Aggression and welfare in a common aquarium fish, the Midas cichlid

Many species of fishes are aggressive when placed in small aquaria. Aggression can negatively affect the welfare of those individuals toward whom it is directed. Animals may behave aggressively in order to defend resources such as food, shelter, mates, and offspring. The decision to defend depends on the distribution of resources and on ecological factors such as number of competitors, amount of available space, and amount of habitat complexity. This study tested the effects of these factors on aggression in a common aquarium fish, the Midas cichlid (Amphilophus citrinellus). The study found that time spent behaving aggressively was not associated with small-scale differences in group size or available space. Aggression was significantly lower in a large aquarium with a complex habitat. Aquaria of sizes typically used in the companion animal (pet) hobby did not provide optimal welfare for cichlids housed with aggressive conspecifics. The public should be aware that this and similar species require larger aquaria with complex habitat, which elicit more natural behavior.     

Aggression in closely related Malawi cichlids varies inversely with habitat complexity

Within the past 2 million years, the cichlids of Lake Malawi have diversified into well over 500 species resulting in one of the worlds largest lacustrine fish radiations. As a result, many of the habitats within the lake support a high diversity of species. In these highly species rich communities, male cichlids must acquire and defend a territory to successfully reproduce. Within the rock-dwelling cichlids of Lake Malawi (mbuna), this has resulted in the formation of polyspecific leks on the heterogeneous rocky benthos. Aggression is fairly common in these leks and has been tied not only to individual reproduction but to the larger phenomenon of community assembly and the maintenance of biological diversity. In this study, I examined the patterns of aggressive acts of four species within the mbuna genus Maylandia at two locations in the southern Lake Malawi. The number of aggressive acts of two sympatric species was examined at each location. At each site, one species defends territories over bedrock and the other over cobble. The number of aggressive acts across the four species was compared. The influence of habitat type on male aggression was examined and the targets of male aggression were identified to evaluate several hypotheses concerning the evolution of male aggression. The results show that aggression quantitatively varied among species, was largely directed towards heterospecifics, and was strongly influenced by habitat type. The aggressive behavior of one sympatric species pair, Maylandia benetos and Maylandia zebra, was observed under controlled laboratory conditions. Laboratory results support field observations: the bedrock associated species performed more aggressive acts and aggressive acts were directed equally at con- and heterospecifics. The results of this study suggest that habitat complexity plays a larger role in shaping aggressive behavior than other suggested factors such as competition for resources.     

Loopholes in the regulation of invasive species: genetic identifications identify mislabeling of prohibited aquarium plants

Numerous invasive aquatic species introductions can be traced to the aquarium trade. Many potentially harmful aquarium species may be difficult to identify based on morphology alone. As such, some prohibited or invasive species may be available for purchase if they are mislabeled as species without restrictions. Here we compare molecular identifications to internet vendors’ identifications for accessions of a popular genus of aquarium plants that are difficult to distinguish morphologically (Myriophyllum; watermilfoils). Specifically, we identified the extensive mislabeling of M. heterophyllum—an invasive species in the northeastern and western US. Furthermore, genotypes of M. heterophyllum found in our aquarium survey have also been found in invasive populations, suggesting their potential introduction through escape from aquaria, water gardens, or nurseries. Two additional taxa were sold under incorrect names. Finally, our survey revealed that Myriophyllum taxa present in the aquarium trade generally have poorly known distributions and ecologies, and therefore their invasive potential is unknown. Our study confirms that molecular identification methods can provide a valuable tool to survey commercial pathways for potentially harmful species that are otherwise difficult to identify.     

