Habitat structure directly affects aggression in convict cichlids Archocentrus nigrofasciatus

Aggressive behavior can be an important factor in determining how animals use and divide space and resources. Previous studies have shown that aggression in fishes can be influenced by a variety of factors, including water temperature and resource levels. In this study, we tested if the amount of habitat structure in the environment affected aggression levels in female convict cichlids Archocentrus nigrofasciatus. We performed a laboratory experiment in which we placed female convict cichlids into an aquarium with low or high amounts of habitat structure and monitored the dominant female's behavior toward the subordinate female. Aggressive behavior in convict cichlids primarily consists of chases and bites. We found that the total time the dominant female spent chasing the subordinate female was greater when there was a low amount of habitat structure as compared to when there was a high amount of habitat structure. We also found that both the average duration of a chasing bout and the number of bites directed at the subordinate fish increased when there was a low amount of structure, but the number of chases did not. These results indicate that increased habitat structural complexity decreases aggressive behavior in convict cichlids [Current Zoology 56 (1): 52-56, 2010].     

Experimentally induced helper dispersal in colonially breeding cooperative cichlids

The ‘benefits of philopatry’ hypothesis states that helpers in cooperatively breeding species derive higher benefits from remaining home, instead of dispersing and attempting to breed independently. We tested experimentally whether dispersal options influence dispersal propensity in the cooperatively breeding Lake Tanganyika cichlids Neolamprologus pulcher and N. savoryi. Cooperative groups of these fishes breed in densely packed colonies, surrounded by unoccupied, but apparently suitable breeding habitat. Breeding inside colonies and living in groups seems to benefit individuals, for example by early detection and deterrence of predators. We show that despite a slight preference of both species for habitat with a higher stone cover, 40% of the preferred habitat remained unoccupied. On average, the colonies contained a higher number of (1) predators of adults, juveniles and eggs, (2) shelter competitors, and (3) other species including potential food competitors, compared to the outside colony habitat. Apparently, habitat differences cannot explain why these cichlids breed in colonies. Accordingly, dispersal may not be limited by a lack of suitable breeding shelters, but by the relatively higher risk of establishing an outside- compared to a within-colony breeding territory. To test whether cichlids prefer within- to outside-colony breeding territories, we provided breeding shelters inside the colony and at the colony edge and studied helper dispersal. As expected, significantly more shelters were occupied within the colony compared to the edge. New breeding pairs with several helpers occupied these shelters. We conclude that although breeding habitat is plentiful outside the colonies, helpers delay dispersal to obtain a higher quality breeding position within the group or colony eventually, or they disperse in groups. Our results suggest that (1) group augmentation and Allee effects are generally important for dispersal decisions in cooperatively breeding cichlids, consistent with the ‘benefits of philopatry hypothesis’, and (2) habitat saturation cannot fully explain delayed dispersal in these species.     

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.     

Habitat light sets the boundaries for the rapid evolution of cichlid fish vision,while sexual selection can tune it within those limits

Cichlid fishes' famous diversity in body coloration is accompanied by a highly diverse and complex visual system. Although cichlids possess an unusually high number of seven cone opsin genes, they express only a subset of these during their ontogeny, accounting for their astonishing interspecific variation in visual sensitivities. Much of this diversity is thought to have been shaped by natural selection as cichlids inhabit a variety of habitats with distinct light environments. Also, sexual selection might have contributed to the observed visual diversity, and sexual dimorphism in coloration potentially co‐evolved with sexual dimorphism in opsin expression. We investigated sex‐specific opsin expression of several cichlids from Africa and the Neotropics and collected and integrated data sets on sex‐specific body coloration, species‐specific visual sensitivities, lens transmission and habitat light properties for some of them. We comparatively analysed this wide range of molecular and ecological data, illustrating how integrative approaches can address specific questions on the factors and mechanisms driving diversification, and the evolution of cichlid vision in particular. We found that both sexes expressed opsins at the same levels—even in sexually dimorphic cichlid species—which argues against coevolution of sexual dichromatism and differences in sex‐specific visual sensitivity. Rather, a combination of environmental light properties and body coloration shaped the diversity in spectral sensitivities among cichlids. We conclude that although cichlids are particularly colourful and diverse and often sexually dimorphic, it would appear that natural rather than sexual selection is a more powerful force driving visual diversity in this hyperdiverse lineage.     

