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ABSTRACT
Cultivation of eastern oysters using aquaculture gear increases habitat for temperate reef fish. Cunner (Tautogolabrus adspersus), scup (Stenotomus chrysops) and tautog (Tautoga onitis) inhabit a variety of complex natural and manmade habitats, including oyster aquaculture cage farms. Underwater video was recorded on two cage farms and a rock reef during May–September 2018 to quantify fish behavioural interactions and to assess ecological services provided by aquaculture gear, relative to natural structured seafloor. To collect video, action cameras were mounted on four study cages at a high‐density farm of 40–100 commercial cages (dense farm), on four single cages interspersed on low relief seafloor (sparse farm) and adjacent to four boulders on a rock reef (rock reef), within an embayment off Milford, Connecticut in Long Island Sound (NW Atlantic). Video was recorded hourly in 8‐min segments from 7 AM to 7 PM. Behaviours associated with habitat provisioning (e.g., courtship/reproduction, escape from predators, foraging, sheltering, schooling/grouping, territoriality) were observed for all three fish species on cages and boulders. Foraging and sheltering activity in cunner was significantly higher on cages than boulders while territorial behaviour occurred more frequently on boulders. Instances of escape from predators, foraging and sheltering behaviours in scup and tautog were significantly higher on cages than on boulders. Courtship/reproduction, grouping and territoriality were also higher on cages than boulders in tautog. Our results suggest that oyster cages confer ecological services that fulfil the basic biological and functional requirements of reef‐oriented fish, and provide habitat attributes afforded by natural rock reefs.
Introduction
Aquaculture gear intended for the grow-out of shellfish can confer ecosystem benefits beyond that of food production, including enhancement of habitat for wild fish (Alleway et al. 2019; Ferriss et al. 2021; Martínez-Baena et al. 2022). Multi-tiered oyster aquaculture cages, used in the cultivation of eastern oysters (Crassostrea virginica), contribute structurally-complex habitat to marine environments that may also provide ecological services to shelter-oriented fish (O'Brien et al. 2018; Theuerkauf et al. 2021; Lefcheck et al. 2021). Fish rely on ecosystem benefits, such as increased forage, shelter sites, breeding habitat, nursery areas and protection from predation (e.g., Forrest et al. 2009; Corrigan et al. 2024; Mercaldo-Allen et al. 2025), for successful growth, reproduction and survival (Brown et al. 2022). Comparing relative frequencies of habitat-related behaviours, such as territoriality where fish defend preferred territory, and occupancy, where fish shelter on or alongside structure, may serve as indicators of the quality of habitat (Lindell 2008; Armbruster et al. 2024; Mercaldo-Allen et al. 2025). Behavioural observations of fish on and around oyster cages may help identify benefits provided to fish and provide insights into the functional value of shellfish aquaculture gear in fish community ecology.
Temperate reef fish species in southern New England include the black sea bass (Centropristis striata), cunner (Tautogolabrus adspersus), scup (Stenotomus chrysops) and tautog (Tautoga onitis) (Weiss 1997), which reside on both natural and manmade structured habitat (e.g., Steimle et al. 1999). Black sea bass and scup, managed jointly by the Mid-Atlantic Fishery Management Council and the Atlantic States Marine Fisheries Commission, have federally designated essential fish habitat (EFH), while tautog and cunner are not federally managed nor have designated EFH. Black sea bass, scup and tautog are all economically valued within the commercial and recreational fishery, while cunner, although ecologically important, are not a managed or commonly-targeted species.
Shelter constitutes a functional requirement for cunner and tautog, which demonstrate high site fidelity and limited home ranges, particularly during early life stages (e.g., Olla et al. 1974; Auster 1989; Able and Fahay 1998; Tupper and Boutilier 1997; Able et al. 2005). Scup, another sympatric reef species, also prefers structure but has a broader habitat distribution, and is found in both open and rocky seafloor environments (Steimle et al. 1999). These structure-oriented fish are also commonly observed on oyster cage farms, where they occur at similar or higher abundance than on natural rock reefs (Mercaldo-Allen et al. 2020, 2021, 2023). The high affinity for oyster aquaculture gear shown by temperate reef species suggests that cages possess habitat characteristics that meet the basic functional and biological needs of fish, in much the same way as natural hard bottom seafloor and other types of man-made structured habitats (e.g., jetties, shipwrecks, wharves, artificial reefs) (Auster 1989; Steimle et al. 1999; Steimle and Shaheen 1999).
Visual assessment of fish behavioural interactions with oyster farm cages and boulders may enhance our understanding of habitat provisioning by shellfish aquaculture gear. The utility of underwater video for cataloguing fish activity in difficult to sample complex seafloor habitats, including oyster aquaculture farms and natural rock reefs, has been well established (e.g., Muething et al. 2020; Ferriss et al. 2021; Mercaldo-Allen et al. 2021, 2023; Shinn et al. 2021; Veggerby et al. 2024; Ambrose and Munroe 2025). Action cameras capture fish activity with less disturbance than other direct visual methods (e.g., diver surveys), enable the use of timers to record at intervals across a day or tidal cycle, allow for concurrent spatial and temporal replication and create a permanent archive of footage that can be revisited to confirm fish identification or conduct further analysis (Zarco-Perello and Enriquez 2019; Mercaldo-Allen et al. 2021, 2023; Farrell et al. 2023). Documenting fish interactions with their surroundings can provide information on species-specific habitat use and preferences (Shinn et al. 2021; Armbruster et al. 2024). Video sampling may be particularly valuable for assessing nuanced behaviour of wild fish around aquaculture gear and natural structure (Muething et al. 2020; Shinn et al. 2021), allowing for extended monitoring of fish movements. Recent behavioural analysis of black sea bass in underwater videos recorded on cage farms and rock reefs has found that cage structures provide ecological services to this species similar to that afforded by boulder habitat (Mercaldo-Allen et al. 2021, 2023, 2025; Armbruster et al. 2024). However, behavioural interactions of cunner, scup and tautog associated with oyster aquaculture gear relative to natural structured habitat have yet to be described.
Understanding habitat interactions of wild fish on shellfish farms can inform decision makers regarding fisheries resource management, aquaculture permitting processes and identify potential benefits provided to fish by oyster aquaculture cages (Mercaldo-Allen et al. 2021; Farrell et al. 2023; Ambrose and Munroe 2025). Despite growing evidence of provisioning of habitat by shellfish aquaculture gear to economically important fish species, uncertainty related to how and to what extent aquaculture structures function relative to comparable natural habitats has impeded the consideration of habitat provisioning by oyster aquaculture gear into the regulatory decision-making process, including the EFH consultation process. This often puts oyster aquaculture industry growth at odds with habitat conservation and restoration goals. The ability for resource managers and permitting agencies to quantitatively consider the benefits of an oyster aquaculture farm proposal against its potential adverse effects provides an opportunity for a balanced review and may allow for consideration of how growth in shellfish cage-based aquaculture can support fisheries management.
