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Predatory suspension feeders: an unusual feeding mode switch in Olivella columellaris (Caenogastropoda: Olividae) and its possible ecological effects

Predatory suspension feeders: an unusual feeding mode switch in Olivella columellaris... In analyses of trophic networks, the complexities of animal communities often have to be reduced by grouping species that share similar features. Which group-defining features are most meaningful in a given context is a notoriously problematic question (Simberloff & Dayan, 1991; Wilson, 1999; Blondel, 2003; Blaum et al., 2011), even in seemingly simple cases such as the categorization of marine benthos according to feeding modes (Cadée, 1984). According to the classical definition (Hunt, 1925), suspension or filter feeders collect detritus and micro-organisms from the water, deposit feeders extract similar materials from the sediment, and carnivores, including both predators and scavengers, feed on animal bodies. While this seems straight-forward, it has long been known that some benthic polychaetes (Dauer, Maybury & Ewing 1981; Taghon & Greene, 1992), crustaceans (Mills, 1967; Riisgård & Schotge, 2007), bivalves (Navarro et al., 2008; Törnroos et al., 2015), gastropods (Navarro & Chaparro, 2002; Kamimura & Tsuchiya, 2006) and ophiuroids (Miller, Bock & Turner 1992; Loo et al., 1996) switch between suspension- and deposit-feeding depending on circumstances. Switching between microphagous suspension-feeding and macrophagous carnivory is far less common, probably due to incompatible constraints on digestive functions (Jumars, Dorgan & Lindsay 2015). Examples exist, though, such as the polychaete Hediste (previously Nereis) diversicolor (Müller, 1776) discussed by Hartmann-Schröder (1996) and Riisgård & Kamermans (2001). Here we report a new case, the gastropod Olivella columellaris G.B. Sowerby I, 1825, and suggest that its unusual feeding versatility might explain its conspicuous dominance on many sandy beaches of the tropical East Pacific. Olividae (Caengastropoda: Neogastropoda), marine gastropods typically found on soft sediments in shallow water (Kantor et al., 2017), generally are predators and scavengers (Marcus & Marcus, 1959; Kantor, 1991; Kantor & Tursch, 2001; Robinson & Peters, 2018). However, O. columellaris and O. semistriataGray, 1839, which form the subgenus PachyolivaOlsson, 1956, in the subfamily Olivellinae, represent exceptions as they are passive filter feeders. The two species often dominate intertidal sandy beach communities on the tropical American west coast and have been confused frequently as they resemble each other morphologically and behaviourally (Troost et al., 2012). Their foot consists of a posterior main part, the metapodium, and a small, crescent-shaped anterior propodium that carries two elongate, lateral appendages. As described first for O. semistriata (misidentified as O. columellaris; Schuster, 1952; Seilacher, 1959), the animals use these appendages to expose mucus membranes to the backwash, collecting suspended particles from the seawards-flowing water (Fig. 1). The particle-loaded mucus is eaten at short intervals. Propodial appendages characterized by their ability to spread and retract mucus sheets in a controlled manner are unique in gastropods and represent an autapomorphy of Pachyoliva (Troost et al., 2012). Figure 1. Open in new tabDownload slide Suspension-feeding Olivella columellaris. A. Suspension-feeding animals are burrowed in the sediment except for the propodium (anterior foot), the ventral side of which faces into the flow (arrows). The lateral propodial appendages (arrowheads) support mucus sheets that bulge into semi-spheres (asterisks) in the flow. The proboscis with the mouth opening (white circle) is located dorsally. B. Snail with partly missing left propodial appendage. Due to the asymmetry, the current (arrows) rotates the propodium clockwise in this view. Figure 1. Open in new tabDownload slide Suspension-feeding Olivella columellaris. A. Suspension-feeding animals are burrowed in the sediment except for the propodium (anterior foot), the ventral side of which faces into the flow (arrows). The lateral propodial appendages (arrowheads) support mucus sheets that bulge into semi-spheres (asterisks) in the flow. The proboscis with the mouth opening (white circle) is located dorsally. B. Snail with partly missing left propodial appendage. Due to the asymmetry, the current (arrows) rotates the propodium clockwise in this view. We studied a population of O. columellaris near Colan, Peru (4°59′15″S, 81°4′30″W) in August 2013 (cool season) and March 2014 (hot season). The micro- to mesotidal study beach experiences tidal ranges from 0.5 m (neap tide) to 2.2 m (spring tide), resulting in horizontal movements of the water line between 10 and >80 m. At any season and time of day, and also at all stages of the tidal cycle, dense accumulations of O. columellaris were suspension-feeding in the backwash zone. This suggested tidal migrations. To determine population densities of benthic species, short rigid plastic tubes (28 cm diameter, 20 cm length) were pushed into the