The zoogeography, ecology and taxonomy of the genus Labeotropheus Ahl, 1927, of Lake Malawi (Pisces: Gichlidae)

Two species of Labeotropheus are currently recognized, but an earlier study of aquarium specimens caused doubt to be cast upon their specific distinctness. This doubt was removed by a field and laboratory study which showed that L. fuelleborni differed from L. trewavasae morphologically, in coloration, in geographic distribution, preferred habitat and ability to penetrate depth. The two species practise assortative mating.
Labeotropheus fuelleborni feeds predominantly from upper surfaces of rocks while L. trewavasae favours under surfaces. Although both species are specialized to crop epilithic algae they also feed secondarily on other food resources (e.g. plankton, insects) when these are available.
Gene flow between many populations of rock-frequenting cichlids is restricted by their sedentary nature and apparent inability to cross open water between rocky areas. Consequently, numerous geographic races, incipient species and species are found in the lake. This suggests that intralacustrine allopatric speciation may account for much of the species diversity of cichlids of rocky zones.
It is recommended that biologists working on cichlids from Lake Malawi should ascertain the precise location at which their specimens were caught and avoid using aquarium reared fish.     

Effects of habitat complexity,dominance and personality on habitat selection: Ideal despotic cichlids

Habitat structure can impede visibility and movement, resulting in lower resource monopolization and aggression. Consequently, dominant individuals may prefer open habitats to maximize resource gain, or complex habitats to minimize predation risk. We explored the role of dominance on foraging, aggression and habitat choice using convict cichlids (Amatitlania nigrofasciata) in a two‐patch ideal free distribution experiment. Groups of six fish of four distinct sizes first competed for shrimp in one‐patch trials in both an open and complex habitat; half the groups experienced each habitat type first. Following these one‐patch trials, each group then chose between habitat types in a two‐patch trial while competing for food. Finally, each fish underwent an individual behavioural assessment using a battery of “personality” tests to determine if behaviour when alone accurately reflected behaviour within a social context. In the one‐patch trials, dominant fish showed similar food consumption between habitats, but chased more in the complex habitat. In the two‐patch choice trials, dominants preferred and defended the complex habitat, forming an ideal despotic distribution with more than half the fish and competitive weight in the open habitat. Within the groups, individual fish differed in foraging and chasing, with repeatabilities of 0.45 and 0.23 across all treatments. Although a higher foraging rate during the individual assessment predicted foraging rate and use of the complex habitat during the group trials, aggression and boldness tests were not reflective of group behaviour. Across groups, heavier dominants and those with higher foraging rate in the open habitat used the open habitat more, suggesting that both risk and energetic state affect habitat preference in dominant convict cichlids.     

From shallow to deep divergences: mixed messages from Amazon Basin cichlids

Cichlids are a conspicuous component of Amazonian ichthyofauna, filling a wide range of niches. Yet taxonomy of many groups is still poorly known in the Amazon, and most of the yet-to-be discovered species are concentrated there. We analyzed 230 individuals sampled from six major Amazonian River Basins representing 56 morpho-species, 34 nominal and 22 undescribed species in 18 cichlid genera. We used four different single-locus species-discovery (SLSD) methods, delimiting between 53 (mPTP) and 57 (GMYC) species/lineages. When detected, species/lineages are hierarchically geographically structured. Many groups such as the Geophaginae and the Cichlinae have recently diversified, and species of genera such as Cichla and Symphysodon hybridize or have a history of hybridization; thus, these species will not be detected by SLSD methods. At the same time, for example, the genera Apistogramma and Biotodoma harbor cryptic species. For all these reasons, species/lineage diversity of Amazonian cichlids is significantly underestimated. The diversity of Amazonian cichlids is particularly remarkable given that the 570 species of Neotropical cichlids, many of which are from the Amazon Basin, are found in just 1.7% of the freshwater aquatic habitat in which the ~ 2,000 species of the East African rift lake cichlids evolved.

     

Lake Malawi cichlid evolution along a benthic/limnetic axis

Divergence along a benthic to limnetic habitat axis is ubiquitous in aquatic systems. However, this type of habitat divergence has largely been examined in low diversity, high latitude lake systems. In this study, we examined the importance of benthic and limnetic divergence within the incredibly species‐rich radiation of Lake Malawi cichlid fishes. Using novel phylogenetic reconstructions, we provided a series of hypotheses regarding the evolutionary relationships among 24 benthic and limnetic species that suggests divergence along this axis has occurred multiple times within Lake Malawi cichlids. Because pectoral fin morphology is often associated with divergence along this habitat axis in other fish groups, we investigated divergence in pectoral fin muscles in these benthic and limnetic cichlid species. We showed that the eight pectoral fin muscles and fin area generally tended to evolve in a tightly correlated manner in the Lake Malawi cichlids. Additionally, we found that larger pectoral fin muscles are strongly associated with the independent evolution of the benthic feeding habit across this group of fish. Evolutionary specialization along a benthic/limnetic axis has occurred multiple times within this tropical lake radiation and has produced repeated convergent matching between exploitation of water column habitats and locomotory morphology.     