The goal of this work is to catalogue and describe behaviour of three temperate reef fish species, cunner, scup and tautog, around oyster aquaculture cages on shellfish farms and on boulders at natural rock reefs, in order to document the ecological services and habitat function provided by cages, relative to natural structured seafloor. Comparing fish behaviour on cages versus boulders provides context for whether artificial structures confer services to fish in the same way as hard bottom rock reef habitat.
Materials and Methods
Study Sites
We recorded underwater video on eastern oyster farms with high (dense) and low (sparse) cage density versus boulders on a rock reef. The three subtidal study sites were located west of Charles Island, near Milford Connecticut, USA, within central Long Island Sound (northwest Atlantic; Figure 1). Detailed methods and fish abundance were previously reported in Mercaldo-Allen et al. (2023).
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The dense cage farm site, with 40–100 commercial cages, was located on a 0.11 km2 shellfish lease, permitted for up to 200 multi-tiered off-bottom cages. An adjacent 0.25 km2 lease, co-located at the dense cage farm site, was permitted for up to 250 off-bottom cages. Both were working commercial farms where gear was intermittently deployed, hauled for maintenance and/or removed for harvest, hence the number of cages in the vicinity varied on any particular day and across the spring to fall study period and limited our ability to test the effect of specific cage density itself. We placed four study cages at the dense cage farm site, spaced 47.5 m apart.
The sparse cage farm site was located at the intersection of two large shellfish leases, each 0.20 km2 in size, where traditional on-bottom oyster aquaculture with no off-bottom gear was underway. Here, the sand and shell bottom were characterized by low vertical relief, contained areas of live oysters and empty shell valves, and was devoid of rocks or boulders. We placed five study cages in a line along the delineation between the two on-bottom aquaculture leases, at a distance of 82 m apart. This discrete ‘farm’ of five study cages was designated a ‘sparse’ farm, relative to the five study cages on the ‘dense’ actively worked commercial farm, where as many as 100 cages were present within the study area. The rock reef site, containing 70% cobble and boulder substrate, served as a structured reference site for natural hard bottom habitat. The reef was horseshoe-shaped, patchy and covered 0.25 km2 of seafloor. We selected four boulders on the reef for study, located a minimum of 10 m apart and out of visual range of one another. Boulders were selected as our reference site as rock reefs are the natural structured seafloor most utilized by temperate reef fish in this region.
Distances between the three study sites (dense farm, sparse farm and rock reef) ranged from 745 to 1537 m (Mercaldo-Allen et al. 2020). Water depths at high tide measured 4.6 m at the dense cage farm, and 6.1 m at the sparse cage farm and rock reef sites.
Oyster Aquaculture Cages, Camera Deployments and Recording Methods
We studied commercially-available shelf and bag style oyster aquaculture bottom cages (Ketcham Supply, New Bedford, Massachusetts, USA) constructed of 11.43 cm mesh, heavy duty 8-gauge vinyl coated wire, with 2 reinforced 3.81 cm wire mesh feet. Wire mesh feet elevated cages 15.2 cm off the seafloor, with 10 bricks added to each cage foot for ballast. Cages measured 1.22 × 0.91 × 0.61 m3, and had three shelves, each holding 2 bags of oysters (6 bags per cage). Bags, made from 2.3 cm plastic mesh, measured 1.07 m long by 0.52 m wide. We stocked each cage with 150–200 seed oysters measuring 2.5 to 4.5 cm in size. Cage handling methods and oyster stocking densities reflect general industry practices.
Methods for deployment and retrieval of action cameras on oyster cages and adjacent to boulders are detailed in Mercaldo-Allen et al. (2021, 2023). Briefly, cages were brought onto the boat deck where we attached two cameras to each of four study cages, one camera positioned to record across the cage top and the other with a view across two cage sides and the interface between the cage and the seafloor (Figure 2a). We also constructed T-platform stands to mount cameras among boulders that provided a similar perspective to cage-mounted cameras and minimized added structure to the rock reef site, as described in Mercaldo-Allen et al. (2021, 2023). Divers attached two cameras to each of four T-platform mounting stands placed adjacent to the four study boulders, with one camera positioned to record across the top boulder surface while the other camera captured the side of the boulder and the boulder-seafloor interface (Figure 2b).
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Video recording followed methods outlined in Mercaldo-Allen et al. (2021, 2023). Briefly, we programmed time-synched GoPro Hero Silver 3+ cameras to record video at 30 frames per second, 1920 × 1080 resolution 10 megapixel, with a wide-angle lens (firmware v03.02). We placed cameras inside a polycarbonate waterproof case with BacPac attachment which accommodated a timer. Each camera lens was fitted with a Polar Pro magenta filter, with a 0.5 stop reduction in exposure, to reduce natural green coloration in video. Intervalometer Blink timers (CamDo), paired with GoPro cameras, were used to delay onset of video recording and extend battery life. Cameras were unbaited. Video recording began approximately 24 h after cameras were deployed, which was intended to reduce deployment-related disturbance effects on fish behaviour. Video was recorded for 8 min every hour from 7 AM to 7 PM, yielding 13 recordings per camera deployment, to collect footage over a complete tidal cycle and most daylight hours.
We collected video on the dense farm during all weekly camera deployments over the 17-week study period from May to September 2018. We recorded during alternate weeks at the other two sites, due to logistic constraints, with eight deployments on the sparse farm and nine deployments on the rock reef. Analysis of these videos for fish abundance and community composition, as well as measurements of current speed (cm s−1), light illuminance (lumens m−2), seawater temperature (°C) and salinity (PSU) at the three study sites, were reported previously (Mercaldo-Allen et al. 2023).
Video Analysis of Fish Behaviour
The behavioural coding software Observer XT (v14.2 and 15.0; Noldus Information Technology) software was used for reviewing and scoring time-synched video that was recorded simultaneously on cages and boulders with top and side cameras. Water clarity varied between deployments and among hours within a single deployment, however visibility generally extended the full length of the cage/boulder. To minimize the effect of variable water clarity across videos, only fish within the immediate cage or boulder vicinity were included, regardless of the total extent of field of view within any single video clip. Video footage was also reviewed to qualitatively describe the epifaunal community colonizing cages and boulders, as reported in Mercaldo-Allen et al. (2023).