Fish communities on staghorn coral: effects of habitat characteristics and resident farmerfishes

Branching corals, like many in the genus Acropora, provide structurally complex habitats for reef fishes and other organisms. Fluctuations in the abundance, distribution and characteristics of thicket-forming staghorn Acroporids may contribute to changes in the abundance and species composition of reef fishes due to changes in the availability of shelter habitat and food. Farming damselfishes of the genus Stegastes can occur in high abundances in staghorn corals and actively defend food and nest space against organisms that threaten these resources. Here we assess the value of staghorn as habitat for fishes in the central South Pacific, and how the presence of territorial farming damselfishes may influence the assemblage of fishes that associate with staghorn corals. Surveys of 185 Acropora pulchra patches located in the lagoons surrounding the island of Moorea, French Polynesia revealed 85 species of fish from 25 families. Total fish abundance and species richness values ranged from no fish on a patch to a high of 275 individuals and 26 species. Patch area was the most important characteristic in explaining variation in attributes of the fish assemblage, with other characteristics explaining little of the species composition or trophic structure. Behavioral observations revealed that farming damselfishes were most aggressive toward corallivores, herbivores, and egg predators, while they ignored most carnivores and omnivores. Despite this pattern, we observed positive covariance between Stegastes and the group of fishes that elicited the strongest aggressive response when the effect of patch area was removed, suggesting these fishes remain drawn to the resources produced or enhanced by Stegastes on A. pulchra.     

The freshwater aquarium trade as a vector for incidental invertebrate fauna

The aquarium trade has a long history of transporting and introducing fish, plants and snails into regions where they are not native. However, other than snails, research on species carried “incidentally” rather than deliberately by this industry is lacking. I sampled invertebrates in the plankton, and from water among bottom stones, of 55 aquaria from 43 New Zealand households. I recorded 55 incidental invertebrate taxa, including copepods, ostracods, cladocerans, molluscs, mites, flatworms and nematodes. Six were known established non-indigenous species, and eight others were not previously recorded from New Zealand. Of the latter, two harpacticoid copepod species, Nitokra pietschmanni and Elaphoidella sewelli, are not native to or known from New Zealand, demonstrating the aquarium trade continues to pose an invasion risk for incidental fauna. The remaining six species were littoral/benthic rotifers with subtropical/tropical affinities; these may or may not be native, as research on this group is limited. A variety of behaviours associated with the set-up and keeping of home aquaria were recorded (e.g., fish and plants in any home were sourced from stores, wild caught, or both, and cleaning methods varied), which made prediction of “high risk” behaviours difficult. However, non-indigenous species had a greater probability of being recorded in aquaria containing aquatic plants and in those that were heated. Methods for disposal of aquarium wastes ranged from depositing washings on the lawn or garden (a low risk for invasion) to disposing of water into outdoor ponds or storm-water drains (a higher risk). It is recommended that aquarium owners be encouraged to pour aquarium wastes onto gardens or lawns—already a common method of disposal—as invasion risk will be minimised using this method.     

A risk assessment of aquarium trade introductions of seaweed in European waters

Aquaculture and maritime traffic have been identified as the main vectors for introductions of alien marine species. Except for one notorious case of Caulerpa taxifolia, the role of aquarium trade towards the introduction of alien seaweeds has been largely unassessed. Here, we address the risk of accidental release of seaweed species from the aquarium trade market in European waters. We assessed the importance and diversity of seaweed species in the European online aquarium retail circuit. Our web survey revealed more than 30 genera available for online sale into Europe, including known introduced and invasive species. A second aspect of the study consisted in sampling algal diversity found in aquaria. While allowing direct and accurate identification of the specimens, this approach was targeting not only ornamental species, but also seaweeds that may be accidentally present in the aquarium circuit. By DNA-barcoding we identified no less than 134 taxa, 7 of which are flagged as introduced in Europe and 5 reported as invasive. Climate envelope models show that at least 23 aquarium species have the potential to thrive in European waters. As expected by the tropical conditions in most aquaria, southern Atlantic regions of Europe and the Mediterranean are the most vulnerable towards new introductions. Further predictions show that this risk will increase and shift northwards as global warming proceeds. Overall our data indicate that aquarium trade poses a potential risk of new seaweed introductions, and calls for a cautious approach.     

