Commensalism between cichlid fishes through differential tolerance of guarding parents toward intruders

In the shallow waters of Lake Tanganyika, a benthic cichlid Xenotilapia boulengeri rests in a cluster around the nesting sites of substrate-brooding cichlids Lepidiolamprologus attenuatus and L. elongatus . Experimental removal of parent Lepidiolamprologus guarding free-swimming young drastically reduced the number of X. boulengeri around these nesting sites within a few days, suggesting that their presence has a positive effect on attracting X. boulengeri . Xenotilapia boulengeri were less frequently disturbed by the scale-eating cichlids Perissodus microlepis and Plecodus straeleni when they remained near nesting sites of these species than when stayed away. This was attributed to the territorial behaviour of guarding parents who drove away scale-eaters at greater distances, while allowing X. boulengeri , harmless to their young, to approach the nesting site. It is concluded that a clumped distribution of X. boulengeri results from differential tolerance of breeding parents toward intruders that produces a safety zone from scale-eaters.     

The effect of environmental factors on the nesting and courtship behaviour ofTilapia zillii in Lake Kinneret (Israel)

Tilapia zillii is a polyphilic, guarding nest spawner which is widespread in west and north Africa and in the Middle East. A study of the nesting and courtship behaviour of this species in Lake Kinneret (Sea of Galilee), revealed that nesting and brood care differ according to the exposure of the nesting site to wave action and the nature of the substrate. The nests are simple and parental care is abbreviated in exposed, sandy sites whereas elaborate nests with brood chambers are excavated in sheltered areas with clayey substrates, and parental care is extensive. The implications of these findings to studies on cichlid breeding and the conservation ofT. zillii in Lake Kinneret are discussed.     

The Role of Urine in Parent-Offspring Communication in a Cichlid Fish

The fry of the Midas cichlid, Cichlasoma citrinellum, preferred over blank water, a urine solution from either parent or from a nonparental adult of either sex. They failed to show a preference for the urine of a predator, C. managuense, or for the mucus from the mother. It is possible that steroid titers in urine could provide information on sex, and that peptide chains in urine could indicate species. The chemosensory acuity of fry is high, as they were able to detect a “concentration” of pheromone as low as one sibling per 2 1 of water. This high acuity implies that olfaction rather than gustation mediated the chemosensory responses described above.     

Utilization of the “Nesting” site territory and relationships with conspecific individuals during parental care in the males of masked greenling <Emphasis Type="Italic">Hexagrammos octogrammus</Emphasis> (Hexagrammidae,Scorpaeniformes)

Territorial behavior of males of masked greenling Hexagrammos octogrammus guarding the nest site with eggs until hatching of embryos in the aquarium is studied based on visual observations and videorecording. Formation of defensed territory around the nest and dynamics of aggressive contacts between the guarding male, conspecific individuals, and other fish species are described. Factors determining the shape of the nesting territory, as well as the features of behavior of masked greenling in the aquarium in comparison to behavior of this species in natural conditions, are discussed.     

Antipredator Behaviour and Suppressed Aggression by Convict Cichlids in response to Injury-released Chemical Cues of Conspecifics but not to those of an Allopatric Heterospecific

In aquatic environments, chemical cues serve as an important source of information for the detection of predation risk. Here, we investigate the response of convict cichlids, Cichlasoma nigrofasciatum, to injury-released chemical cues. We exposed pairs of juvenile convict cichlids first to dechlorinated tap water (control), then later to one of two test stimuli: 1. chemical cues from injured convict cichlids; or 2. chemical cues from injured mosquito fish, Gambusia affinis. Gambusia are allopatric and phylogenetically unrelated to convict cichlids. Gambusia skin was used to control for a general response to injured fish. In response to conspecific cues, convict cichlids significantly increased time spent near the bottom of test aquaria and time under a shelter object. In response to Gambusia skin, convict cichlids tended to increase time spent near the tank bottom but did not increase use of the shelter object. There was a trade-off between antipredator and agonistic behaviours. In response to convict cichlid cues, there was a significant reduction in the frequency of approaches and bites. Gambusia skin extract had no significant effect on aggressive behaviour. These data suggest a species-specific antipredator response to conspecific alarm pheromones in a New World cichlid fish and demonstrate a trade-off between predator avoidance and intraspecific aggression. Further, the presence of an alarm response in this model species sets the stage for the use of chemical cues as a research tool to manipulate predation risk in studies of the interaction between predation risk and reproductive behaviour.     