Fish within the immediate vicinity of the cage or boulder were positively identified visually using morphological features (e.g., Weiss 1997; Collette and Klein-MacPhee 2002) and swimming behaviour. Small-bodied cunner are olive-grey to brown in colouration with a more pointed snout than young black sea bass or tautog. They have a rounded caudal fin, a horizontal stripe through the eye, and have a false eyespot at the anterior of the dorsal fin. Although cunner have no sex-specific morphological differences, spawning males can exhibit a blue-phase colouration, while females and immature males show no colour changes relative to reproductive status or maturity (Pottle and Green 1979a, 1979b). Tautog can be distinguished from cunner by a stouter body shape, pronounced lips and a blunt snout. This species exhibits sexual dimorphism, where juveniles and females have mottled grey to black colouration on a pale olive to brownish or grey body and large adult males are dark grey to black with a distinct white chin, blunt forehead and white blotches on their flanks. In addition, there are non-dimorphic males which are also sexually mature but exhibit similar external coloration to female tautog (Olla and Samet 1977; Hostetter and Monroe 1993). Subdued colouration in small males, females and juvenile tautog may serve as camouflage while more contrasting colouration in large sexually mature males may aid in attracting mates or establishing territorial dominance (Olla et al. 1981; Horodysky et al. 2013). Large mature females may exhibit a white saddle or striped pigmentation on their mid-flanks in association with reproductive readiness (Olla and Samet 1977). Scup have a slender, copper or silver appearance, and are deep-bodied, with a forked tail and often irregular dark vertical bars on their body and are not sexually dimorphic (Collette and Klein-MacPhee 2002). These distinct differences in physical morphology and colour pattern made it possible to accurately distinguish among these species, sexes and life-history stages in video.
Fish activity was scored using a behavioural ethogram (Table 1). Behaviour categories were selected based on relevance to habitat provisioning and included courtship and reproduction (males pursue gravid females, swimming close together or touching, release of gametes), escape from predators (rapid retreat into structure when pursued by predators or aggressively challenged by other fish), foraging (picking and grazing on cage surfaces, lines, rigging, mesh bags and boulder surfaces), sheltering (hiding inside cages and bags, station keeping using small fin movements to hold place in currents above or alongside cage or boulder), schooling or grouping (moving together in a synchronized school or forming loose aggregations above or alongside the cage or boulder) and territoriality (agonistic aggressive chasing or biting, fin flaring). Fish transiting through the farm or reef was commonly observed but not quantified, as this behaviour did not relate directly to habitat use.
TABLE 1 Ethogram describing behavior of cunner, scup and tautog observed on oyster cages at shellfish farms and adjacent to boulders on rock reefs.
| Courtship and reproduction | Flanks or tail touching, swimming in close proximity, spawning runs, release of gametes |
| Escape from predators | Fish retreats into structure associated with a predator or another fish |
| Foraging | Fish picking food (out of water column, off lines/bags/cages) |
| Schooling or grouping | Shoaling, fish of the same species aggregating in numbers of two or more |
| Sheltering |
Fish enters the structure. Not predator/territory associated. Station-keeping or resting, using small fin movements to maintain position. |
| Territoriality | Active defence of space or resources. Fish chases another from an area, displacement, agonistic displays, fin flaring, yawning |
A daily index was calculated for each behaviour (courtship/reproduction, escape, foraging, grouping, shelter and territoriality) for each species by summing events across all video recorded: 8 min per hourly interval × 13 h = 104 total min per day. This daily index was calculated for each cage/boulder replicate at each study site. The daily indices were normalized to the daily average MaxN abundance for each replicate/date combination, to remove the effect of fish abundance on occurrence of behaviour events. MaxN abundance, defined as the maximum number of fish of a given species present in a single frame within each 8-min video segment, was reported previously (Mercaldo-Allen et al. 2023). Normalized daily indices for each behaviour were then compared across the sampling time series. A global repeated measures analysis was first performed to compare fish behaviour between habitats; that is, two cage farms considered replicates and compared to the single rock reef site. When global tests were significant, pairwise comparisons across study sites were performed. As all three study sites could not be sampled simultaneously, data were pooled by month prior to analysis. The repeated measures analysis used marginal distributions and a percentile bootstrapping method, with 20% trimmed means as the measure of central tendency, as this approach has no assumptions of normality or homoscedasticity, which are both common problems in ecological data (Wilcox 2023). Hochberg's sequentially rejective approach was used to control familywise error within pairwise comparisons (Wilcox 2023). All statistical analysis was conducted using the software program R version 4.4.3 ().
Results
Behaviours associated with habitat provisioning (e.g., courtship/reproduction, escape from predators, foraging, sheltering, schooling/grouping, territoriality) were observed for all three fish species on both cages and boulders. Generally, nearly all of these behaviours occurred more frequently on oyster aquaculture cages as compared to boulders. We quantified a total of 5679 behavioural events in this study, inclusive of 2395 for cunner, 1420 for scup and 1864 for tautog.
For cunner, sheltering and foraging were the two most common behaviours. Significantly more sheltering behaviour (p = 0.0088, Figure 3a) was observed on cage versus boulder habitat. Sheltering was significantly higher on the dense (p < 0.0001) and sparse farms (p < 0.0001) compared to the rock reef. No difference in sheltering was noted between the two farm sites (p = 0.56). Significantly more foraging (p = 0.028, Figure 3b) was observed on cage versus boulder habitat. Foraging was significantly more frequent on the dense (p = < 0.0001) and sparse farm (p = 0.0038) compared to the rock reef. Foraging activity was significantly higher on the dense versus sparse farm (p < 0.0001). Territorial behaviour was significantly higher on the boulder versus cage habitats (p = 0.0024, Figure 3c) and occurred more frequently on the rock reef compared to the dense and sparse farms (p = 0.023, p = 0.0034, respectively). No difference in territoriality was detected between the two farm sites (p = 0.55). No significant difference in escape from predators (p = 0.38, Figure 3d), grouping (p = 0.73, Figure 3e) or courtship/reproduction (p = 0.82, Figure 3f) behaviours were detected between cages and boulders; since the global test of habitat type was not significant, pairwise comparisons of sites were not performed. Behavioural events observed in cunner for each of the six categories (escape, foraging, grouping, reproduction, shelter, territoriality) summed by sampling date and study site are shown in Table S1.