sand. The sediment within the plastic rings was removed to 8 cm depth, sieved (1.5 mm mesh) and the retained organisms were identified and counted. The median density of O. columellaris in samples taken in the backwash zone at high tide in August was 5,817 individuals m–2 (range 1,095–19,033, n = 12). At the same spots at the subsequent low tide, it was 0 individuals m–2 (range 0–9, n = 12), confirming that tidal migrations occurred (Fig. 2). Densities of all other invertebrates combined—mainly the gastropod Mazatlania sp., bivalves (Donax sp.) and occasional polychaetes and small crustaceans—never exceeded 90 individuals m–2 in these 24 samples. Figure 2. Open in new tabDownload slide Densities of Olivella columellaris in the feeding zone in the backwash at high tide and at the identical locations during the following low tide. Data-points represent four independent samples taken at each of three consecutive high tides and again at each subsequent low tide on 2 and 3 August 2013. Figure 2. Open in new tabDownload slide Densities of Olivella columellaris in the feeding zone in the backwash at high tide and at the identical locations during the following low tide. Data-points represent four independent samples taken at each of three consecutive high tides and again at each subsequent low tide on 2 and 3 August 2013. We recorded six instances of a polychaete protruding from the proboscis of an O. columellaris (Fig. 3A). All were found in the cool season (August: 2,699 O. columellaris examined) but none in the hot season (March: 16,960 O. columellaris examined), which may reflect seasonality in polychaete density. The polychaetes clearly were alive as they occasionally moved. Snails carrying worms showed normal crawling behavior and, when placed on beach sand, burrowed in their usual manner. Sometimes a gastropod bent its propodium dorsally, apparently using the opposable left and right halves of the propodium to manipulate the polychaete in its proboscis (Supplementary Material Video S1). Similar usage of the propodium has been described in Callianax (formerly Olivella) biplicata, a closely related scavenger/predator (Kelly et al., 2021). Figure 3. Open in new tabDownload slide Carnivory in Olivella columellaris. A. Individual with a polychaete (Nephtys sp.) in its proboscis (compare Supplementary Material Video S1). B. Extraction of the polychaete (compare Supplementary Material Video S2). C. Extracted polychaete; arrowheads mark the position of the gastropod's mouth opening before extraction. The part of the worm that had been located deepest in the snail's digestive tract appears to consist of remains of the integument only (arrows; compare Supplementary Material Video S2). D. Specimen feeding on shrimp meat (asterisk), with proboscis marked by arrow (compare Supplementary Material Video S3). Figure 3. Open in new tabDownload slide Carnivory in Olivella columellaris. A. Individual with a polychaete (Nephtys sp.) in its proboscis (compare Supplementary Material Video S1). B. Extraction of the polychaete (compare Supplementary Material Video S2). C. Extracted polychaete; arrowheads mark the position of the gastropod's mouth opening before extraction. The part of the worm that had been located deepest in the snail's digestive tract appears to consist of remains of the integument only (arrows; compare Supplementary Material Video S2). D. Specimen feeding on shrimp meat (asterisk), with proboscis marked by arrow (compare Supplementary Material Video S3). Six cases of polychaete feeding observed in a sample of almost 20,000 snails do not seem to suggest a major trophic role for predation in O. columellaris compared to their ubiquitous suspension-feeding in the backwash. However, worms barely protruding from the snails’ probosces might have been overlooked, and predation on smaller prey that the gastropods can swallow whole would mostly be missed in the natural habitat. Thus, carnivory could contribute significantly to the energy budget of this suspension feeder. Moreover, the gastropods occasionally lose parts of their propodial appendages, which probably affects the efficiency of their suspension feeding (Fig. 1B). In benthic polychaetes, feeding mode switches enable survival after the loss of body parts that are essential for feeding by one particular mode (Lindsay & Woodin, 1995). Similarly, the ability to switch to carnivory should increase the fitness of O. columellaris when its propodial appendages are damaged or when suspension feeding is impossible for other reasons. Three of the polychaetes were extracted with forceps (Fig. 3B; Supplementary Material Video S2). They died within 1–2 h, whereas the snails exhibited normal activities after the procedure. Dead polychaetes were conserved in ethanol (local aguardiente being the only source available) and deposited at the Instituto del Mar del Peru, Callao, Peru (acc. no, 07-000005); all other collected animals were released alive into their habitat within 24 h. The parts of the extracted polychaetes that had been located within the gastropods’ digestive tracts showed increasing structural decay with distance from the snails’ mouth openings (Fig. 3C). One snail