentration de ces tubes dans ces zones est telle que la pratique du chalutage y est très difficile. Situé dans la partie externe de la baie de la Vilaine, ce faciès couvre plus de 100 km2. Deux types de chaluts ont été adaptés à l'échantillonnage de ces fonds encombrés (chalut à perche String et chalut à panneaux avec grille ventrale) et utilisés au cours de trois campagnes du navire de recherche Gwen Drez. Une marée expérimentale sur le Marika, navire de pêche professionnel, a été réalisée pour apprécier l'intérêt commercial du secteur. Les principaux résultats montrent que, au sein de la même communauté des fonds côtiers vaso-sableux, des différences d'occurrence et d'abondance opposent le faciès à Haploops de son voisinage immédiat. Le benthos y est plus diversifié et marqué par l'abondance relative en crustacés décapodes, alors que les mollusques gastéropodes et céphalopodes y sont moins bien représentés. Quelques espèces commerciales y trouvent un habitat préférentiel, en particulier les roussettes, le tacaud, la dorade grise et l'étrille, ainsi que la coquille Saint-Jacques et, en faible abondance, la baudroie et le Saint-Pierre. Certaines espèces, au contraire, ne sont bien représentées qu'à l'extérieur du faciès, notamment les poissons plats (sole, plie, céteau), le merlu, le merlan et le bar. L'analyse de la fraction juvénile des principaux poissons montre que les vasières à Haploops jouent le rôle de nourricerie pour la petite roussette, la dorade grise, le Saint-Pierre, mais que la nourricerie de la baie de Vilaine ne se prolonge pas dans le faciès ; les juvéniles de merlu, merlan, bar, plie, sole et céteau demeurent en zone externe. Les fonds à Haploops constituent un habitat de moindre intérêt que les fonds de la baie de Vilaine

L'étude réalise un premier inventaire du peuplement halieutique et du rôle écologique d'un habitat original, : les vasières consolidées à Haploops. Haploops est un petit crustacé amphipode qui élabore des tubes souples et la concentration de ces tubes dans ces zones est telle que la pratique du chalutage y est très difficile. Situé dans la partie externe de la baie de la Vilaine, ce faciès couvre plus de 100 km2. Deux types de chaluts ont été adaptés à l'échantillonnage de ces fonds encombrés (chalut à perche String et chalut à panneaux avec grille ventrale) et utilisés au cours de trois campagnes du navire de recherche Gwen Drez. Une marée expérimentale sur le Marika, navire de pêche professionnel, a été réalisée pour apprécier l'intérêt commercial du secteur. Les principaux résultats montrent que, au sein de la même communauté des fonds côtiers vaso-sableux, des différences d'occurrence et d'abondance opposent le faciès à Haploops de son voisinage immédiat. Le benthos y est plus diversifié et marqué par l'abondance relative en crustacés décapodes, alors que les mollusques gastéropodes et céphalopodes y sont moins bien représentés. Quelques espèces commerciales y trouvent un habitat préférentiel, en particulier les roussettes, le tacaud, la dorade grise et l'étrille, ainsi que la coquille Saint-Jacques et, en faible abondance, la baudroie et le Saint-Pierre. Certaines espèces, au contraire, ne sont bien représentées qu'à l'extérieur du faciès, notamment les poissons plats (sole, plie, céteau), le merlu, le merlan et le bar. L'analyse de la fraction juvénile des principaux poissons montre que les vasières à Haploops jouent le rôle de nourricerie pour la petite roussette, la dorade grise, le Saint-Pierre, mais que la nourricerie de la baie de Vilaine ne se prolonge pas dans le faciès ; les juvéniles de merlu, merlan, bar, plie, sole et céteau demeurent en zone externe. Les fonds à Haploops constituent un habitat de moindre intérêt que les fonds de la baie de Vilaine

Effects of food availability on the larval survival and development of Crepidula onyx were studied in four experiments by feeding the larvae with different concentrations of the chrysophyte Isochrysis galbana and by starving the larvae for different periods of time. Food concentration had a clear impact on the survival, growth and development time of C onyx veligers. Larval development occurred only at 10(4) cells ml(-1) and higher algal concentrations. No shell increment was detected in the veligers cultured for 12 days at 10(2) Cells ml(-1) I. galbana or the blank control. At 10(3) cells ml(-1), there was only a slight increase in shell length over 12 days. At 10(4) cells ml(-1), about 40\% of the larvae became competent in 18 days. At 10(5) and 10(6) cells ml(-1), more than 90\% of the larvae reached competence in 7 days. Initial starvation negatively affected the larval development, but the sensitivity differed among parameters measured on day 5: lower survivorship was detected only for larvae that had suffered 3 days or longer initial starvation, whereas one-day initial starvation caused shorter shells and lower percentage of competent larvae. Three days of continuous feeding was required for 50\% of the larvae to reach competence. After feeding for 3 days, most larvae could become competent to metamorphose even under starvation. The time of starvation was also critical: larvae that suffered 1-day food deprivation in the first 2 days of larval release had shorter shells and lowered percent competent larvae than those that suffered the same length of food deprivation in later stages of development. Our study thus indicates that both food concentration and short-term starvation have detrimental effects on the larval development of this species, and that once the larva has consumed certain amount of food, starvation may induce metamorphosis. (C) 2003 Elsevier B.V. All rights reserved.