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For scup, sheltering, foraging and grouping were the most frequently observed behaviours. Significantly more sheltering activity was observed on cage versus boulder habitat (p < 0.001, Figure 4a). Significantly more sheltering was observed at the dense (p = 0.0038) and sparse farms (p = 0.0072) when compared to the rock reef. No difference in sheltering was noted between the two farm sites (p = 0.80). Significantly higher instances of foraging (p = 0.0022, Figure 4b) were observed on cage versus boulder habitat. Significantly more foraging was observed at the dense (p = 0.0038) and sparse (p < 0.001) farms compared to the rock reef. There was no difference in foraging between farm sites (p = 0.63). Territoriality was observed infrequently with no significant difference detected between the boulder and cage habitats (p = 0.79, Figure 4c). Significantly higher escape activity was observed on cage versus boulder habitat (p = 0.0082, Figure 4d). Higher frequency of escape was observed on the dense farm compared to the rock reef (p < 0.001). There was no significant difference in escape between the sparse farm and the rock reef after controlling for multiple comparisons (p = 0.042; adjusted α 0.025) or between the two farm sites (p = 0.97). There was no significant difference in grouping (p = 0.11, Figure 4e) or courtship/reproductive activity (p = 0.68, Figure 4f) between cage and boulder habitats (Figure 4f). Behavioural events observed in scup for each of the six categories (escape, foraging, grouping, reproduction, shelter, territoriality) summed by sampling date and study site are shown in Table S2.
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For tautog, sheltering and foraging were the most commonly observed behaviours. All six of the quantified behaviours were significantly more frequent on cage versus boulder habitats, including sheltering (p = 0.005, Figure 5a), foraging (p < 0.001, Figure 5b), territoriality (p < 0.001, Figure 5c), escape from predators (p < 0.001, Figure 5d), grouping (p = 0.019, Figure 5e) and courtship and reproduction (p = 0.015, Figure 5f). Significantly more courtship and reproduction, escape from predators, foraging, sheltering and territoriality occurred at the dense farm than on the rock reef (all p < 0.001). Significantly more escape from predators (p < 0.001), foraging (p = 0.004), grouping (p < 0.001), courtship and reproduction (p = 0.038), sheltering (p = 0.015) and territoriality (p < 0.001) occurred at the sparse farm compared to the rock reef. There was no significant difference in grouping behaviour between the dense farm and the rock reef (p = 0.12). There was no significant difference between the dense and sparse farms for escape (p = 0.76), foraging (p = 0.78), grouping (p = 0.75), courtship/reproduction (p = 0.20), sheltering (p = 0.90) and territoriality (p = 0.54). Behavioural events observed in tautog for each of the six categories (escape, foraging, grouping, reproduction, shelter, territoriality) summed by sampling date and study site are shown in Table S3.
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Qualitative assessments of colonizing organisms from video recorded on the cage farms and rock reefs were used to describe habitat attributes of the cages and boulders, as previously reported (Mercaldo-Allen et al. 2023). Visual inspection of video from cages noted growth of colonizing organisms on cages, bags, lines and rigging at the dense and sparse cage farms. The two visually dominant species were the colonial hydroid Campanularia spp. and lacy crust bryozoan Membranipora membranacea. No algae were observed. Cages had light to moderate colonizer density with up to 80% coverage by late summer. Boulders on the rock reef were sparsely colonized by Campanularia spp. and the spiral-tufted bryozoan Crisularia turrita.
Discussion
We observed cunner, scup and tautog demonstrating a variety of behaviours on oyster cage farms associated with habitat provisioning, including courtship and reproduction, escape from predators, foraging, grouping or schooling, sheltering, and territoriality, similar to that observed on rock reef habitats. Interactions of reef fish with their environment are highly influenced by the structural complexity and spatial arrangement of habitat. Although cage farms share similar physical attributes with rock reefs (DeAlteris et al. 2004; Lefcheck et al. 2021), cages and boulders differ spatially and dimensionally in the configuration of the structure they provide to fish and these differences influence how these structures function as habitat. Observations of fish behaviour associated with oyster aquaculture cages on farms and boulders on a rock reef provide insights into the functional role provided by these habitats to temperate reef fish species.
Sheltering
Sheltering behaviour was demonstrated by all three temperate reef species on oyster cages and boulders. Availability of suitable shelter influences carrying capacity and population size in cunner and tautog (Olla et al. 1974; Auster 1987, 1989). These species rely on shelter to avoid competition with conspecifics, conserve energy in high current flow and for protection during periods of low activity, night-time torpor or in winter, often situating themselves alongside or under an object (Dew 1976; Olla et al. 1974, 1980; Auster 1987; Tupper and Juanes 2017). Small-bodied cunner and young tautog remain close to the home site, with generally localized movements (Olla et al. 1974; Nitschke et al. 2002) and utilize shelter for protection from predation (Sharf et al. 2006). Scup and tautog frequently demonstrated station-keeping, using small fin movements to maintain position on the upper surface or inside cages. This sheltering behaviour provides fish with respite from active swimming during high current flow, and may result in energetic savings and increased opportunities for feeding (Gerstner 1998). Interestingly, we also observed a school of adult yellow jack, a southern species, station-keeping above a cage across several recording intervals on three occasions during August and September.
We documented a higher frequency of sheltering behaviour in cunner on cages than boulders. Use of cages as shelter by cunner suggests that aquaculture gear provides this species with high quality structured habitat (Nitschke et al. 2002) much like rock reefs. Higher association of cunner with oyster gear but not adjacent marsh habitat in Barnegat Bay, New Jersey, suggested that cages may have provided a structural component absent from the structured marsh edge (Shinn et al. 2021). Cunner was also highly associated with farm gear in Great Bay, New Hampshire where hard bottom substrate was absent (Glenn 2016). The variably sized openings and crevices in cages and boulders create an assortment of interstitial spaces for small cunner while the epifaunal community on surfaces contribute refuge and cover (Hales and Able 2001; Nitschke et al. 2002). Once entering the cage, cunner often swam into the individual bags of oysters, with the smaller mesh size of the bags offering additional protection from predators. Occasionally, cunner initiated a flashing behaviour, rubbing their body against the cage surface, perhaps in an attempt to remove parasites, scratch an itch or remove loose scales. On cages and boulders, cunner often swam slowly and continuously, remaining in the same vicinity throughout the entire video recording. Notably, cunner were almost always in motion, with limited observations of station keeping.
Tautog demonstrated sheltering behaviour more often on cages than on boulders and this was consistently noted throughout the season. Tautog could be seen utilizing space under, within, and alongside cages as well as on the cage top. Fish of all life stages were observed to rest for varying lengths of time on the upper cage surface. Large males and gravid females were observed resting for extended periods on the cage top, possibly recovering from reproductive activity (Olla and Samet 1977). Tautog sometimes laid on their side atop the cage, swaying in the current. There were instances of large tautog sitting on the cage top engaged in buccal pumping, where a fish opens and closes its mouth to bring water in and over the gills, an activity related to gill ventilation among teleost fishes (Ferreira and Wilson 2024). Cages appeared to protect juveniles and females from large reproductive males. Fish also sometimes remained stationary beneath cages. On the reef, adult, juvenile and young of the year tautog all occasionally sheltered alongside or atop of a boulder. In July, a large tautog was observed nested between boulders within the colonizing community. For shelter-oriented tautog, addition of complex structure that creates new habitat may contribute to increased fish production (Olla et al. 1974).