carried a worm that protruded from the proboscis by 7 mm when the animals were found (Supplementary Material Video S1). After 10 h in captivity, the snail spontaneously released the worm, which now measured only 5 mm in total. Both ends of this polychaete appeared similarly decomposed, as if two snails had fed on it from opposite sides. The extracted polychaetes belonged to the genus NephtysCuvier, 1817 (Annelida: Nephtyidae), but their state of preservation did not permit further identification. The geographically closest previous record of this genus is N. simoniPerkins, 1980, which Hartman (1941) reported as N. magellanicaAugener, 1912 from the Bahia de Sechura, some 60 km south of our study site (see also Moreno et al., 2021). Evidently, O. columellaris ingested polychaetes in a prolonged process that depended on the progress of digestion in the gastropod's intestinal tract. To see whether the usually suspension-feeding O. columellaris generally switches to carnivory when given the opportunity, we offered shrimp meat to individuals in aquaria, not later than 2 h after they had been collected. Five of the nine gastropods tested grabbed the meat with their propodia and fed eagerly (Fig. 3D; Supplementary Material Video S3) in a similar manner as carnivorous Olivellinae do (Kelly et al., 2021). They did not use their propodial appendages in the process. The remaining four specimens ignored the food. Analogous feeding tests with O. semistriata, the other Pachyoliva, were conducted at Playa Grande, Costa Rica (10°20′10″N, 85°51′0″W). Although carnivory in the wild has never been reported for O. semistriata, it responded like O. columellaris in these tests (Supplementary Material Video S4). Tidal migrations have been interpreted as a necessary behavioural adaptation in Pachyoliva species, which seem to depend on suspension feeding in the continuously moving backwash zone (Troost et al., 2012; Seilacher, 1959). Vanagt et al. (2008) inferred from laboratory tests with O. columellaris (misidentified as O. semistriata) that an endogenous, circatidal clock controls migratory behaviour. This interpretation conflicts with observations in natural habitats of an opportunistic exploitation of suitable flow conditions, independently of the tidal progression (Morse & Peters, 2016). Rather than being driven by an endogenous rhythm, tidal migrations, which the animals doubtlessly perform, may result from collective opportunism: each snail stays where it can feed and moves from where it cannot (Morse & Peters, 2016; McLachlan, Wooldridge & van der Horst, 1979). Unlike suspension feeding in the backwash, carnivory is not necessarily linked to a particular beach zone. Therefore, our observation of carnivory in O. columellaris adds to the doubts about a strict dependence of feeding on tidal migrations and further questions the postulated requirement for an endogenous clock. Conducting the first systematic studies of the intertidal fauna in North Peru, Olsson (1923–1924: 122) found sandy beaches in our study region ``devoid of much interest'' as they were ``characterized by the abundance of a few species'', including O. columellaris (as O. semistriata). Similarly, O. columellaris (as O. semistriata) contributed 66% of the macrobenthos on beaches in Ecuador, with an average density of 243 individuals m–2 (Aerts et al., 2004). These figures reflect total counts across the entire beach and underestimate the density of O. columellaris in the backwash zone. At our study site, the median density in the backwash was about 6,000 individuals m–2. If 6,000 gastropods are distributed homogeneously over 1 m2 in a hexagonal array, the distance between any two neighbours will be 13.9 mm, which is about the shell length of large specimens. Even if only 1% of these snails were inclined to predaceous behaviour at any time, potential prey in the backwash zone would always be less than 140 mm away from a motivated predator. While the high-density feeding zone usually is only a few meters wide, it is rolling across the beach slope continuously with the tides. Thus, every intertidal location periodically experiences dense accumulations of potentially carnivorous gastropods, which must generate considerable pressure on prey species as well as on competing carnivores. Textbook wisdom has it that predator populations will decline if they drive their prey towards extinction. But O. columellaris can keep suspension-feeding when this happens—and which predator could have a more profound ecological impact than one that thrives regardless of prey availability? From a community-ecological perspective, the interpretation of O. columellaris as a suspension feeder that occasionally switches to carnivory appears misleading. 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Predatory suspension feeders: an unusual feeding mode switch in Olivella columellaris (Caenogastropoda: Olividae) and its possible ecological effects

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Oxford University Press
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© The Author(s) 2022. Published by Oxford University Press on behalf of The Malacological Society of London.
