The Nesting Site of a Piscivorous Cichlid Lepidiolamprologus profundicola as a Safety Zone for Juveniles of a Zooplanktivorous Cichlid Cyprichromis leptosoma in Lake Tanganyika

Juvenile Cyprichromis leptosoma, an endemic cichlid fish in Lake Tanganyika, form large schools near the nesting sites of the piscivorous cichlid Lepidiolamprologus profundicola. The female L. profundicola guarding the nesting site drove away piscivorous fish that approached the nest. After the disappearance of the guarding L. profundicola, the frequency of predatory dashes toward the school of juvenile C. leptosoma near the nesting site increased markedly. This suggests that juvenile C. leptosoma use the breeding territory of L. profundicola as a safety zone from their potential predators. The relationship between C. leptosoma and L. profundicola should be regarded as a type of commensalism.     

Parental care in a monogamous mouthbrooding cichlidXenotilapia flavipinnis in Lake Tanganyika

Breeding pairs ofXenotilapia flavipinnis held their territories on the sandy bottom and repeated several breeding cycles. Females mouthbrooded the eggs and early larvae but afterward males took over the mouthbrooding role. When the young became free swimming, they were guarded by both parents and remained in the spawning territory. Males played a leading role in the guarding, while females were more active in foraging during the guarding period. It was concluded that males’ active participation in the parental care could accelerate the gonadal recovery of females and consequently could maximize the fecundity of serially monogamous pairs.     

Cichlid-catfish mutualistic defense of young in Lake Malawi,Africa

Summary Bagrid catfish and cichlid parents engage in a mutualistic defense of their young from predators. Over 50% of the catfish broods observed contained cichlid young, primarily of three species: Crytocara pleurostigmoides, C. pictus and Rhamphocromis sp. Three catfish broods, monitored for over 50 days, had a survivorship rate 6 times greater during periods when cichlids were present than when cichlids were absent. During two and a half h of observations of catfish broods without cichlids, I observed 23 strikes by predators on catfish young. However, when cichlids were present no foraging attacks were observed upon the catfish, but 32 occurred against cichlids in two and a half hours of observations. When the catfish parents were experimently removed the cichlid young were consumed first then the catfish young. When cichlids were present the catfish young survived over 80% longer than those in broods without cichlids. It is concluded that the parental catfish increase the survival of their own young be allowing cichlids into the brood. Because of an asymmetry of costs between the two species, cichlid mothers are more likely to abandon these interspecific broods than are the catfish.     

The cichlid fishes of Guyana, South America, with notes on their ecology and breeding behaviour

Field studies in Guyana, north-eastern South America, from 1957 to 1962 included observations on the ecology and behaviour of cichlids in natural waters. Collections made were examined at the British Museum (Natural History) together with other cichlid material from Guyana. This paper records these observations and gives keys for the field indentification of these cichlids. Fifteen genera comprising at least 35 species occur in Guyana, mostly species with a wide distribution in tropical South America. In the predatory Cichla ocellaris which grows larger than other South American cichlids, growth continues long after the fish matures. Males tend to be larger than associated females. The behaviour of Cichla ocellaris and Cichlasoma festivum when guarding their young suggests that the caudal ocellus has a vital role in the orientation of young to parent. Most of these cichlids in which breeding habits are known are substratum-spawners. An attachment gland, producing viscous threads, was found on the heads of recently hatched Geophagus jurupari young, although this species carries the young in the mouth.     

Interspecific brood-mixing in Tanganyikan cichlids

Synopsis We collected schools of young, guarded by parents, of six common cichlid species to investigate the frequency and origin of interspecific brood-mixing. The main host species were a piscivore Lepidiolamprologus elongatus and a scale-eater Perissodus microlepis; more than half of their schools included heterospecific young, accounting for 20–40% of the total young. Most of the foreign young belonged to four biparental mouth-brooders whose parents have a habit of carrying their young in their mouths. Many of these young were smaller than the largest young brooded by their own parents. We concluded that adoption of young before independence results from farming-out, a behavior by which parents actively transfer their young to foster parents.     