Frequency of sheltering behaviour in scup occurred consistently across the summer months and was more common on cages than boulders. Of the three temperate reef species, scup most often demonstrated station keeping behaviour, sometimes hanging in the water column alongside the cage, hiding underneath or within the cage, or commonly on the cage top. Access to structured habitat is thought to provide necessary shelter for scup, particularly at the early life stages (Steimle et al. 1999; Mercaldo-Allen et al. 2023). We frequently observed young-of-the-year scup entering cages through the top or side wire mesh. Although grouping was more common on cages, we did observe a school of seventeen young-of-the-year transiting through the boulder area.
Shellfish aquaculture gear has been previously shown to create shelter for wild fish. In Australia, sheltering behaviour in fish occurred more frequently around oyster farms than on unstructured seafloor and fish generally passed through low relief areas without interacting with habitat (Martínez-Baena et al. 2022). A number of fish species at a variety of life stages demonstrated sheltering seeking behaviour in and around oyster gear on farms in Barnegat Bay, New Jersey (Shinn et al. 2021). Trestles used for oyster cultivation in France were preferred resting locations for sole (Solea solea), which sheltered beneath these structures (Laffargue et al. 2006). In related video studies in Long Island Sound, we found that the open shelves and interstitial spaces characteristic of shelf and bag style oyster aquaculture cages shelter temperate reef fish species over a range of body sizes and across life history stages (Armbruster et al. 2024; Mercaldo-Allen et al. 2021, 2023, 2025). Anecdotally, we observed a wider variety of fish sizes on the farms relative to the reef, likely a reflection of the more complex architecture of the shelter afforded by cages relative to boulders.
Foraging
For shelter-oriented species that remain in close contact with structure, availability of abundant high-quality forage increases habitat value (Brown et al. 2022). Epifaunal organisms that colonize shellfish aquaculture gear can represent a food source for fish (Veggerby et al. 2024). Cunner, scup and tautog employ a scan-and-pick feeding modality (e.g., Olla et al. 1975; Dew 1976; Auster 1987) that makes them suited to consuming organisms on the surfaces of cages and boulders. The open framework of shelf and bag cages enable fish access to interior spaces that provides extensive surface area for epifauna while on rock reef areas, solid boulders are often thickly settled with colonizers. Both cages and boulders were inhabited by similar colonizing organisms that created cover for fish and their prey. These emergent communities harbour a variety of potential prey including hydroids, bryozoa, molluscs, small crustaceans and fish that constitute key diet components for these temperate reef fish in Long Island Sound (Olla et al. 1975; Bowman et al. 1987; Clark et al. 2006).
Foraging behaviour in cunner was more frequent on cages than boulders. We observed cunner pecking at the colonizing community covering cages, rigging and lines and boulder surfaces and plucking food particles from the water column. Cunner feed opportunistically on planktonic, epibenthic or infaunal species, e.g., small molluscs and crustaceans, on the seafloor, on rock surfaces and from the water column (e.g., Collette and Klein-MacPhee 2002; Olla et al. 1975; Shumway and Stickney 1975; Whoriskey 1983; Auster 1987). Cunner will also dart into currents to ingest food or attack prey and then return to cover, perhaps as a means to conserve energy (Olla et al. 1975; O'Brien et al. 2018). This strategy allows cunner to effectively forage in areas with high tidal flow (Auster 1987). We frequently observed cunner grazing on cages and boulder surfaces in groups of two or more.
Tautog foraging activity occurred at higher frequency on cages than boulders. Observations of feeding activity on the farms and the rock reef trended upward during the summer, perhaps reflecting an increase in food availability as colonizers became more established on cages and boulders over time. Adult, juvenile and young-of-the-year tautog pecked at the interior and exterior cage surfaces, on mesh oyster bags, cage lines and rigging and occasionally on growth covering camera mounts. Juvenile tautog bit at the knots in cage lines and consumed food passing by in the water column. Tautog are considered feeding generalists, consuming a variety of colonizing and encrusting organisms (e.g., bryozoans and hydroids), benthic molluscs and small crustaceans (Clark et al. 2006; Taylor et al. 2023). Over the course of ontogeny, tautog develop strong pharyngeal jaws that enable them to consume hard shell prey, including mussels and crabs (Dorf 1994; Clark et al. 2006). On one occasion, we observed an adult male tautog swimming around the cage while consuming a large spider crab. On the reef, tautog were occasionally observed foraging on benthic organisms where boulders met the seafloor, as well as on hydroids and bryozoans atop and alongside boulders.
Scup foraged more frequently on cages than boulders and the highest feeding activity was noted in June and July at the dense farm. Juvenile, adult and young-of-the-year scup picked at the epifaunal community that colonized cages, lines and rigging and covered boulder surfaces. Opportunistic and visual benthivores, scup browse on a wide variety of small invertebrates and occasionally, small fish (e.g., Bowman et al. 1987; Morse 1978; Michelman 1988; Steimle et al. 2000; Byron and Link 2010). Field observations by divers have documented scup using scan and pick feeding to consume prey from rock surfaces (Auster 1987). At the rock reef, scup occasionally foraged on boulder substrate. We frequently observed large schools of scup at all life stages feeding on and above cages and also documented numerous instances where a lone adult scup grazed on cages or lines. Scup, which are found on both sandy and rocky substrate (Steimle et al. 1999; Mercaldo-Allen et al. 2020), may also feed along edge habitat at the border between structured cage farms and low relief seafloor (Sambrook et al. 2016; Mercaldo-Allen et al. 2021, 2023).
The role of shellfish farm infrastructure as forage habitat for fish has been documented worldwide. On Australian oyster farms, foraging occurred more frequently on aquaculture gear than unstructured seafloor and the feeding activity associated with gear was similar to use of natural biogenic habitats (Martínez-Baena et al. 2022). Video surveillance around Japanese oyster rafts found black sea bream feeding on the sessile organisms growing on shellfish gear (Tsuyuki and Umino 2018). In Washington State, Veggerby et al. (2024) found that oyster farms provided foraging habitat for a variety of fish species and that aquatic epiphytes attached to gear may concentrate invertebrate prey. Sessile biofouling organisms that represent the base of the food web for fish may serve a similar ecological role on aquaculture gear (DeAlteris et al. 2004).