Extreme microallopatric divergence in a cichlid species from Lake Malawi

We demonstrate significant population structuring on an extremely small spatial scale between adjacent demes of a Lake Malawi haplochromine cichlid species of the mbuna group, Pseudotropheus callainos, separated by only 35 m of habitat discontinuity. This substantiates further the notion that intralacustrine allopatric divergence may help to explain the high level of species richness of the mbuna in comparison to other Malawian cichlids, as well as of the Malawian haplochromines as a whole.     

Sex differences in biparental care as offspring develop: a field study of convict cichlids (<Emphasis Type="Italic">Amatitlania siquia</Emphasis>)

Parental investment theory states that parents should contribute more to older offspring. Differences between the sexes also influence how each parent contributes to offspring in biparental species. Here, we examined a naturally occurring population of biparental convict cichlids in Costa Rica to determine how each parent cared for offspring during two distinct offspring development stages. Consistent with the predictions of the reproductive value hypothesis, we hypothesized that the levels of parental contribution would be relative to the value that each parent places on a brood. We predicted that female parents would contribute more than male parents because female convict cichlids have lower future reproductive success than males. Additionally, we predicted that both parents should contribute more to older offspring, either due to the young’s increased susceptibility to predation (i.e., the vulnerability hypothesis) or because of the longer period of time parents have been interacting with older offspring (i.e., feedback hypotheses). This increase in investment by males should coincide with a change in the coordination of care between parents. Detailed observations of parental pairs in their natural habitat supported these predictions. Females contributed more to broods than males and were relatively unaffected by offspring age while males spent significantly more time with older, free-swimming fry. Additionally, males tended to leave younger offspring more than females did, and were more likely to do so consecutively with younger offspring. This suggests that the coordination of duties between parents changes as parental investment changes. Overall, these data support both the reproductive value and the vulnerability hypotheses, but not necessarily the feedback hypothesis.     

Habitat use and its implications to functional morphology: niche partitioning and the evolution of locomotory morphology in Lake Tanganyikan cichlids (Perciformes: Cichlidae)

Animal locomotory morphology, i.e. morphological features involved in locomotion, is under the influence of a diverse set of ecological and behavioral factors. In teleost fish, habitat choice and foraging strategy are major determinants of locomotory morphology. In this study, we assess the influence of habitat use and foraging strategy on important locomotory traits, namely the size of the pectoral and caudal fins and the weight of the pectoral fin muscles, as applied to one of the most astonishing cases of adaptive radiation: the species flock of cichlid fishes in East African Lake Tanganyika. We also examine the course of niche partitioning along two main habitat axes, the benthic vs. limnetic and the sandy vs. rocky substrate axis. The results are then compared with available data on the cichlid adaptive radiation of neighbouring Lake Malawi. We find that pectoral fin size and muscle weight correlate with habitat use within the water column, as well as with substrate composition and foraging strategies. Niche partitioning along the benthic–limnetic axis in Lake Tanganyikan cichlids seems to follow a similar course as in Lake Malawi, while the course of habitat use with respect to substrate composition appears to differ between the cichlid assemblages of these two lakes.     

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 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.     

Systematics and historical biogeography of Greater Antillean Cichlidae

A molecular phylogenetic analysis recovers a pattern consistent with a drift vicariance scenario for the origin of Greater Antillean cichlids. This phylogeny, based on mitochondrial and nuclear genes, reveals that clades on different geographic regions diverged concurrently with the geological separation of these areas. Middle America was initially colonized by South American cichlids in the Cretaceous, most probably through the Cretaceous Island Arc. The separation of Greater Antillean cichlids and their mainland Middle American relatives was caused by a drift vicariance event that took place when the islands became separated from Yucatan in the Eocene. Greater Antillean cichlids are monophyletic and do not have close South American relatives. Therefore, the alternative hypothesis that these cichlids migrated via an Oligocene landbridge from South America is falsified. A marine dispersal hypothesis is not employed because the drift vicariance hypothesis is better able to explain the biogeographic patterns, both temporal and phylogenetic.     