Phylogenetic relationships of the lamprologine cichlid genus Lepidiolamprologus (Teleostei: Perciformes) based on mitochondrial and nuclear sequences, suggesting introgressive hybridization

Using sequences of the mitochondrial NADH dehydrogenase subunit 2 gene (ND2, 1047bp) and a segment of the non-coding mitochondrial control region, as well as nuclear sequences including two introns from the S7 ribosomal protein and the loci TmoM25, TmoM27, and UME002, we explore the phylogenetic relationships of Lepidiolamprologus, one of seven lamprologine cichlid genera in Lake Tanganyika, East Africa. Analyses consisted of direct optimization using POY, including a parsimony sensitivity analysis, and maximum likelihood and Bayesian inference for comparison. With respect to Lepidiolamprologus, the results based on the mitochondrial dataset were robust to parameter variation in POY. Lepidiolamprologus cunningtoni was resolved in a large clade sister to ossified group lamprologines, among which the remaining Lepidiolamprologus were nested. In addition to L. attenuatus, L. elongatus, L. kendalli, and L. profundicola, Neolamprologus meeli, N. hecqui, N. boulengeri, N. variostigma, and two undescribed species were resolved in a two-pore Lepidiolamprologus clade sister to Lamprologus callipterus and two species of Altolamprologus. Lepidiolamprologus nkambae, in marked conflict with morphological and nuclear DNA evidence, nested outside of the two-pore Lepidiolamprologus clade, suggesting that the mtDNA signal has been convoluted by introgressive hybridization.     

FACTORS AFFECTING THE AGE OF FIRST BREEDING OF THE KITTIWAKE RISSA TRIDACTYLA

At a Kittiwake colony in Northumberland, 80% of those birds which returned to their natal colony to breed were males and these supplied 52% of all male recruits. More females breed away from their natal colony than males. There was no differences in the proportions of young fledged from sites in the centre or at the edge of the colony, or by parents of different experience, which returned to breed. Kittiwakes breed for the first time at ages from 3 to 8 years, but most at 4 or S years old. Males arrive back at the colony at an earlier age than females and breed for the first time one year earlier. Males obtaining sites at the centre of the colony first breed at an earlier age than those at the edges. Neither the age nor the area of first breeding appear to be transmitted from parent to offspring. Males breeding first aged 4 years or younger produced more young than those which first bred aged 5 years or older, despite their partners laying smaller clutches. This difference was most marked among those males recruited to sites in the centre of the colony. The advantage of this earlier breeding is counteracted by a lower survival rate among those males which start to breed at the younger ages. In all breeding Kittiwakes, annual reproductive output increases with experience while annual survival rates decrease. Once they had started to breed, many birds failed to breed in one subsequent season. Nearly 60% of these cases of intermittent breeding occurred in the year following first breeding. Intermittent breeding was most frequent among young birds and among females. It is suggested that each breeding involves a cost to the individual in terms of reduced survival, and that deferred and intermittent breeding are means of guarding survival. A model is proposed whereby the age at which a bird starts to breed, the nesting site which it obtains, and its subsequent breeding strategy result in each individual producing an optimal number of reproducing offspring in its lifetime, relative to its quality.     

Intraspecific brood-mixing of the cichlid fishPerissodus microlepis in Lake Tanganyika

Synopsis The frequency and origin of intraspecific brood-mixing in the biparental cichlid fishPerissodus microlepis were investigated by the cohort analysis of schooling young and the underwater observation of guarding parents. The cohort analysis showed that brood-mixing started from the early guarding state when the young were smaller than 10 mm standard length and nearly all schools of young larger than 16 mm contained alien young from up to 6 broods. Brood farming-out of this fish, which was originally proposed to be a way adopted only by a deserted parent, was performed also by paired parents. We suggest that brood-mixing inP. microlepis is attributed mainly to brood farming-out by paired and deserted parents.     

Farming-out of offspring is a predominantly male tactics in a biparental mouthbrooding cichlid <Emphasis Type="BoldItalic">Perrisodus mircolepis</Emphasis>

Synopsis Intraspecific brood-mixing of a Tanganyikan biparental cichlid fish Perissodus microlepis has been attributed to farming-out, a behaviour in which a parent transport some or all of their free-swimming young in the mouth and put them under the care of other guarding parents. We conducted observations under natural conditions to understand how often farming-out occurs and which sex is the primary performer. At most brooding sites, young in a school entirely disappeared before attaining to the size of independence. At many of these sites, farming-out and its precursory behaviours by males preceded disappearance of their young. Females never performed farming-out. We proposed that farming-out by males is an alternative tactics to brood desertion to evade paternal roles for remating.     