Composition and extent of epifaunal communities that provide forage can vary seasonally or across years. We qualitatively noted thicker colonization of cages and boulders in video footage recorded in 2017 as compared to 2018 (Mercaldo-Allen et al. 2021, 2023) and increasing epifaunal coverage of structures was noted over the course of the summer season in 2018. Active feeding on farms and the rock reef was observed among all three fish species, regardless of the recording month or the extent to which cages and boulders were colonized.
Territoriality
Possession of territory is a component of successful mating for both cunner and tautog (Martel and Green 1987; Auster 1989), as well as other reef species (e.g, Reebs 2008; Barneche et al. 2009; LaManna and Eason 2011; Armbruster et al. 2024). In this way, dominant male fish secure access to preferred feeding, shelter and spawning habitat (Gibson 1993).
Interestingly, territorial behaviour in cunner was the only behaviour more common on boulders than cages. Boulders are preferred habitat for cunner (Auster 1987, 1989) and one they appear to actively defend. Conversely, multi-dimensional cages may contain a higher abundance of interstitial shelters than solid boulders, potentially reducing the need for territorial interactions. However, we frequently observed larger cunner chasing conspecifics and small individuals of other species away, in defence of both cages and boulders. Interspecific aggression between cunner and similarly-sized tautog and mummichogs is common, and adults are known to nip and chase these species away from habitat (Able et al. 2005). Intraspecific competition among cunner was frequent in June. On one occasion, we observed as many as sixteen cunner exhibiting territorial chasing and nipping on the reef. In supplemental video recorded on the dense farm in June 2019, a group of six cunner participated in aggressively charged interactions above a cage, swimming at one another, chasing and nipping, and two individuals repeatedly engaged in face-to-face biting (Clark & Phillips, pers. commun.). In August, a pair of cunner exhibited a high level of aggression within the cage structure, demonstrating eight instances of nipping within a 1-min period.
Territoriality in tautog occurred more frequently around cages than on boulders, with activity highest in June and July. Aggressive activity in tautog was similar to seasonal patterns we observed in courtship behaviour. We found that sexually dimorphic males spent much of their time circling around and beneath the cage. These males displayed abrupt and swift bursts of swimming, often heading directly toward a gravid female or nipping at smaller black sea bass, scup or tautog, and on one occasion, a large striped searobin (Prionotus evolans). Territorial males appeared to patrol the cage, apparently guarding females inside. In late May and June, we saw several examples of a persistent male repeatedly and unsuccessfully trying to enter a cage, presumably to displace another fish or reach a female. One adult male tautog swam sideways at a cage, perhaps an attempt to gain entry to the interior. Another large male swam out from beneath the cage at high speed and aggressively tried to enter at the cage side, biting on the wire mesh and thrashing violently. One large tautog rammed the cage multiple times, moving it, while in pursuit of a juvenile black sea bass that retreated inside. These males were too large to pass through the square wire-mesh framework of the cage. Aggressive encounters continued into July. While chasing away other fish, one large male hit one camera so hard that it moved the entire cage. Another large male was observed sticking his head into an opening in the square mesh at the cage top. In one instance, an adult male kept chasing a smaller fish, no matter where it relocated around the cage. One large male tautog swam at a group of smaller black sea bass which dispersed into and under the cage. Interestingly, we also saw one precocious smaller tautog chase a larger male away from the cage.
Previous laboratory studies with tautog found that most behaviour outside of courtship and spawning involved the dominant male tautog aggressively chasing or biting the subordinate male to limit his access to the female or resources (Olla and Samet 1977). Similarly, observations of pairs of juvenile tautog in holding tanks found that the larger fish showed dominance by chasing and nipping at the flanks of the subordinate whenever it approached (Olla and Studholme 1975). As we also observed, peak aggression and territorial activity in tautog coincides with elevated summer seawater temperatures and decreases as water temperatures decline seasonally (Olla et al. 1978, 1980; Arendt, Lucy, Evans, et al. 2001; Arendt, Lucy, Munroe, et al. 2001).
Scup showed limited territoriality with no detectable difference in this behaviour between cages and boulders. Scup are a gregarious shoaling species thought to associate for safety and to increase prospects for feeding and locating mates (Morse 1978). We observed schools of young-of-the-year, adults and mixed schools of multiple life stages swimming around or feeding on the upper surface of the cages. Instances of territorial activity among scup showed a slight uptick as autumn approached, perhaps related to changes in behaviour related to seasonal migration to deeper waters (Thomson et al. 1978). In August, we observed one adult scup chasing another off the cage top and in September, we saw a single instance of a young-of-the-year chasing a conspecific, but this behaviour was rare.
Escape from Predators
Structural complexity is critical to the ecology of temperate reef fishes and is especially important for fish that are small-bodied or at an early life history stage. Shelter-rich environments contribute to greater habitat suitability and decreased predation risk for small species such as cunner (Tupper and Juanes 2017). Complex habitats with interstices exclude large-bodied predators from access to interior spaces and can impede predatory fish who rely on high-speed pursuit to apprehend prey (DeAlteris et al. 2004; Scharf et al. 2006). Shelf and bag oyster cages are constructed of a stiff metal frame with a mesh size that prevents large predators from entering, making the cage interior an effective refuge from predation. Black sea bass, highly associated with oyster cages (Mercaldo-Allen et al. 2023), was the dominant predator during our study and the most aggressive species we observed pursuing smaller cunner, scup and tautog on the farms. Increasing black sea bass abundance in southern New England waters may be influencing interspecies competition and affecting predatory interactions among temperate reef fish (Mercaldo-Allen et al. 2020; Taylor et al. 2023). Intraspecies encounters, where smaller cunner or tautog were chased into cages by larger fish of the same species, was also frequently observed.
Escape from predators occurred at a similar frequency among cunner on both cages and boulders. On the reef, cunner moved in and out of the colonizing community on boulder surfaces as a refuge from other fish. In June, we observed one instance of several cunner actively moving away from a summer flounder as it swam off the top of a boulder. During August, we observed cunner, pursued by black sea bass or tautog, successfully taking refuge in the cage interior and within the oyster bags. Anecdotally, we noted that cunner were generally less visible in video footage when black sea bass were present.
Escape behaviour among tautog was more frequent on cages versus boulders and showed similar patterns on the two farms across the season. We frequently saw young-of-the-year tautog being chased by young-of-the-year black sea bass, and successfully escaping into cages. In September, a young tautog entered the cage through the top mesh to escape a large predatory striped bass, which circled the cage five times before swimming off. Diver observations along the coast of Long Island have similarly observed young tautog using crevices within natural habitat to escape pursuit by striped bass (Olla et al. 1974). Throughout the recording period, we also saw instances of fish retreating quickly into cages in the absence of a visible predator. Although we occasionally glimpsed predatory species (bluefish, striped bass, weakfish) swimming above the cage top, positioning of cameras may have precluded detection of all large predators circling the waters around the cage or boulder, outside the camera field of view.
