Large Shoals Research Articles

Speckle statistics as a tool to distinguish collective behaviors of Zebrafish shoals

Zebrafish have become an important model animal for studying the emergence of collective behavior in nature. Here, we show how to properly analyze the polarization statistics to distinguish shoal regimes. In analogy with the statistical properties of optical speckles, we show that exponential and Rayleigh distributions emerge in shoals with many fish with uncorrelated velocity directions. In the opposite limit of just two fish, the polarization distribution peaks at high polarity, with the average value being a decreasing function of the shoal’s size, even in the absence of correlations. We also perform a set of experiments unveiling two shoaling regimes. Large shoals behave as small domains with strong intra-domain and weak inter-domain correlations. A strongly correlated regime develops for small shoals. The reported polarization statistical features shall guide future automated neuroscience, pharmacological, toxicological, and embryogenesis-motivated experiments aiming to explore the collective behavior of fish shoals.

Multivariate Data Assimilation at a Partially Mixed Estuary

Abstract In 2013 and 2014, multiple field excursions of varying scope were concentrated on the Columbia River, a highly energetic, partially mixed estuary. These experiments included surface drifter and synthetic aperture radar (SAR) measurements during the ONR RIVET-II experiment, and a novel animal tracking effort that samples oceanographic data by employing cormorants tagged with biologging devices. In the present work, several different data types from these experiments were combined in order to test an iterative, ensemble-based inversion methodology at the mouth of the Columbia River (MCR). Results show that, despite inherent limitations of observation and model accuracy, it is possible to detect dynamically relevant bathymetric features such as large shoals and channels while originating from a linear, featureless prior bathymetry in a partially mixed estuary by inverting surface current and gravity wave observations with a 3D hydrostatic ocean model. Bathymetry estimation skill depends on two factors: location (i.e., differing estimation quality inside versus outside the MCR) and observation type (e.g., surface currents versus significant wave height). Despite not being inverted directly, temperature and salinity outputs in the hydrodynamic model improved agreement with observations after bathymetry inversion.

Sex differences in the expression of aggressive behavior and influences on social choice in zebrafish (Danio rerio)

Aggregation behavior (grouping) provides many animals with the ecological benefits of protection from predators, increased access to food, and greater mating opportunities, despite possible costs. Many factors likely contribute to the social choices animals make, and we sought to determine if the individual expression of aggression would correlate with shoalmate choices. We quantified aggressive or submissive tendencies of individual male and female zebrafish and shoalmate preferences within dichotomous choice assays. We hypothesized that fish, regardless of their individual aggressive behaviors, would benefit most and therefore choose to spend time near larger shoals and shoals of the opposite sex. Both sexes spent significantly more time near shoals rather than be alone. Males spent significantly more time with the largest shoal, and females exhibited a similar trend. Both sexes spent more time with shoals of females, rather than males. Males were generally more consistent in their aggressive behaviors across multiple assays and females exhibited more individual-variability. More aggressive male zebrafish were more likely to choose shoals of males over females and were more likely to swim alone, but female zebrafish did not exhibit a correlation between aggressive behaviors and social choices. Our results demonstrate clear sex differences in the expression of individual behavior and influences on shoaling.

A preliminary study on the mechanism of the Liangzhu culture’s migration across the Yangtze river

The Liangzhu culture (5300–4200 cal BP) was the most famous Neolithic culture of who settled near the lower reaches of the Yangtze River in China. The core and initial distribution area of Liangzhu originated around Taihu Lake, located on the south bank of the Yangtze River delta. Recently, archaeological studies believe that the Jianghuai area and Huanghuai area north of the Yangtze River are also important distribution areas of Liangzhu culture. The route for Liangzhu culture migrating across the Yangtze River is inferred as follows: One would have crossed the Yangtze River from Nanjing-Zhenjiang belt and continued to migrate northward; while the other would have crossed the River near the estuary before moving north along the ancient coastline to the Jianghuai during the late period of Liangzhu, or crossed the Yangtze River from the east of the present Beijing-Hangzhou Grand Canal to Jianghuai and Huanghuai. According to the formation of the Yangtze River delta during the Holocene and the evolution of the estuarine sand bar, it is believed that there were large shoals in the Yangtze River channel in the middle-Holocene. The Liangzhu ancestors around 5000 cal BP had the ability of making canoes over 8 m. Based on the archaeological research of the Neolithic period, the evolution of the Yangtze River channel in the Holocene, the history of ancient Chinese shipbuilding, and the modern examples of crossing the Yangtze River with boat, it can be concluded that the present Changzhou–Jiangyin–Zhangjiagang line should be the main and reasonable route for the Liangzhu culture migrating across the Yangtze River.