Behavior and season affect crayfish detection and density inference using environmental DNA

Although the presence/absence of aquatic invertebrates using environmental DNA (eDNA) has been established for several species, inferring population densities has remained problematic. The invasive American signal crayfish, Pacifastacus leniusculus (Dana), is the leading cause of decline in the UK's only native crayfish species, Austropotamobius pallipes (Lereboullet). Methods to detect species at low abundances offer the opportunity for the early detection, and potential eradication, of P. leniusculus before population densities reach threatening levels in areas occupied by A. pallipes. Using a factorial experimental design with aquaria, we investigated the impacts of biomass, sex ratio, and fighting behavior on the amount of eDNA released by P. leniusculus, with the aim to infer density per aquarium depending on treatments. The amount of target eDNA in water samples from each aquarium was measured using the quantitative Polymerase Chain Reaction. We show that the presence of eggs significantly increases the concentration of crayfish eDNA per unit of mass, and that there is a significant relationship between eDNA concentration and biomass when females are egg‐bearing. However, the relationship between crayfish biomass and eDNA concentration is lost in aquaria without ovigerous females. Female‐specific tanks had significantly higher eDNA concentrations than male‐specific tanks, and the prevention of fighting did not impact the amount of eDNA in the water. These results indicate that detection and estimate of crayfish abundance using eDNA may be more effective while females are ovigerous. This information should guide further research for an accurate estimation of crayfish biomass in the field depending on the season. Our results indicate that detection and quantification of egg‐laying aquatic invertebrate species using eDNA could be most successful during periods when eggs are developing in the water. We recommend that practitioners consider the reproductive cycle of target species when attempting to study or detect aquatic species using eDNA in the field.     

Factors Affecting Interspecific Aggression in a Year‐Round Territorial Species,the Jewel Damselfish

Vertebrates live in complex species networks in which interspecific interactions are common. In some contexts, the aggressive behaviours shown in these interspecific interactions are very similar to those shown in intraspecific interactions. It is still an open question whether intra‐ and interspecific aggression share common causality. We studied a year‐round territorial species the jewel damselfish, (Plectroglyphidodon lacrymatus), which cultivate algae they feed on. Territory holders aggressively defend these algae that are an attractive resource for many other species. In this study, we recorded territorial aggression in free‐living individuals and recorded aggressive responses to a standardized territorial intrusion test in captive individuals. Field observations indicated that territorial aggression was selectively targeted towards food competitors. Independent of the size of the species, aggression was more frequent towards common species around their territories. This relationship was confirmed experimentally by confronting the jewel damselfish with novel objects to which the subjects were exposed either frequently or rarely. We suggest that jewel damselfish have to learn which species are competitors and therefore should be chased. In a standardized intrusion test with captive individuals, no significant differences were found in territorial responses towards intra‐ or interspecific intruders. Neither territorial aggression nor the intrusion showed any relationship with plasma androgen levels. Together, these data suggest that experience might be more important in non‐seasonal territorial aggression than circulating hormonal factors.     

Historical distribution and current drivers of guppy occurrence in Brazil

Humans introduce non-native species by means such as the deliberate release of fish into fresh waters and through commercial trade. The guppy Poecilia reticulata Peters, 1859, is commonly kept in aquaria and controls disease vectors, and now it occurs in many areas outside its natural distribution. Its initial habitat in Brazil was identified, and a study was performed to determine whether the density of guppies can be explained by the density of human population, per-capita gross domestic product, level of human impact on the areas where guppies have been found and fish-sampling effort. A total of 1402 guppy records were found; the southeastern region had the oldest records; and the southeastern, northeastern and central-western regions had the maximum records. Low tolerance to the colder climate may be the reason for the lack of guppy records in the southernmost states. It was also observed that the occurrence of this fish is positively, yet weakly, related to the density of human population, indicating that improved regulations regarding its use in controlling disease vectors, the aquarium trade and education of aquarium hobbyists could help prevent the spread of this species and its potential impacts in Brazil.     

How can Habitat Selection Affect the use of a Wetland Complex by Waterbirds?

The factors regulating the distribution of dabbling and diving waterbirds were studied, taking into account habitat selection by the species in a wetland complex of 26 ponds in south-eastern Spain. Such information can be used to management and conservation of wetland threatened bird species. Direct counts and feeding-microhabitat surveys of waterbirds were conducted. The feeding-niche width and the feeding-microhabitat use, as a function of the horizontal spatial gradient in the ponds, were related to the mean size of ponds used for each species. While the more generalist birds, which usually feed close to the shore, probably had available resources in small ponds, specialists that also frequently selected central zones of the pond had proportionally more limited feeding space and, therefore, less resource availability in small ponds. The differences in habitat selection of the different species appeared to encourage their hierarchical disappearance from the wetland complex at the same rate as the ponds diminished in size (‘nested’ pattern). The results support the conservation, restoration or creation of, at least, ponds of greater size in order to preserve extensive open-water zones in the wetlands and maintain the greatest number of specialist, threatened, and area-dependent species, such as the globally endangered White-headed Duck (Oxyura leucocephala).     