Scup demonstrated significantly more instances of escape at the dense farm than on the rock reef, but notably, escape behaviour was similar between the sparse farm and rock reef. In May and July, we observed larger black sea bass attempting to attack young-of-the-year or juvenile scup and in August, saw a school of young-of-the-year scup actively pursued by a large striped bass. One adult scup with a noticeable bite mark was associated with one of the cages, suggesting it may have survived a predator encounter.
Schooling or Grouping
Cunner and tautog are known to occur in loose aggregations associated with structured habitat and reproductive activity (e.g., Collette and Klein-MacPhee 2002; Olla et al. 1975; Martel and Green 1987; Auster 1989) while scup is a shoaling species which socially congregate to locate food, find mates and avoid predation (Morse 1978). We found that oyster farms were at times populated by large schools of scup, smaller groups of cunner and tautog and multi-species aggregations, suggesting that cages provided sufficient habitat resources to support large numbers of co-occurring individuals and species at multiple life stages. Abundance of fish demonstrating schooling or aggregating behaviour varies seasonally related to factors such as food availability and life history stage (Pitcher and Parrish 1993). We noticed larger assemblages of scup and tautog later in the season, when young-of-the-year appeared and cages and boulders contained more biofouling forage on surfaces.
Cunner grouping behaviour was similar on cages and boulders. Fish occurred individually or in small groups on the farms and occurred in larger aggregations on the rock reef (Mercaldo-Allen et al. 2023). Cunner often formed groups, especially if there was abundance of food, staying together perhaps as protection from predation. Fish also aggregated in association with courtship and spawning activities early in the summer.
Grouping behaviour among tautog was generally higher on cages than boulders, however, grouping did not differ between the dense farm and rock reef. Tautog demonstrated little grouping during May and June, which coincides with peak spawning activity and territorial behaviour (Olla et al. 1980). Large male tautog were frequently observed chasing groups of smaller tautog or black sea bass off a cage. Sexually mature tautog remained solitary during early summer spawning but later in the season, when reproductive activity declined, juveniles and adults occurred more frequently in loose aggregations on the rock reef and around cages on the farms. In August and September, we observed groups of four to six fish at times moving over, under, into and alongside cages without aggressive or courtship interactions. Previous observations of tautog held in aquariums noted grouping behaviour taking place within structure during periods of colder temperatures, when fish are less active (Arendt, Lucy, Evans, et al. 2001).
Schooling among scup occurred throughout the sampling season, and at a similar frequency on cage farm and rock reef habitats. Scup at all life stages, including young-of-the-year, commonly schooled on the upper cage surface. Numbers varied from 2 to 3 individuals to as many as 80 fish per school. In May and June, groups of 2–3 juvenile or adult scup occurred around cages. Although scup generally form schools with fish of the same size (Morse 1978), we observed mixed schools of young-of-the-year, juveniles and adults in August and September. Smaller scup appeared to gain benefit from schooling on farms with larger adult fish. Schooling activity on the rock reef increased late in the season with the influx of young-of-the-year. Single adult scup occurred occasionally around cages but young-of-the-year exclusively travelled in large schools. The architecture of multi-dimensional oyster cages appeared to offer sufficient complexity to support the functional needs of these aggregations of scup.
Courtship and Reproduction
We found that cunner courtship and reproductive behaviour occurred at a similar frequency on cages at the two farms and boulders on the rock reef, taking place predominantly in late May, June and July. We observed paired courtship on farms, where a male cunner pursued a gravid female by darting and chasing. Gravid female cunner were easily identified by their broad girth relative to male fish. One male/female pair swam upward together out of view, returning a few minutes later, chasing one another across the cage top, until swimming out of view. In another instance, a gravid female and male swam towards each other on top of the cage. The male then paused and made three passes towards the female, before aligning in parallel, making full body contact for 2 s and then exiting the field of view. In supplemental video collected on the dense farm in 2019, male and female cunner were seen circling and intermittently touching one another above a cage, then aligning side by side and swimming upward and out of view (Clark & Phillips, personal observation). Another set of male and female cunner swam upward together in what appeared to be a spawning run. A third cunner from across the cage attempted to join them, but then returned immediately to the cage surface. This interference spawning, where nonterritorial sneaker males joined a mating pair during the upward spawning rush, has been previously documented in cunner (Pottle et al. 1981; Martel and Green 1987). On the rock reef, we also observed two instances of what appeared to be spawning aggregations of cunner, where two sets of paired fish swam together upwards and out of view.
Diver observations at field locations in New England and Canada have observed territorial male cunner engaging in pair spawning and smaller nonterritorial males participating in group spawning, with activity levels also peaking in June and July (e.g., Wickland 1970; Pottle and Green 1979a, 1979b, 1981; Martel and Green 1987; Auster 1989). Successful courtship and spawning in cunner relies on establishment and defence of territory by dominant males (Martel and Green 1987). Similar to our observations, these studies found cunner courtship and spawning activity to be characterized by chasing and darting in paired or small groups, followed by vertical swimming to a few meters above the seafloor, with close contact or physical touching of fish and subsequent release of gametes (e.g., Wickland 1970; Pottle and Green 1979b; Pottle et al. 1981; Martel and Green 1987). Although release of gametes by cunner was not captured in our videos, we observed courtship activity followed by upward swimming, suggesting that spawning likely took place above cages or boulders, beyond our camera view.
We found the frequency of tautog courtship and reproductive activity to be higher on cages as compared to boulders. These behaviours occurred most often in the late afternoon over the period May through mid-August. Courtship behaviour near cages was observed between discrete pairs of male and female fish and among small groups of tautog. Previous laboratory and field studies have shown that spawning occurs in heterosexual pairs or as a group with a single female and multiple small males (e.g., Olla and Samet 1977; Olla et al. 1981; Dixon 1997). Smaller males without the sexually dimorphic characteristics of large tautog (e.g., prominent forehead and pronounced jaw), may be less able to compete directly for females (Olla and Samet 1977; Auster 1989). These non-dimorphic males may achieve reproductive success through interference spawning alongside a male and female pair or in a spawning group (Olla and Samet 1977; Hostetter and Munroe 1993). We also saw large males chasing off what appeared to be these younger ‘sneaker’ males that were attempting synchronized swimming with a gravid female.