Uncharted coral reefs from the Inhambane Province, Mozambique

Mozambique has the third-largest coastline in the western Indian Ocean with a large and diverse reef area. Those reefs provide food and income for a large proportion of coastal communities but remain scarcely studied. As a baseline assessment, we sampled four uncharted shallow reefs (Baixo Africa, Baixo Zambia, Pomene and Baixo Silvia) within a 56 km stretch line from 5 to 32 meters deep. The fish abundance and assemblage composition were assessed by underwater visual census (25x2m, n = 112). The benthic community was characterized by digital images (0.5x0.5m, n = 164) and the invertebrate megafauna by direct counting (25x2m, n = 52). A total of 209 reef fish species was observed, with similar mean richness (22.4 ± 7.2 species.100 m−2) at all reefs. Invertivorous (n = 79) and planktivorous fish species (n = 36) represented 55% of species total. Trophic composition was similar among sites and with large contribution of planktivores. The presence of large shoals (>100 ind.100 m−2, e.g., surgeonfish, fusiliers, snappers) and large-bodied fishes (>90 cm) (e.g., groupers) was a clear indication of local productivity and health conditions of reefs. Seventy-nine taxa were observed on the benthic cover with the largest richness found at Baixo Silvia (28.5 ± 7.4, species.100m −2, mean ± SD). Turf algae dominated the benthos (31.5% ± 16.4) at all sampled reefs. Coral cover was variable between sites (9.3 – 25.8%), dominated by branching and plate corals, and <1% of colonies showed signs of diseases/bleaching. The baseline assessment indicated large biomass of fishes and negligible evidence of large impacts on coral formations which indicated a good conservation status of those areas. Based on these findings, a marine protected area is proposed for establishment in collaboration with environmental authorities and the local community.

Dynamic visual noise promotes social attraction, but does not affect group size preference, in a shoaling fish

Gathering information from the environment allows animals to make informed behavioural decisions, with individuals sampling information either privately or via social cues from group members. Environmental noise, however, may disrupt the ability of animals to gather information in their environment. Therefore, social interactions could be disrupted by environmental noise, or adapted to mitigate the costs associated with compromised perception in noisy environments. Here we tested whether the presence of water caustics, a natural form of visual noise prevalent within shallow aquatic habitats, affects the social decisions of shoaling fish. In a classic experimental paradigm, individual three-spined sticklebacks, Gasterosteus aculeatus, were given a choice between two conspecific shoals differing in size while immersed in different levels of visual noise. Fish showed greater social attraction towards conspecifics as noise levels increased, measured by a reduction in the distance they maintained from others. While sticklebacks only showed clear preferences for groups that had at least four more members than another group, there was no evidence that visual noise level influenced fish's preference to associate with these larger shoals, nor did it change the preferences for other shoal sizes. Our findings suggest that, at the group sizes tested, visual noise increases individual stickleback's social attraction towards conspecifics, but does not affect their decision to associate with the larger of two groups. Such changes to social behaviour may reflect the consequences of having a reduced ability to sample private, but not social, information in noisy environments.

Transcriptomic Diversity in the Livers of South African Sardines Participating in the Annual Sardine Run.

During austral winter, the southern and eastern coastlines of South Africa witness one of the largest animal migrations on the planet, the KwaZulu-Natal sardine run. Hundreds of millions of temperate sardines, Sardinops sagax, form large shoals that migrate north-east towards the subtropical Indian Ocean. Recent studies have highlighted the role that genetic and environmental factors play in sardine run formation. In the present study, we used massively parallel sequencing to assemble and annotate the first reference transcriptome from the liver cells of South African sardines, and to investigate the functional content and transcriptomic diversity. A total of 1,310,530 transcripts with an N50 of 1578 bp were assembled de novo. Several genes and core biochemical pathways that modulate energy production, energy storage, digestion, secretory processes, immune responses, signaling, regulatory processes, and detoxification were identified. The functional content of the liver transcriptome from six individuals that participated in the 2019 sardine run demonstrated heterogeneous levels of variation. Data presented in the current study provide new insights into the complex function of the liver transcriptome in South African sardines.