Effects of food availability on the larval survival and development of Crepidula onyx were studied in four experiments by feeding the larvae with different concentrations of the chrysophyte Isochrysis galbana and by starving the larvae for different periods of time. Food concentration had a clear impact on the survival, growth and development time of C onyx veligers. Larval development occurred only at 10(4) cells ml(-1) and higher algal concentrations. No shell increment was detected in the veligers cultured for 12 days at 10(2) Cells ml(-1) I. galbana or the blank control. At 10(3) cells ml(-1), there was only a slight increase in shell length over 12 days. At 10(4) cells ml(-1), about 40\% of the larvae became competent in 18 days. At 10(5) and 10(6) cells ml(-1), more than 90\% of the larvae reached competence in 7 days. Initial starvation negatively affected the larval development, but the sensitivity differed among parameters measured on day 5: lower survivorship was detected only for larvae that had suffered 3 days or longer initial starvation, whereas one-day initial starvation caused shorter shells and lower percentage of competent larvae. Three days of continuous feeding was required for 50\% of the larvae to reach competence. After feeding for 3 days, most larvae could become competent to metamorphose even under starvation. The time of starvation was also critical: larvae that suffered 1-day food deprivation in the first 2 days of larval release had shorter shells and lowered percent competent larvae than those that suffered the same length of food deprivation in later stages of development. Our study thus indicates that both food concentration and short-term starvation have detrimental effects on the larval development of this species, and that once the larva has consumed certain amount of food, starvation may induce metamorphosis. (C) 2003 Elsevier B.V. All rights reserved.

Effects of food availability on the larval survival and development of Crepidula onyx were studied in four experiments by feeding the larvae with different concentrations of the chrysophyte Isochrysis galbana and by starving the larvae for different periods of time. Food concentration had a clear impact on the survival, growth and development time of C onyx veligers. Larval development occurred only at 10(4) cells ml(-1) and higher algal concentrations. No shell increment was detected in the veligers cultured for 12 days at 10(2) Cells ml(-1) I. galbana or the blank control. At 10(3) cells ml(-1), there was only a slight increase in shell length over 12 days. At 10(4) cells ml(-1), about 40\% of the larvae became competent in 18 days. At 10(5) and 10(6) cells ml(-1), more than 90\% of the larvae reached competence in 7 days. Initial starvation negatively affected the larval development, but the sensitivity differed among parameters measured on day 5: lower survivorship was detected only for larvae that had suffered 3 days or longer initial starvation, whereas one-day initial starvation caused shorter shells and lower percentage of competent larvae. Three days of continuous feeding was required for 50\% of the larvae to reach competence. After feeding for 3 days, most larvae could become competent to metamorphose even under starvation. The time of starvation was also critical: larvae that suffered 1-day food deprivation in the first 2 days of larval release had shorter shells and lowered percent competent larvae than those that suffered the same length of food deprivation in later stages of development. Our study thus indicates that both food concentration and short-term starvation have detrimental effects on the larval development of this species, and that once the larva has consumed certain amount of food, starvation may induce metamorphosis. (C) 2003 Elsevier B.V. All rights reserved.

The Atlantic slipper limpet, Crepidula fornicata, is a sequential hermaphrodite whose size at sex-change is plastic with respect to social and population cues. As an organism allocates energy between growth, reproduction and maintenance, an increased cost of one process may affect another. In this paper, I evaluate whether the presence of an epibiotic sponge (Cliona celata) affects the growth, condition and sex-change of C. fornicata individuals. Population surveys demonstrate a variable effect of Cliona presence on C. fornicata condition. The results of a twelve-week in situ experiment demonstrated a decrease in growth when C. celata was present. Regarding the timing of sex-change, the results of the study were inconclusive. The results show that C. celata negatively affects C. fornicata growth and condition in some contexts, though more precise studies are required to determine whether sex-change is also affected.

The Atlantic slipper limpet, Crepidula fornicata, is a sequential hermaphrodite whose size at sex-change is plastic with respect to social and population cues. As an organism allocates energy between growth, reproduction and maintenance, an increased cost of one process may affect another. In this paper, I evaluate whether the presence of an epibiotic sponge (Cliona celata) affects the growth, condition and sex-change of C. fornicata individuals. Population surveys demonstrate a variable effect of Cliona presence on C. fornicata condition. The results of a twelve-week in situ experiment demonstrated a decrease in growth when C. celata was present. Regarding the timing of sex-change, the results of the study were inconclusive. The results show that C. celata negatively affects C. fornicata growth and condition in some contexts, though more precise studies are required to determine whether sex-change is also affected.

The Atlantic slipper limpet, Crepidula fornicata, is a sequential hermaphrodite whose size at sex-change is plastic with respect to social and population cues. As an organism allocates energy between growth, reproduction and maintenance, an increased cost of one process may affect another. In this paper, I evaluate whether the presence of an epibiotic sponge (Cliona celata) affects the growth, condition and sex-change of C. fornicata individuals. Population surveys demonstrate a variable effect of Cliona presence on C. fornicata condition. The results of a twelve-week in situ experiment demonstrated a decrease in growth when C. celata was present. Regarding the timing of sex-change, the results of the study were inconclusive. The results show that C. celata negatively affects C. fornicata growth and condition in some contexts, though more precise studies are required to determine whether sex-change is also affected.