Environmental determinants of butterflyfish social systems

Synopsis Butterflyfishes (Chaetodontidae) display a variety of social systems, including monogamous pair-bonds, harems, and schooling with group spawning. The range of reproductive options available to butterflyfishes is shaped by their general life history characteristics, such as broadcast spawning with widely dispersed pelagic larvae, large body size and low adult mortality. The distribution and quality of food resources are major determinants of group size and mobility, thereby influencing the relative costs and benefits of available options, and determining specific social systems. Planktivorous and corallivorous butterflyfishes exemplify the relationship between food resources and social systems. Pelagic plankton is a patchy, but temporally and spatially unpredictable food resource which is efficiently exploited by fish in mobile schools. Neither sex is able to monopolize food resources necessary for the other sex, and plantivorous butterflyfishes appear constrained to spawn in groups. In contrast, corals are stable and predictable in space and time, favoring residence in one area and territorial defense of that space by coral-feeding butterflyfishes. Females defend food resources from other females, and males defend territories containing a female from other males. Males attempt to defend areas containing more than one female, but are unsuccessful. A monogamous social system results. This system favors the evolution of cooperative behavior between mates to increase female fecundity, as long as the male has an opportunity of sharing in that reproduction. Mate removal experiments conducted on two monogamous coral-feeding species,Chaetodon multicinctus andChaetodon quadrimaculatus reveal a division of labor between male and female pair-mates. Paired males assume most of the territorial defense activities, allowing their mates to feed more.     

Factors correlating with deterioration of giant kelp Macrocystis pyrifera (Laminariales, Heterokontophyta) in an aquarium setting

Survival of the giant kelp, Macrocystis pyrifera (Linnaeus) C. Agardh, in its natural habitat is governed by abiotic and biotic factors such as temperature, light, nutrients, current velocity, and predators. Factors affecting the survival of the alga in an aquarium setting, however, have not been investigated. The National Museum of Marine Biology and Aquarium (NMMBA), in subtropical Taiwan, is the only aquarium in the world that displays giant kelp that does not have naturally occurring specimens in nearby waters. Giant kelp displayed in aquaria often deteriorates within a 3-month period, yet the cause of this mortality is unknown. We investigated abiotic and biotic parameters affecting survival of giant kelp in aquaria over a 3-month period. The results indicated that temperature, salinity, pH, light, and nutrient concentrations did not affect giant kelp survival. However, the massive proliferation of epiphytic diatoms on kelp blades (from 7?×?10² cells cm^?2 initially to 3?×?10⁴ cells cm^?2 after 1 month) was identified as being the most likely candidate affecting survival of giant kelp in an aquarium setting. Potential factors that may stimulate epiphyte proliferation include lack of epiphytic algae control via predation, high nutrient concentrations, a weak current, and a generally stable environment.     

Behavioural and Genetic Data Suggest that Bechstein's Bats Predominantly Mate Outside the Breeding Habitat

Investigating mating systems of species with a cryptic lifestyle often requires a combination of behavioural and genetic data. We used such a combined approach to investigate the mating system of the communal breeding Bechstein's bat (Myotis bechsteinii). Although females of this species are philopatric, gene flow among colonies is high. Gene flow occurs if dispersing males mate within the colony to which they moved. Males could gain local matings by defending resources or females in their breeding habitat. Alternatively, mating may take place at swarming sites, apart from the breeding habitat of the females, where males and females of several colonies meet. Whether or not males defend resources or females in the breeding habitat is of importance for understanding the mating system. Detailed observations of individual foraging and roosting behaviour over 4 yr suggest that males do not defend resources or females to gain matings. Moreover, paternity assignment based on microsatellite data of four complete juvenile cohorts showed that local males fathered less than 25% of the juveniles born in the colony where they settled. Even more striking, none of the males that had immigrated into our study area reproduced with the local females.     