Pairs and Harems in a Cichlid Fish,Lamprologus brichardi1

L. brichardi is a substrate brooding cichlid with facultative polygamy. The social organization was studied in the field for a 6-week period. The mating structure was examined in detail in the laboratory. Two types of social groupings are described:

• 1 Aggregations of sexually mature but nonterritorial fish, also frequently visited by territory holders in the vicinity.
• 2 Reproductive units (families) mainly consisting of the reproducing pair members and offspring from several broods. All family members defend a common territory around the shelter site. Occasionally a male has access to two females each with a separate territory (harem).

The factors influencing mating structure were investigated in the laboratory:

• 1 Females select breeding sites rather than partners.
• 2 Without competitors for breeding sites, and with an equal or nearly equal sex ratio, harems were established nearly as often as pairs.
• 3 Young males are physically able to mate and form a harem; but they are usually prevented from doing so by more competitive (larger) males.
• 4 Competition for breeding sites is not a prime influence on harem formation, although it is of great importance in determining the composition and size of the breeding population.
• 5 Just as many pairs as harems were formed with and without predators, even though, with predators, no young survived.
• 6 In L. brichardi the formation of harems is not predominantly determined by the distribution of suitable spawning sites. The monopolization of females is only slightly influenced by the distance between their territories.
• 7 In L. brichardi it is not necessary for harem formation that the male is bigger than the female.
• 8 Behavioural protocols and data on growth rates, as well as spawning intervals, did reveal any consistent difference between pairs and harmes.

Of the variables tested, male competition for females was therefore the sole determinant of who should mate.     

Impact of heron (<Emphasis Type="Italic">Ardea cinerea</Emphasis> L. and <Emphasis Type="Italic">A. alba</Emphasis> L.) habitat on coastal juvenile fish assemblages in Rybinsk Reservoir

It has been determined that the stocking density of juvenile fish in an open reservoir shallow-water site in the heronry area in the nesting period is higher than that in the reference site. The juvenile roach Rutilus rutilus L., which is one of the dominant species, in the bird impact zone significantly differs in higher values for both the size–weight characteristics and the body weight-growth rates. The habitat-dependent phenotypic differences in various biotopic assemblages of the juvenile fish are revealed. It has been proven that the environmental impact of the heron-colony life activities on the fish populations in the reservoir shallow-water site is temporary, which is limited by the bird-nesting season.     

Theoretical considerations for the maintenance of interspecific brood care by a Nicaraguan cichlid fish: behavioral plasticity and spatial structure

An herbivorous cichlid fish Cichlasoma (Theraps) nicaraguense has been reported to perform brood care for the fry of a piscivorous cichlid Cichlasoma (Nandopsis) dovii in a Nicaraguan lake. It was suggested that interspecific brood care (IBC) evolved to propagate C. dovii as a predator of another herbivorous fish Neetroplus nematopus, which is superior to C. nicaraguense in competition for breeding territory. However, it is still unclear how IBC is maintained in the face of cheaters that do not provide IBC. In this study, to elucidate the conditions for maintenance of IBC, we developed a spatially structured model, assuming a trade-off between reproduction and IBC (i.e., intra- and interspecific competition) and behavioral plasticity in IBC. The model predicted the following maintenance conditions: the probability of expressing (or suppressing) IBC in response to competitor invasion (or exclusion) is low, or, if the probability is high, then IBC is less effective for exclusion of competitors. Furthermore, we also predicted that IBC can be maintained without spatial structure if it is plastic behavior. Our model contributes to understanding of maintenance of IBC by providing empirically testable predictions. We discuss how this model is applicable to more general relationships, such as those in competitive or mutualistic systems.     