Spatial and Temporal Patterns in Macroherbivore Grazing in a Multi-Species Tropical Seagrass Meadow of the Great Barrier Reef

Macroherbivory is an important process in seagrass meadows worldwide; however, the impact of macroherbivores on seagrasses in the Great Barrier Reef (GBR) has received little attention. We used exclusion cages and seagrass tethering assays to understand how the intensity of macroherbivory varies over space and time in the seagrass meadows around Green Island (Queensland), and what impact this has on overall meadow structure. Rates of macroherbivory were comparatively low, between 0.25–44% of daily seagrass productivity; however, rates were highly variable over a one-year period, and among sites. Loss of seagrass material to macroherbivory was predominantly due to fish; however, urchin herbivory was also taking place. Macroherbivory rates were of insufficient intensity to impact overall meadow structure. No macroherbivory events were identified on video cameras that filmed in the day, indicating that feeding may be occurring infrequently in large shoals, or at night. While relatively low compared to some meadows, seagrass macroherbivory was still an important process at this site. We suggest that in this highly protected area of the GBR, where the ecosystem and food webs remain largely intact, macroherbivory was maintained at a low level and was unlikely to cause the large-scale meadow structuring influence that can be seen in more modified seagrass systems.

Ontogenetic change in the diet of Semaprochilodus insignis (Characiformes: Prochilodontidae) during migration between two limnologically distinct environments in the Amazon Basin

Abstract Semaprochilodus insignis is an Amazonian migratory fish species, moving in large shoals between white- and black-water rivers. It has long been classified as a detritivorous fish. However, it is possible that the trophic plasticity of S. insignis could be higher than previously assumed. The objective of this study was to investigate the relative contributions of autotrophic energy sources to the diet of S. insignis in the Negro and Solimões rivers and to determine if the species undergoes an ontogenetic change in the diet. We found variations between the δ13C and δ15N values of S. insignis between the rivers. In the Negro River, periphyton (84%) was the principal energy source for adults, while juveniles in the Solimões River foraged predominantly on terrestrial plants/C3 macrophytes (50%) and phytoplankton (42%). These variations in isotopic signatures are likely associated with migratory movements of S. insignis at different life stages and hydrological periods. Instead of the previously assumed dietary classification, we suggest that S. insignis varies its diet ontogenetically, with adults acting as illiophagous in black-water while the young are detritivorous in white-water rivers. The results show that this species creates complex links between food chains, thus emphasizing the importance of conserving flooded areas.

Molecular identification of Brachygenys and Haemulon species (Perciformes: Haemulidae) from the Brazilian coast

Abstract The fishes of the Haemulidae family are currently allocated to 19 genera with a worldwide distribution in the tropical and subtropical waters of the world’s oceans. Brachygenys and Haemulon are important genera of reef fish in Brazil, as they occur in large shoals, which are both ecologically and commercially valuable. This study identified the Brazilian species of the genera Brachygenys and Haemulon using DNA barcodes. While we found only a single lineage in Brachygenys chrysargyrea, Haemulon melanurum, H. parra, and H. squamipinna, more than one molecular operational taxonomic unit (MOTU) was identified in H. atlanticus, H. aurolineatum, and H. plumieri, indicating the possible existence of discrete populations or cryptic species.

Unknown facts of the biology of Polybius henslowii – Effect of gender on claw morphology and diet

Polybius henslowii is known for both its unusual swimming behaviour and for forming large shoals during its pelagic phase. Although being a common species, many aspects of its biology remain unknown. In this work, P. henslowii's sexual dimorphism was examined in both dietary composition and morphologies of carapace and claw. Males were generally larger than females both on carapace and claw sizes. Claw morphology differed between males and females. Diet was similar between genders. Hence, we suggest that the sexual dimorphism in claw morphology results from gender-specific behaviours instead of diet. The most common dietary items were sand (51.2%), squid (34.1%) and crustaceans (26.8%). The same items were also the most important in volume, but with squids being more important than sand. The diet composition of P. henslowii was consistent with previous studies but also presenting sediment as an important item.

Complex Flow Patterns in the Scheldt Estuary: Field Measurements and Validation of a Hydrodynamic Model

AbstractThe morphology of the Scheldt estuary creates challenges for its different functions. Large shoals and adjacent channels with sharp bends result in challenging conditions for navigation. In...