Networks and dominance hierarchies: does interspecific aggression explain flower partitioning among stingless bees?

1. The distribution of consumers among resources (trophic interaction network) may be shaped by asymmetric competition. Dominance hierarchy models predict that asymmetric interference competition leads to a domination of high quality resources by hierarchically superior species. 2. In order to determine the competitive dominance hierarchy and its effect on flower partitioning in a local stingless bee community in Borneo, interspecific aggressions were tested among eight species in arena experiments. 3. All species tested were strongly mutually aggressive in the arena, and the observed interactions were often lethal for one or both opponents. Aggression significantly increased with body size differences between fighting pairs and was asymmetric: larger aggressors were superior over smaller species. Additional aggression tests involved dummies with surface extracts, and results suggest that species‐ and colony‐specific surface profiles are important in triggering the aggressive behaviour. 4. Sixteen stingless bee species were observed foraging on 41 species of flowering plants. The resulting bee–flower interaction network showed a high degree of generalisation (network‐level specialisation H₂’ = 0.11), corresponding to a random, opportunistic distribution of bee species among available flower species. 5. Aggressions on flowers were rare and only occurred at a low level. The dominance hierarchy obtained in the arena experiments did not correlate significantly with plant quality, estimated as the number of flowers per plant or as total bee visitation rate. 6. Our findings suggest that asymmetries in interference competition do not necessarily translate into actual resource partitioning in the context of complex interacting communities.     

Competitive ability and the potential for lotteries among territorial reef fishes

Stegastes diencaeus and S. dorsopunicans are mutually territorial Caribbean damselfishes. S. diencaeus is larger, grows faster and lives longer than S. dorsopunicans. S. diencaeus is a habitat specialist that shares its primary habitat mainly with S. dorsopunicans. Field manipulations show that both S. diencaeus and S. dorsopunicans readily take over living space from smaller, but not larger, heterospecific neighbors. Natural changes in the use of living space by both species occur frequently and adult S. diencaeus often aggressively usurp the living areas of smaller S. dorsopunicans. Lunar and seasonal patterns of juvenile recruitment by S. diencaeus and S. dorsopunicans are similar. Large size bestows competitive superiority on S. diencaeus by giving its adults a superior ability to aggressively acquire living space, and by enabling its juveniles to quickly escape the period when they lack a size advantage. Hence they spend much of their lives as competitive dominants. There is no evidence that competitive advantages arising from large size are offset either by other adult attributes or by differences in temporal patterns of recruitment that affect priority of access to space. The lottery hypothesis for species coexistence relies on patterns of abundance being determined by patterns of recruitment to vacant space because different species have equal space-holding abilities. These data show that the existence of such a mechanism is doubtful.     

The aggressive behavior of Nile tilapia introduced into novel environments with variation in enrichment

Many studies show environmental enrichment is correlated with benefits to captive animals; however, one should not always assume this positive relationship given that enrichment increases the amount of resources that a territorial animal must defend and possibly affects its aggressive dynamics. In this study, we tested if environmental enrichment affects aggressive interactions in the aggressive fish Nile tilapia (Oreochromis niloticus). We compared fights staged between pairs of male tilapia of similar size (= matched in resource holding potential) in a novel arena that was either barren or enriched, to examine whether enrichment enhances territory value in line with theoretical predictions, with the potential for compromised welfare. We evaluated time elapsed until the first attack (latency), frequency of aggressive interactions and fight duration. We detected fight dynamic differences at the pair level. Higher resource value generated increased aggression but had no effect on fight duration or latency. This conclusion is in line with game theory predictions concerning resource value and contradicts the theory that enrichment of the environment will serve welfare purposes.     