Behavior and physiology of coexistence in two species of Dascyllus (Pomacentridae,Teleostei)

Synopsis In the Gulf of Aqaba Dascyllus aruanus and D. marginatus, form distinct populations organised into harems of single males and groups of females, living permanently in Stylophora and Acropora corals. Each harem consists of 4 sexual casts: dominating males, adult females, sex-changing fish and juveniles. The sexual performance of each adult fish in such social units is controlled by the presence or absence of a dominating male. In corals occupied by mixed groups of both species, only those females spawn that belong to the same species as the dominating male. Direct observation in the sea shows that such situations are produced by indiscriminative settling mechanisms of groups of juvenile fish. Evidence is also presented, demonstrating that dispersion of coral colonies suitable for occupancy and predation by piscivorous fish can modulate such complicated reproduction.     

One cost of being gold: selective predation and implications for the maintenance of the Midas cichlid colour polymorphism (Perciformes: Cichlidae)

In the colour‐polymorphic Midas cichlid fish species complex (Amphilophus citrinellus spp.), gold morphs occur at much lower frequencies (< 10%) than dark individuals. This might be surprising because gold coloration is dominant and coded for by a single Mendelian locus. Furthermore, gold individuals are considered to be competitively advantaged over dark ones because they grow faster and win aggressive encounters more often compared to dark individuals of equal size. However, one might expect a cost of being gold in terms of natural selection as a result of predation. We tested whether the Jaguar cichlid (Parachromis managuensis), a major fish predator of Midas cichlids, preys differentially on colour variants of goldfish (Carassius auratus auratus), which were used as a proxy for Midas cichlids because of their similarity in colour. Size‐matched pairs of prey fish (gold and dark) were offered to the predator and the time until the fish were attacked was recorded. The gold morph was attacked first more often (approximately 70%) but not faster than the dark morph. This suggests that the predator perceives the gold individual first, and/or that the predator exhibits a preference or higher motivation to attack the gold prey fish. The increased risk of predation of gold prey fish suggests for the Midas cichlid system that being gold may carry significant costs in terms of natural selection as a result of its major piscivorous predator. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 350–358.     

Sounds Modulate Males’ Aggressiveness in a Cichlid Fish

Acoustic signals are produced in many fish species during agonistic or courtship interactions. A way to test the biological role of these sounds is the use of acoustic playback experiments. However, sounds are usually associated with visual displays and playback experiments performed in fish so far, often failed to match acoustic and visual stimuli. To avoid this mismatch issue, we experimentally separated or coupled visual and acoustic channels to test the role of sounds produced during male–male aggressive interactions in a cichlid fish, Metriaclima zebra. Results show that aggressive behaviour is based on visual stimuli and that acoustic signals alone never trigger aggression. Furthermore, the association between visual and acoustic channels lowers the level of aggressiveness found when fish can only interact visually. This suggests that acoustic signals used during a dispute may complement visual displays to modulate males’ behaviour by reducing their aggressiveness and the risk of escalated fights.     

Cooperative Breeding in the Lake Tanganyika Cichlid Julidochromis ornatus

Cooperative breeding has been described for several cichlids from the genus Julidochromis (Perciformes: Cichlidae) under laboratory conditions, but field evidence is scarce. Here we describe the breeding system of the cichlid Julidochromis ornatus (Boulenger) in Lake Tanganyika (Zambia). Groups defended a breeding shelter under a large flat stone. Smaller group members stayed and fed under or close to the stone, actively guarded by the larger group members. Six out of 28 groups were newly established by breeders, joined by subordinates from a large pool of independent fish (comprising 50–70% of the total population), and four groups were seen to dissolve during a total of 77 observation days. Breeding groups consisted of a large breeding male and female with zero to five smaller subordinates (average 2). Larger breeders and subordinates were found in larger groups. All group members participated in territory defence and -maintenance, but the breeders were only present at the shelter 48% of the time, in contrast to the subordinates which guarded the breeding shelter 94% of the time. Smaller group members showed submissive behaviours to larger group members. We conclude subordinates in J. ornatus are helpers, but we did not find evidence that helpers increased the group’s current reproductive success. Personal observations combined with a literature review revealed at least 19 species of Lake Tanganyika cichlids show evidence of cooperative breeding, entirely confined to the substrate breeding tribe of the Lamprologini (24% of 80 species in total): 2 Chalinochromis spp., 5 Julidochromis spp., 12 Neolamprologus spp. More effort should be put into detecting cooperative breeding in American and Asian substrate breeding cichlid species.     