Positive selection of the long-wavelength opsin gene in South American cichlid fishes

Hydrobiologia - Cichlid fishes use their vision for recognition of other fishes, feeding, and in habitat selection. Neotropical cichlids can be found in a range of different habitats differing in...     

Paradoxical role of C1561T glutamate carboxypeptidase II (GCPII) genetic polymorphism in altering disease susceptibility

Although olfaction could play a crucial role in underwater habitats by allowing fish to sense a variety of nonvolatile chemical signals, the importance of olfaction in species-rich cichlids is still controversial. In particular, examining whether cichlids rely on olfaction for reproduction is of primary interest to understand the mechanisms of speciation. In the present study, we explored the V1R (also known as ora) genes, which are believed to encode reproductive pheromone receptors in fish, in the genomes of Lake Victoria cichlids. By screening a bacterial artificial chromosome library, we identified all six intact V1R genes (V1R1 to V1R6) that have been reported in other teleost fish. Furthermore, RT-PCR and in situ hybridization analyses showed that all of the V1R genes were expressed in the olfactory epithelium, indicating that these receptors are functional in cichlids. These observations indicate that cichlids use V1R-mediated olfaction in some ways for their social behaviors.     

Acoustic diversity in Lake Malawi’s rock-dwelling cichlids

The cichlids of Lake Malawi are one of the world’s most species rich and phenotypically diverse groups of extant vertebrates. The extraordinary variability of this group’s color patterns, reproductive behaviors, and trophic morphologies are well documented. More recently, an additional axis of phenotypic diversity has been identified. Lake Malawi cichlids have been shown to use species-specific acoustic communication in both aggressive and reproductive encounters. However, documentation of acoustic signals used by this group is limited to a small number of taxa observed within the confines of the laboratory. This study examines the acoustic signals produced by six species spanning four genera of rock-dwelling cichlids recorded in their natural habitat, the shallow waters surrounding Thumbi West Island, Lake Malawi. Four acoustic parameters were quantified and compared between species: trill duration, number of pulses per trill, pulse duration, and pulse period. Using these characteristics, sympatric species within the genus Maylandia were easily distinguished. Furthermore, a comparison of this data to previously published acoustic data reveals possible geographic dialects within species.     

Trophic divergence of Lake Kivu cichlid fishes along a pelagic versus littoral habitat axis

Local adaptation to the littoral and pelagic zones in two cichlid haplochromine fish species from Lake Kivu was investigated using morphometrics. Cranial variation and inferred jaw mechanics in both sexes of the two species across the two habitat types were quantified and compared. Comparisons of littoral versus pelagic populations revealed habitat‐specific differences in the shape of the feeding apparatus. Also, kinematic transmission of the anterior jaw four‐bar linkage that promotes greater jaw protrusion was higher in the pelagic zone than in the littoral zone for both species. Inferred bite force was likewise higher in pelagic zone fish. There were also sex‐specific differences in craniofacial morphology as males exhibited longer heads than females in both habitats. As has been described for other cichlids in the East African Great Lakes, local adaptation to trophic resources in the littoral and pelagic habitats characterizes these two Lake Kivu cichlids. Similar studies involving other types of the Lake Kivu fishes are recommended to test the evidence of the observed trophic patterns and their genetic basis of divergences.     

Stream fish assemblage and habitat structure in a tropical African river basin (Nyagui River, Zimbabwe)

Few studies have been carried out on stream ecology in southern Africa although many species are endangered. This study investigated the stream fish assemblage and their habitat associations over a period of 3 months (October 2004 to January 2005), in view of the proposal to build a dam across the Nyagui River. Twenty-four fish species were collected and were separated into groups based on preferred microhabitats. The first group, dominated by Barbus paludinosus , comprised species collected from the upstream stations with slow flow, shallow depth (pools) and fine substrate type. Species associated with riffles, which included Chiloglanis neumanni , Labeobarbus marequensis and Opsaridium zambezense , comprised the second group on the downstream. The last group comprised species preferring pools with rock substrate and slow flow such as Pharyngochromis acuticeps and Pseudocranilabrus philander . The species were consistently associated with their habitat types throughout the sampling period. This relationship may be explained by the fish's morphological adaptations. Species richness increased from nine in the upstream section to twenty in the downstream section and this was related to increasing habitat complexity downstream. The construction of the Kunzvi Dam across the Nyagui River is likely to lead to loss of rheophilic species while cichlids and introduced species may increase.