Courtship behaviour was most conspicuous in large sexually dimorphic male tautogs, which spent much of their time patrolling and circling cages. Males were frequently observed moving rapidly toward a gravid female and/or pursuing her around the cage. These rushes were previously described in laboratory studies by Olla and Samet (1977), where a male tautog courted the female by rushing toward her at speed without displacement, followed by parallel swimming in what appeared to be a spawning run. We commonly observed one to two smaller tautog, apparently females, swimming into a cage while a male circled just outside. Smaller males were also observed trying to join a large male/female pair in a spawning run. On several occasions in June and July, we observed three to four tautog swimming in close proximity, around and under cages with flanks touching, the female positioned between the males. Under laboratory holding conditions, male and female pairs were seen engaging in close physical contact, with nuzzling or rubbing of flanks (Bridges and Fahay 1968) and in group spawning, where males move alongside females with flanks touching (Olla and Samet 1977). Notably in July, a male and female tautog were observed swimming upward from the cage in unison, the female adopting a curved body posture, and the male keeping pace close alongside. Another male and female tautog swam nose to tail at the cage side, male behind the female. During early August, we observed a single instance of spawning, when a cloudy substance, likely sperm, appeared to be released by a tautog just above a cage.
We noted little obvious courtship behaviour in scup. A single instance of scup reproductive behaviour occurred in early August, where an adult swam upward in front of a cage on the dense farm, releasing several small clouds of what appeared to be sperm. In preliminary video collected at the dense farm during 2017, an adult scup also released gametes, likely sperm, as it swam along the cage side directly in view of the camera (Clark & Phillips per. observation 8/2017). A substance that appeared to be floating eggs was also observed. To our knowledge, these instances are the first spawning events visually documented for scup in Long Island Sound.
Shellfish farming areas have the potential to provide spawning habitat for fish. High densities of black sea bream (Acanthopagrus schlegelii) eggs, sampled around oyster rafts in Hiroshima Bay, Japan, suggests that these farms provided habitat for spawning populations (Kawai et al. 2021). Our direct observations of courtship and spawning activity on oyster aquaculture cages in Long Island Sound provides empirical evidence that the structured habitat created by shellfish farms supports the occurrence of natural reproductive activity in temperate reef fish. Newly settled young-of-the-year fish observed in video collected on cages suggests that oyster aquaculture gear may provide habitat for the offspring of reef-oriented species that spawn in the vicinity of the farms (Mercaldo-Allen et al. 2023).
Implications for EFH Designations
Behavioural observations of scup indicate habitat provisioning by oyster cages, making a case for consideration of oyster aquaculture gear as fish habitat in the aquaculture permitting review process. In a related study of black sea bass, assessment of behaviour across life stages provided evidence that cages confer ecological benefits similar to natural habitat, affording equal or greater quality habitat (Mercaldo-Allen et al. 2025). Unlike black sea bass, artificial structures (such as aquaculture cages) are not included in the EFH description for scup, despite a documented affinity for natural and artificial structured habitat across multiple life history stages for both species. Where artificial structures are included in the EFH description, such as with black sea bass, the addition of aquaculture structures for such species would result in potential positive (in unstructured habitats) or neutral effects (in structured habitats) in EFH assessments and consultations for such species, as opposed to strictly considering placement of gear as an adverse effect on all designated and mapped EFH. Prior to inclusion of ecological services within the permitting decision-making process, further research is required to assess whether aquaculture practices (e.g., temporary or seasonal removal of cages from the water for maintenance and harvest) can affect the value and scale of services provided by aquaculture gear (Mercaldo-Allen et al. 2025). In addition, it may require the appropriate fisheries management councils to amend the EFH definitions for scup to include artificial structures as EFH for associated life history stages. Results of this work provide compelling evidence of beneficial associations between cages and structure-oriented scup and similar provision of services as habitats currently defined as EFH for the species.
Conclusions
Direct visual observations of wild fish behaviour associated with oyster aquaculture cages and boulders enabled documentation of the provision of habitat services to temperate reef fish. Cages on oyster farms provide similar functions for fish as do boulders on rock reefs. These ecological benefits include creation of surface area for colonization by species that provide forage, shelter from predation and respite from current flow, habitat where courtship and spawning activity can occur and provide sufficient resources to support communities of schooling or grouping fish. Territorial defence of shelter associated with cage structures, particularly by cunner and tautog, may be interpreted as an indicator of habitat quality. Behavioural interactions of young-of-the-year fish with cages, including feeding, predator escape, schooling/grouping and sheltering may indicate a nursery role function of aquaculture gear. These findings suggest that oyster farms may serve as EFH for managed species like scup for which manmade structures are not already included in habitat designations.
Author Contributions
All authors agree to be listed and have made substantial contributions to this work. Gillian Phillips: data curation, investigation, methodology, validation, writing – original draft preparation (lead). Renee Mercaldo-Allen: conceptualization, funding acquisition, methodology, project administration (lead), resources, supervision (lead), writing – original draft preparation (lead), writing – review and editing. Peter Auster: conceptualization. Paul Clark: data curation, investigation, methodology, validation. Mark Dixon: investigation, methodology, resources. Dylan Redman: investigation, methodology, resources. Barry Smith: investigation, methodology, resources. Alison Verkade: conceptualization. Christopher Schillaci: funding acquisition, writing – original draft preparation. Julie Rose: conceptualization, formal analysis, funding acquisition, project administration (lead), supervision (lead), visualization. All authors reviewed a first draft and approved submission.
Acknowledgements
We thank NOAA's Northeast Fisheries Science Center and Office of Aquaculture for funding. Field operations were conducted aboard the Milford Laboratory's 15-m NOAA R/V Victor Loosanoff. We thank David Carey, Kristin DeRosia-Banick and Shannon Kelly of the State of Connecticut, Bureau of Aquaculture, Jimmy Bloom of Norm Bloom and Sons and James Markow of Noank Aquaculture Cooperative for the loan of seed oysters and deployment assistance, shellfish growers Gary Salce of G & B Shellfish and Charles Viens of Charles Island Oyster Farms for access to leased shellfish beds, NOAA divers Calandrea DeCastro, Keith Golden, and Jerry Prezioso for additional dive support, and Peter Hudson, Deaven Maull and Max Mauro for support of field operations, Ryan Morse for assistance with the Zenodo data repository, and Grace Cajski, Samuel Pletcher and Ryan Rubino for assistance in behavior scoring. Use of tradenames does not imply endorsement.
Funding
We thank NOAA's Northeast Fisheries Science Center and Office of Aquaculture for funding.
Ethics Statement
This study was carried out in strict accordance with the Guidelines for the Use of Fishes in Research ().
Conflicts of Interest
The authors declare no conflicts of interest.
Data Availability Statement
Data is available from Zenodo repository, Phillips et al. 2025: .
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