Decadal morphological evolution of the mouth zone of the Yangtze Estuary in response to human interventions

AbstractThe morphology of the Yangtze Estuary has changed substantially at decadal time scales in response to natural processes, local human interference and reduced sediment supply. Due to its high sediment load, the morphodynamic response time of the estuary is short, providing a valuable semi‐natural system to evaluate large‐scale estuarine morphodynamic responses to interference. Previous studies primarily addressed local morphologic changes within the estuary, but since an overall sediment balance is missing, it remains unclear whether the estuary as a whole has shifted from sedimentation to erosion in response to reduced riverine sediment supply (e.g. resulting from construction of the Three Gorges Dam). In this paper we examine the morphological changes of two large shoals in the mouth zone (i.e. the Hengsha flat and the Jiuduan shoal) using bathymetric data collected between 1953 and 2016 and a series of satellite images. We observe that the two shoals accreted at different rates before 2010 but reverted to erosion thereafter. Human activities such as dredging and dumping contribute to erosion, masking the impacts of sediment source reduction. The effects of local human intervention (such as the construction of a navigation channel) are instantaneous and are likely to have already resulted in new dynamic equilibrium conditions. The morphodynamic response time of the mouth zone to riverine sediment decrease is further suggested to be &gt;30 years (starting from the mid‐1980s). Accounting for the different adaptation time scales of various human activities is essential when interpreting morphodynamic changes in large‐scale estuaries and deltas. © 2019 John Wiley &amp; Sons, Ltd.

Bivalves boost biodiversity

Bivalves boost biodiversity

Juvenile convict cichlids shoaling decisions in relation to shoal size and age

AbstractMany animals form groups and socialize in response to evolutionary pressures such as predation, food availability, and mate acquisition. Evidence of social choice based on various phenotypic characters (Group Phenotypic Composition [GPC]) has been observed in several animal species. In addition to the physical characteristics of the social group, it is also interesting to consider how decisions of who to socialize with might be expected to change for an individual over time. Younger individuals with limited life experience may discriminate differently between social groups than older conspecifics who have had the opportunity to learn and who may be faced with different ecological or environmental pressures. Here, we used a traditional two‐choice design to explore the shoaling behavior of juvenile convict cichlids and determine whether the number of fish and/or the size/life stage of the individuals within a shoal influenced social choices. We found that juvenile convict cichlids spent more time shoaling with similarly sized juvenile individuals and also preferred to shoal with larger shoals, but not when shoals were comprised of adult fish. The size of the individuals in a shoal was a more influential factor than the size of the shoal itself. Size of individual juveniles was correlated with tendency to visit shoals, but was not correlated with overall time spent shoaling, regardless of shoal composition. As juveniles, convict cichlids can make discriminatory choices that are influenced by specific aspects of shoal composition.

Comparison of Fish Catch Between an Electric Seine and Backpack Electrofishers in Shoal Habitat of the Chipola River, Florida

Abstract We conducted a gear comparison study to determine the best fish-sampling methodology in shoal habitat to assist with the development of a long-term fish community monitoring program on the Chipola River, Florida. Shoal habitat occurs throughout the southeastern United States, is shallow (&amp;lt; 1 m), and contains rock substrate and outcroppings, which limits boat electrofishing to the habitat margins. We compared the relative effectiveness of an electric seine and backpack electrofishers. Twenty-five randomly selected sites (4 m wide by 5 m long) per gear type were block netted and sampled across four shoals from August 28 to October 3, 2014. Habitat variables were recorded for each site. Using backpack electrofishers, 1,155 fish representing 21 species were collected. Using the electric seine, 1,036 fish representing 22 species were collected. We found no significant differences between gear types in mean total catch per site, mean species richness per site, Shannon's diversity, Simpson's dominance, Pielou's evenness, or the rate of species accumulation. Therefore, we consider the two gear types to be equally effective for block-netted sites in large river shoals. Differences in available equipment, funding, and personnel were compared and may ultimately determine which gear type is used in future studies.

Quantity discrimination in angelfish (Pterophyllum scalare) is maintained after a 30-s retention interval in the large but not in the small number range.

The ability to discriminate between sets that differ in the number of elements can be useful in different contexts and may have survival and fitness consequences. As such, numerical/quantity discrimination has been demonstrated in a diversity of animal species. In the laboratory, this ability has been analyzed, for example, using binary choice tests. Furthermore, when the different number of items first presented to the subjects are subsequently obscured, i.e., are not visible at the moment of making a choice, the task requires memory for the size of the sets. In previous work, angelfish (Pterophyllum scalare) have been found to be able to discriminate shoals differing in the number of shoal members both in the small (less than 4) and the large (4 or more) number range, and they were able to perform well even when a short memory retention interval (2-15s) was imposed. In the current study, we increased the retention interval to 30s during which the shoals to choose between were obscured, and investigated whether angelfish could show preference for the larger shoal they saw before this interval. Subjects were faced with a discrimination between numerically small shoals (≤4 fish) and also between numerically large (≥4 fish) shoals of conspecifics. We found angelfish not to be able to remember the location of larger versus smaller shoals in the small number range, but to exhibit significant memory for the larger shoal in the large number range as long as the ratio between these shoals was at least 2:1. These results, together with prior findings, suggest the existence of two separate quantity estimation systems, the object file system for small number of items that does not work with the longer retention interval and the analogue magnitude system for larger number of items that does.