Social Reversal of Sex‐Biased Aggression and Dominance in a Biparental Cichlid Fish (Julidochromis marlieri)

In biparental species, aggression, dominance, and parental care are typically sexually dimorphic. While behavioral dimorphism is often strongly linked to gonadal sex, the environment—either social or ecological—may also influence sex‐biased behavior. In the biparental cichlid fish Julidochromis marlieri, the typical social environment for breeding pairs consists of large females paired with smaller males. While both sexes are capable of providing territory defense and parental care, the larger female provides the majority of defense for the pair, while the smaller male remains in the nest guarding their offspring. We examine the contributions of sex and relative mate size to these sex‐biased behaviors in monogamous J. marlieri pairs. Both female‐larger and male‐larger pairs were formed in the laboratory and were observed for territorial aggression (against conspecifics and heterospecifics), dominance, and parental care. In female‐larger pairs, territorial aggression and intra‐pair dominance were female‐biased, while in male‐larger pairs this bias was reversed. For both pairing types, the presence of an intruder amplified sex differences in territorial aggression, with the larger fish always attacking with greater frequency than its mate. Though less robust, there was evidence for plasticity of sex‐bias for some egg care related behaviors in the inverse direction. Our study suggests that relative mate size strongly influences the sex bias of aggression and dominance in J. marlieri and that this aspect of the social environment can override the influence of gonadal sex on an individual's behavior. The remarkable plasticity of this species makes Julidochromis an exciting model that could be used to address the relationship between proximate and ultimate mechanisms of behavioral plasticity.     

Intraguild predation between spawning smallmouth bass (Micropterus dolomieu) and nest‐raiding crayfish (Orconectes rusticus): implications for bass nesting success

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Patterns of Fish Use and Piscivore Abundance within a Reconnected Saltmarsh Impoundment in the Northern Indian River Lagoon, Florida

Nearly all saltmarshes in east-central, Florida were impounded for mosquito control during the 1960s. The majority of these marshes have since been reconnected to the estuary by culverts, providing an opportunity to effectively measure exchange of aquatic organisms. A multi-gear approach was used monthly to simultaneously estimate fish standing stock (cast net), fish exchange with the estuary (culvert traps), and piscivore abundance (gill nets and bird counts) to document patterns of fish use in a reconnected saltmarsh impoundment. Changes in saltmarsh fish abundance, and exchange of fish with the estuary reflected the seasonal pattern of marsh flooding in the northern Indian River Lagoon system. During a 6-month period of marsh flooding, resident fish had continuous access to the marsh surface. Large piscivorous fish regularly entered the impoundment via creeks and ditches to prey upon small resident fish, and piscivorous birds aggregated following major fish movements to the marsh surface or to deep habitats. As water levels receded in winter, saltmarsh fish concentrated into deep habitats and emigration to the estuary ensued (200% greater biomass left the impoundment than entered). Fish abundance and community structure along the estuary shoreline (although fringed with marsh vegetation) were not analogous to marsh creeks and ditches. Perimeter ditches provided deep-water habitat for large estuarine predators, and shallow creeks served as an alternative habitat for resident fish when the marsh surface was dry. Use of the impoundment as nursery by transients was limited to Mugil cephalus Linnaeus, but large juvenile and adult piscivorous fish used the impoundment for feeding. In conclusion, the saltmarsh impoundment was a feeding site for piscivorous fish and birds, and functioned as a net exporter of forage fish to adjacent estuarine waters.     

Trophic interrelationships amongst cichlid fishes in a tropical African reservoir (Lake Chivero,Zimbabwe)