Female–female aggression is linked to food defence in a poison frog

Habitat occupancy by territorial animals is expected to depend on the distribution of critical resources. Knowledge on female territoriality is scarce, but it has been suggested as a mechanism to defend limited resources for reproduction. A previous study showed female intrasexual aggression to be associated with territorial behaviour in the strawberry poison frog Oophaga pumilio, a diurnal aposematic species with complex maternal care. Here, we investigate the link between spatial distribution of resources important for reproduction and female distribution and behaviour. We observed focal females in their natural habitat in Costa Rica, and recorded the distribution of ecological predictor variables in a grid system. We used the data for calculating home range and territory sizes and for connecting female habitat use to the distribution of potential resources by computing spatial habitat occupancy models. Even though we found females to occupy large home ranges, they were highly aggressive towards other females only inside a small part of their home range, here termed core area. Among the ecological factors, the sustained abundance of ants (main food item of the frogs), the presence of leaf litter and suitable rearing sites for tadpoles predicted female site occupancy patterns. The number of ants per grid was twice as high in the core areas compared to the rest of the female home ranges. Our results suggest that female spacing behaviour is principally driven by the spatial distribution of its main food resource, but that hiding places (leaf litter) and tadpole‐rearing sites also play a role. The defence of areas with sustainably high abundance of ants could be relevant, as egg production and maternal care are energetically highly demanding in this prolonged‐breeding species. Regarding the link between resource defence and maternal care, the reproductive strategy of female strawberry poison frogs resembles that of the females of small mammals comprising same‐sex competition for food and high investment in producing and rearing young.     

Spawning behavior of snubnose darters (Percidae) in natural and laboratory environments

Eight species of snubnose darters (genus Etheostoma) were observed and filmed to compare the spawning behavior in the field and in aquaria. These darters exhibited substrate searching, male lateral quivers, and head jabbing behavior in both the field and aquaria. In streams, snubnose darters attached eggs singly to cobble and larger rock substrates, while in aquaria several species were found to attach eggs to fine aquarium gravel as well as cobble and larger rocks. In general, these results support previous aquarium studies of snubnose darters. Information on spawning behavior in snubnose darters as well as darters of the closely related subgenus Etheostoma is synthesized in order to address the phylogenetic affinities of some species in this group of fish.     

Interspecific aggressive behavior in tropical reef stomatopods and its possible ecological significance

Summary Aggressive behavior may play a role in resource partitioning in two Eniwetok stomatopods Gonodactylus incipiens and Haptosquilla glyptocercus. These animals inhabit cavities in coral rubble which they defend vigorously. H. glyptocercus is the more aggressive species and is more than twice as abundant as G. incipiens in the habitat studied. The latter attains larger sizes and also occurs in habitats where H. glyptocercus is rare or absent. A third rarer species, G. falcatus, is also less aggressive than H. glyptocercus and is found in a different microhabitat.     

Influence of hybridization on animal space use: a case study using coyote range expansion

Hybridization between animal species is likely to increase as distributional and reproductive barriers continue to break down due to anthropogenically driven changes in habitat and climate. Yet, the influence of hybridization on ecological interactions and ecosystem function remains understudied. Animal space use, an important component of ecosystem dynamics, is a complex relationship between intrinsic factors, which hybridization can influence, and extrinsic factors, such as environmental heterogeneity. Using the coyote Canis latrans, a well‐studied species with a long history of hybridization with wolves and dogs Canis spp., we sought to assess the influence of hybridization relative to environmental factors in determining animal space use. We conducted a meta‐regression analysis of 67 datasets on coyote home range size across North America and generated models to predict coyote home range size. Climate (latitude) and environmental variability played important roles in determining patterns of coyote space use, likely through their influence on availability of prey resources. However, we found hybridization to be the preeminent factor driving variation in coyote space use, with non‐introgressed populations having considerably smaller home ranges than those from within the Canis hybrid zone of eastern North America. This pattern was upheld despite the variation in environmental factors between areas inside and outside the Canis hybrid zone. Our findings suggest that hybridization may serve as an important factor affecting ecosystems, as hybrids may have altered space requirements, and presumably different niche dimensions, compared to parental species. This, in turn, may influence the role that particular species play within communities. Synthesis Hybridization between animal species is likely to increase due to anthropogenically driven changes in habitat and climate. We used a meta‐regression analysis (n = 67) to examine the relationship between hybridization and coyote Canis latrans space use across North America. We found that independent of environmental factors, introgressed coyotes had larger home ranges. Our findings suggest that hybridization may serve as an important factor affecting ecosystems, as hybrids may have altered space requirements, and presumably different niche dimensions, compared to parental species. This, in turn, may influence the role that particular species play within communities.