Non-consumptive effects of native and invasive predators on juvenile Caribbean parrotfish

Non-consumptive effects of predators can have important impacts on aquatic food webs, but there are few data on how predators change the behaviour of Caribbean reef fishes. Such changes may include behavioural responses to the invasive predatory lionfish (Pterois volitans/P. miles). This study used an aquarium experiment to examine the behaviour of herbivorous parrotfish (Scarus iseri) in the absence of other fish (control), with a non-piscivore present, and with a predatory threat from a native grouper or lionfish. Treatments were repeated with and without additional parrotfish shelters to examine the potential effects of degraded reefs (loss of refuges). Using video, parrotfish behaviours (sheltering, swimming in open areas, foraging, aggressive conspecific interactions, bite rates, and shoaling behaviour) were recorded for groups of four parrotfish. Compared to the control, the average number of parrotfish hiding was reduced by 65 % and foraging shoals were 10 % larger when threatened by grouper, likely as a specific response to an ambush predator. When exposed to lionfish, parrotfish reduced their bite rates by 50 %, possibly to be more vigilant of this predator’s unique stalking behaviour. The absence of additional shelter had limited effects although parrotfish formed 10 % larger shoals when swimming in open water, potentially as a defensive behaviour because of a perceived lack of refuges. The reduction in parrotfish bite rates caused by lionfish may have important demographic consequences. Furthermore, parrotfishes are important grazers of macroalgae, and these behavioural changes may exacerbate the direct effects of lionfish predation and potentially affect reef benthic dynamics.

The effect of temporally variable environmental stimuli and group size on emergence behavior

How animals trade-off food availability and predation threats is a strong determinant of animal activity and behavior; however, the majority of work on this topic has been on individual animals, despite the modulating effect the presence of conspecifics can have on both foraging and predation risk. Although these environmental factors (food and predation threat) vary spatially within habitats, they also vary temporally, and in marine habitats, this can be determined by not only the diel cycle but also the tidal cycle. Humbug damselfish, Dascyllus aruanus, live in small groups of unrelated individuals within and around branching coral heads, which they collectively withdraw into to escape a predation threat. In this study, we measured the proportion of individuals in the colony that were outside the coral head before and after they were scared by a fright stimulus and compared the responses at high tide (HT) and low tide (LT). We found that a greater proportion of the shoal emerged after the fright stimulus at HT and in larger groups than at LT or in smaller groups. We also quantified the pattern of emergence over time and discovered the rate of emergence was faster in larger shoals as time progressed. We show that shoals of fish change their behavioral response to a predation threat in accordance with the tide, exemplifying how temporally variable environmental factors can shape group movement decisions.

Discrimination of large quantities: Weber's law and short-term memory in angelfish, Pterophyllum scalare

The ability to discriminate between different quantities has important ecological relevance for animals when engaging in behaviours such as forming groups, foraging or trying to avoid predators. Quantity discrimination has been shown in a diversity of human and nonhuman animal species. In angelfish this discrimination ability has been investigated using dichotomous choice tests when the numerically different stimulus groups (shoals) of conspecifics were fully visible to the test fish. Here, using a new procedure we investigated whether test fish were able to discriminate between the contrasting shoals using their memory. After a period of full visual access to the contrasted shoals on the two sides of their test tank, the test fish was required to make a choice while being able to see only a single member of the stimulus shoals on each side. With this cognitively more demanding procedure we tested discrimination between numerically large shoals (≥ four fish per stimulus shoal). As in our previous studies, we found that angelfish consistently chose the larger of the two shoals when the shoals differed by a 2:1 or higher ratio, but not those that differed by a 3:2 or 4:3 ratio. The results followed Weber's law in that performance became poorer as the ratio between the two stimulus shoals approached one. In addition, when we kept the absolute difference between the contrasted shoals constant, discrimination was less accurate as the shoal sizes increased. This pattern of results lends support for the analogue magnitude representational system in the angelfish, a nonverbal approximation system believed to be employed by a diversity of human and nonhuman animal species. Furthermore, our results also demonstrate that angelfish remember the different shoals presented to them, i.e. they make their choice based upon mental representation of the different quantities.