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0260-1230
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Abstract

In analyses of trophic networks, the complexities of animal communities often have to be reduced by grouping species that share similar features. Which group-defining features are most meaningful in a given context is a notoriously problematic question (Simberloff & Dayan, 1991; Wilson, 1999; Blondel, 2003; Blaum et al., 2011), even in seemingly simple cases such as the categorization of marine benthos according to feeding modes (Cadée, 1984). According to the classical definition (Hunt, 1925), suspension or filter feeders collect detritus and micro-organisms from the water, deposit feeders extract similar materials from the sediment, and carnivores, including both predators and scavengers, feed on animal bodies. While this seems straight-forward, it has long been known that some benthic polychaetes (Dauer, Maybury & Ewing 1981; Taghon & Greene, 1992), crustaceans (Mills, 1967; Riisgård & Schotge, 2007), bivalves (Navarro et al., 2008; Törnroos et al., 2015), gastropods (Navarro & Chaparro, 2002; Kamimura & Tsuchiya, 2006) and ophiuroids (Miller, Bock & Turner 1992; Loo et al., 1996) switch between suspension- and deposit-feeding depending on circumstances. Switching between microphagous suspension-feeding and macrophagous carnivory is far less common, probably due to incompatible constraints on digestive functions (Jumars, Dorgan & Lindsay 2015). Examples exist, though, such as the polychaete Hediste (previously Nereis) diversicolor (Müller, 1776) discussed by Hartmann-Schröder (1996) and Riisgård & Kamermans (2001). Here we report a new case, the gastropod Olivella columellaris G.B. Sowerby I, 1825, and suggest that its unusual feeding versatility might explain its conspicuous dominance on many sandy beaches of the tropical East Pacific. Olividae (Caengastropoda: Neogastropoda), marine gastropods typically found on soft sediments in shallow water (Kantor et al., 2017), generally are predators and scavengers (Marcus & Marcus, 1959; Kantor, 1991; Kantor & Tursch, 2001; Robinson & Peters, 2018). However, O. columellaris and O. semistriataGray, 1839, which form the subgenus PachyolivaOlsson, 1956, in the subfamily Olivellinae, represent exceptions as they are passive filter feeders. The two species often dominate intertidal sandy beach communities on the tropical American west coast and have been confused frequently as they resemble each other morphologically and behaviourally (Troost et al., 2012). Their foot consists of a posterior main part, the metapodium, and a small, crescent-shaped anterior propodium that carries two elongate, lateral appendages. As described first for O. semistriata (misidentified as O. columellaris; Schuster, 1952; Seilacher, 1959), the animals use these appendages to expose mucus membranes to the backwash, collecting suspended particles from the seawards-flowing water (Fig. 1). The particle-loaded mucus is eaten at short intervals. Propodial appendages characterized by their ability to spread and retract mucus sheets in a controlled manner are unique in gastropods and represent an autapomorphy of Pachyoliva (Troost et al., 2012). Figure 1. Open in new tabDownload slide Suspension-feeding Olivella columellaris. A. Suspension-feeding animals are burrowed in the sediment except for the propodium (anterior foot), the ventral side of which faces into the flow (arrows). The lateral propodial appendages (arrowheads) support mucus sheets that bulge into semi-spheres (asterisks) in the flow. The proboscis with the mouth opening (white circle) is located dorsally. B. Snail with partly missing left propodial appendage. Due to the asymmetry, the current (arrows) rotates the propodium clockwise in this view. Figure 1. Open in new tabDownload slide Suspension-feeding Olivella columellaris. A. Suspension-feeding animals are burrowed in the sediment except for the propodium (anterior foot), the ventral side of which faces into the flow (arrows). The lateral propodial appendages (arrowheads) support mucus sheets that bulge into semi-spheres (asterisks) in the flow. The proboscis with the mouth opening (white circle) is located dorsally. B. Snail with partly missing left propodial appendage. Due to the asymmetry, the current (arrows) rotates the propodium clockwise in this view. We studied a population of O. columellaris near Colan, Peru (4°59′15″S, 81°4′30″W) in August 2013 (cool season) and March 2014 (hot season). The micro- to mesotidal study beach experiences tidal ranges from 0.5 m (neap tide) to 2.2 m (spring tide), resulting in horizontal movements of the water line between 10 and >80 m. At any season and time of day, and also at all stages of the tidal cycle, dense accumulations of O. columellaris were suspension-feeding in the backwash zone. This suggested tidal migrations. To determine population densities of benthic species, short rigid plastic tubes (28 cm diameter, 20 cm length) were pushed into the sand. The sediment within