The diet of seven cichlid species of Lake Chivero, Zimbabwe, was investigated from October 2004 to April 2005 to determine the degree of dietary overlap and interspecific competition. These fish could be separated into four groups. Two microphages, Oreochromis niloticus and O. macrochir, fed mostly on blue-green algae and their diets, in all size classes, overlapped almost completely. This suggested a high degree of competition between the two species and may account for the fact that O. niloticus, introduced into the lake in the 1980s is now the dominant species. There was also a significant dietary overlap between the two macrophages Tilapia sparrmanii and T. rendalli, which were macrophages with macrophytes as a significant component of their diet. Interspecific competition between these two species may account for the increase in the abundance of T. sparrmanii and a decrease in the numbers of T. rendalli over the last two decades. The diet of two small omnivores Pharyngochromis acuticeps and Pseudocrenilabrus philander consisted of algae, zooplankton, plant material, fish and detritus and also overlapped extensively, but these species probably avoid competition by living in different habitats; P. philander is usually associated with vegetation while P. acuticeps tends to live in more open waters. The last species, Serranochromis robustus, was largely piscivorous. There were no ontogenetic variations in diet, apart from P. acuticeps where small fish fed mainly on zooplankton becoming omnivorous in larger ones, and S. robustus where the diet shifted from zooplankton in specimens <50 mm SL to fish in larger size classes. Data available from the 1960s and 1970s indicate that the diets of some species have changed with invertebrates such as chironomid larvae being previously more important in all diets, while most species now take a higher proportion of algae. This reflects changes in the lake, which is eutrophic and has experienced severe algal blooms since 1960. Handling editor: C. Strumbauer     

Initial Six-year Expansion of an Introduced Piscivorous Fish in a Tropical Central American Lake

Two populations of the piscivorous fish Gobiomorus dormitor (the guabina) were examined in two separate crater lakes in Nicaragua, Central America. At Lake Apoyo, G. dormitor were introduced in May 1991 by local fishermen and have invaded successfully; at Lake Xiloá, the population is naturally occurring. To provide baseline life history data for G. dormitor in both lakes and to test hypotheses related to population growth and invasion success, this study aimed to (1) document and define the population growth and depth distribution of the fish through video transects in Lake Apoyo; (2) compare diet composition between lakes; and (3) compare life history parameters between lakes, including relative growth rates and reproduction. Videotaped transect studies, designed originally to examine cichlid breeding, began in both lakes in 1990 and have documented the subsequent increase of G. dormitor. Yearly means generally fit the exponential growth model. G. dormitor foraged mainly in shallow waters in Lake Apoyo and preyed mostly upon the atherinid Melaniris cf. sardina throughout seasons. In Lake Xiloá, cichlid fry constituted the majority of the diet overall and cannibalism was more common, although this trend varied with the cichlid breeding season. Relative growth rates were significantly different, based on otolith mass regressions on standard length. Lake Apoyo G. dormitor generally behaved as a successful invading species, displaying exponential growth, year-round reproduction, and higher foraging success than the natural population of Lake Xiloá. Further research is planned to explore the G. dormitor invasion when the population becomes integrated into the community.     

Cleaning behaviour of the cichlid Mesonauta festivus in the Pantanal wetlands: evidence of a potential freshwater cleaning station

Cleaning interactions are known among several groups of fishes, with a higher number of records for marine fish species. These temporary associations occur between one species that acts as the cleaner and the other species as the client. The interaction usually takes place within the boundaries defined by the cleaner. This site, known as the cleaning station, allows the client fish to strike a typical pose allowing the cleaner to approach, such as wide-open fins and inclined, motionless body. Few studies have reported on the cleaning behaviour of freshwater fish species, and none has reported on behaviour that could be interpreted as establishing a cleaning station. Herein we present a new record of the cleaning interaction in the Neotropics, between the cichlid Mesonauta festivus as the cleaner and three species of anostomid (Leporinus macrocephalus, Leporinus friderici and Schizodon borellii) as clients on Pantanal wetlands, as well as the establishment of a defined cleaning station.     

Effects of loss of male parent on brood survival in a biparental cichlid fish

Synopsis When brood-guarding males of the biparental convict cichlid, Cichlasoma nigrofasciatum, held in experimental ponds, deserted their mate and brood, or were removed from the pond, survival of the young depended on their age when the father left. If they were embryos or free embryos, few broods survived, but if they were up to seven days into the free-swimming juvenile period, most broods survived. The cause of brood mortality is not certain, but predation by conspecifics is most likely (no other fish species were in the ponds). The willingness of a deserted, brood-guarding female to continue defending her young probably depends, in part, on their age. If they are very young, the benefit/cost ratio of guarding them to independence may be so low that she should give up that breeding attempt and begin another. If they are older, the benefit/cost ratio is higher and she should continue to guard them alone.