the plastic rings was removed to 8 cm depth, sieved (1.5 mm mesh) and the retained organisms were identified and counted. The median density of O. columellaris in samples taken in the backwash zone at high tide in August was 5,817 individuals m–2 (range 1,095–19,033, n = 12). At the same spots at the subsequent low tide, it was 0 individuals m–2 (range 0–9, n = 12), confirming that tidal migrations occurred (Fig. 2). Densities of all other invertebrates combined—mainly the gastropod Mazatlania sp., bivalves (Donax sp.) and occasional polychaetes and small crustaceans—never exceeded 90 individuals m–2 in these 24 samples. Figure 2. Open in new tabDownload slide Densities of Olivella columellaris in the feeding zone in the backwash at high tide and at the identical locations during the following low tide. Data-points represent four independent samples taken at each of three consecutive high tides and again at each subsequent low tide on 2 and 3 August 2013. Figure 2. Open in new tabDownload slide Densities of Olivella columellaris in the feeding zone in the backwash at high tide and at the identical locations during the following low tide. Data-points represent four independent samples taken at each of three consecutive high tides and again at each subsequent low tide on 2 and 3 August 2013. We recorded six instances of a polychaete protruding from the proboscis of an O. columellaris (Fig. 3A). All were found in the cool season (August: 2,699 O. columellaris examined) but none in the hot season (March: 16,960 O. columellaris examined), which may reflect seasonality in polychaete density. The polychaetes clearly were alive as they occasionally moved. Snails carrying worms showed normal crawling behavior and, when placed on beach sand, burrowed in their usual manner. Sometimes a gastropod bent its propodium dorsally, apparently using the opposable left and right halves of the propodium to manipulate the polychaete in its proboscis (Supplementary Material Video S1). Similar usage of the propodium has been described in Callianax (formerly Olivella) biplicata, a closely related scavenger/predator (Kelly et al., 2021). Figure 3. Open in new tabDownload slide Carnivory in Olivella columellaris. A. Individual with a polychaete (Nephtys sp.) in its proboscis (compare Supplementary Material Video S1). B. Extraction of the polychaete (compare Supplementary Material Video S2). C. Extracted polychaete; arrowheads mark the position of the gastropod's mouth opening before extraction. The part of the worm that had been located deepest in the snail's digestive tract appears to consist of remains of the integument only (arrows; compare Supplementary Material Video S2). D. Specimen feeding on shrimp meat (asterisk), with proboscis marked by arrow (compare Supplementary Material Video S3). Figure 3. Open in new tabDownload slide Carnivory in Olivella columellaris. A. Individual with a polychaete (Nephtys sp.) in its proboscis (compare Supplementary Material Video S1). B. Extraction of the polychaete (compare Supplementary Material Video S2). C. Extracted polychaete; arrowheads mark the position of the gastropod's mouth opening before extraction. The part of the worm that had been located deepest in the snail's digestive tract appears to consist of remains of the integument only (arrows; compare Supplementary Material Video S2). D. Specimen feeding on shrimp meat (asterisk), with proboscis marked by arrow (compare Supplementary Material Video S3). Six cases of polychaete feeding observed in a sample of almost 20,000 snails do not seem to suggest a major trophic role for predation in O. columellaris compared to their ubiquitous suspension-feeding in the backwash. However, worms barely protruding from the snails’ probosces might have been overlooked, and predation on smaller prey that the gastropods can swallow whole would mostly be missed in the natural habitat. Thus, carnivory could contribute significantly to the energy budget of this suspension feeder. Moreover, the gastropods occasionally lose parts of their propodial appendages, which probably affects the efficiency of their suspension feeding (Fig. 1B). In benthic polychaetes, feeding mode switches enable survival after the loss of body parts that are essential for feeding by one particular mode (Lindsay & Woodin, 1995). Similarly, the ability to switch to carnivory should increase the fitness of O. columellaris when its propodial appendages are damaged or when suspension feeding is impossible for other reasons. Three of the polychaetes were extracted with forceps (Fig. 3B; Supplementary Material Video S2). They died within 1–2 h, whereas the snails exhibited normal activities after the procedure. Dead polychaetes were conserved in ethanol (local aguardiente being the only source available) and deposited at the Instituto del Mar del Peru, Callao, Peru (acc. no, 07-000005); all other collected animals were released alive into their habitat within 24 h. The parts of the extracted polychaetes that had been located within the gastropods’ digestive tracts showed increasing structural decay with distance from the snails’ mouth openings (Fig. 3C). One snail carried a worm that protruded from the proboscis by 7 mm when the animals were found (Supplementary Material Video S1). After 10 h in captivity, the snail spontaneously released the worm, which now measured only 5 mm in total. Both ends of this polychaete appeared similarly decomposed, as if two snails had fed on it from opposite sides. The extracted polychaetes belonged to the genus NephtysCuvier, 1817 (Annelida: Nephtyidae), but their state of preservation did not permit further identification. The geographically closest previous record of this genus is N. simoniPerkins, 1980, which Hartman (1941) reported as N. magellanicaAugener, 1912 from the Bahia de Sechura, some 60 km south of our study site (see also Moreno et al., 2021). Evidently, O. columellaris ingested polychaetes in a prolonged process that depended on the progress of digestion in the gastropod's intestinal tract. To see whether the usually suspension-feeding O. columellaris generally switches to carnivory when given the opportunity, we offered shrimp meat to individuals in aquaria, not later than 2 h after they had been collected. Five of the nine gastropods tested grabbed the meat with their propodia and fed eagerly (Fig. 3D; Supplementary Material Video S3) in a similar manner as carnivorous Olivellinae do (Kelly et al., 2021). They did not use their propodial appendages in the process. The remaining four specimens ignored the food. Analogous feeding tests with O. semistriata, the other Pachyoliva, were conducted at Playa Grande, Costa Rica (10°20′10″N, 85°51′0″W). Although carnivory in the wild has never been reported for O. semistriata, it responded like O. columellaris in these tests (Supplementary Material Video S4). Tidal migrations have been interpreted as a necessary behavioural adaptation in Pachyoliva species, which seem to depend on suspension feeding in the continuously moving backwash zone (Troost et al., 2012; Seilacher, 1959). Vanagt et al. (2008) inferred from laboratory tests with O. columellaris (misidentified as O. semistriata) that an endogenous, circatidal clock controls migratory behaviour. This interpretation conflicts with observations in natural habitats of an opportunistic exploitation of suitable flow conditions, independently of the tidal progression (Morse & Peters, 2016). Rather than being driven by an endogenous rhythm, tidal migrations, which the animals doubtlessly perform, may result from collective opportunism: each snail stays where it can feed and moves from where it cannot (Morse & Peters, 2016; McLachlan, Wooldridge & van der Horst, 1979). Unlike suspension feeding in the backwash, carnivory is not necessarily linked to a particular beach zone. Therefore, our observation of carnivory in O. columellaris adds to the doubts about a strict dependence of feeding on tidal migrations and further questions the postulated requirement for an endogenous clock. Conducting the first systematic studies of the intertidal fauna in North Peru, Olsson (1923–1924: 122) found sandy beaches in our study region ``devoid of much interest'' as they were ``characterized by the abundance of a few species'', including O. columellaris (as O. semistriata). Similarly, O. columellaris (as O. semistriata) contributed 66% of the macrobenthos on beaches in Ecuador, with an average density of 243 individuals m–2 (Aerts et al., 2004). These figures reflect total counts across the entire beach and underestimate the density of O. columellaris in the backwash zone. At our study site, the median density in the backwash was about 6,000 individuals m–2. If 6,000 gastropods are distributed homogeneously over 1 m2 in a hexagonal array, the distance between any two neighbours will be 13.9 mm, which is about the shell length of large specimens. Even if only 1% of these snails were inclined to predaceous behaviour at any time, potential prey in the backwash zone would always be less than 140 mm away from a motivated predator. While the high-density feeding zone usually is only a few meters wide, it is rolling across the beach slope continuously with the tides. Thus, every intertidal location periodically experiences dense accumulations of potentially carnivorous gastropods, which must generate considerable pressure on prey species as well as on competing carnivores. Textbook wisdom has it that predator populations will decline if they drive their prey towards extinction. But O. columellaris can keep suspension-feeding when this happens—and which predator could have a more profound ecological impact than one that thrives regardless of prey availability? From a community-ecological perspective, the interpretation of O. columellaris as a suspension feeder that occasionally switches to carnivory appears misleading. 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Journal of Molluscan StudiesOxford University Press

Published: Jun 27, 2022

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