Reproductive tactics of the Asian cichlids of the genusEtroplus in Sri Lanka

Synopsis The Asian cichlids,Etroplus maculatus (the orange chromide) and E. suratensis (the green chromide) in Sri Lanka reproduce twice during the year when water conditions are favorable for nest construction and maintaining visual contact with offspring. These are the drier premonsoonal and monsoonal seasons when water turbidity decreased and salinity increased. When breeding in isolation orange chromide pairs selected dense vegetation where nests were camouflaged. During the peak breeding cycle (July) orange chromide pairs selected sparse vegetation for nesting as a compromise between survival of young and availability of adult food. These areas were also occupied by foraging non-breeding conspecifics which increased the threat of cannibalism of offspring. Under these pressures most orange chromides nested in colonies which helped decrease both actual and attempted cannibalism. The sympatric green chromide does not forage during nesting and nest site selection was determined mainly by factors favoring offspring survival. Biparental care is exhibited by both species. One member of an orange chromide pair stands guard over offspring while the other leaves the territory to forage — their roles are reversed every few minutes. The total parental investment is equivalent to the full investment of a single parent. Monogamy appears to be maintained by their metabolic constraints. In the green chromide both parents are vigilant over offspring and neither forages thus spending twice as much time in parental investment.     

Biparental mouthbrooding and guarding in a Tanganyikan CichlidHaplotaxodon microlepis

Differences between parental roles of males and females inHaplotaxodon microlepis (Cichlidae) were investigated in Lake Tanganyika, and the early ontogeny and growth of the species were studied in the aquarium. Eggs were mouthbrooded by the female, and it is suggested that small larvae (<9 mm in total length) were also mouthbrooded by females though such samples were not collected. Above this size the larvae began to feed, and parents jointly performed mouthbrooding and guarding until the young grew to 25–30 mm, nearly 2 months after spawning. Males and females mouthbrooded to the same extent, but when a part of the brood was released, females mainly guarded the released brood and males took the mouthbrooding role. Differences in parentalcare patterns betweenH. microlepis and other monogamous mouthbrooding cichlids are discussed.     

Gut Microbiota Dynamics during Dietary Shift in Eastern African Cichlid Fishes

The gut microbiota structure reflects both a host phylogenetic history and a signature of adaptation to the host ecological, mainly trophic niches. African cichlid fishes, with their array of closely related species that underwent a rapid dietary niche radiation, offer a particularly interesting system to explore the relative contribution of these two factors in nature. Here we surveyed the host intra- and interspecific natural variation of the gut microbiota of five cichlid species from the monophyletic tribe Perissodini of lake Tanganyika, whose members transitioned from being zooplanktivorous to feeding primarily on fish scales. The outgroup riverine species Astatotilapia burtoni, largely omnivorous, was also included in the study. Fusobacteria, Firmicutes and Proteobacteria represented the dominant components in the gut microbiota of all 30 specimens analysed according to two distinct 16S rRNA markers. All members of the Perissodini tribe showed a homogenous pattern of microbial alpha and beta diversities, with no significant qualitative differences, despite changes in diet. The recent diet shift between zooplantkon- and scale-eaters simply reflects on a significant enrichment of Clostridium taxa in scale-eaters where they might be involved in the scale metabolism. Comparison with the omnivorous species A. burtoni suggests that, with increased host phylogenetic distance and/or increasing herbivory, the gut microbiota begins differentiating also at qualitative level. The cichlids show presence of a large conserved core of taxa and a small set of core OTUs (average 13–15%), remarkably stable also in captivity, and putatively favoured by both restricted microbial transmission among related hosts (putatively enhanced by mouthbrooding behavior) and common host constraints. This study sets the basis for a future large-scale investigation of the gut microbiota of cichlids and its adaptation in the process of the host adaptive radiation.     

Parental strategy of the cichlid fish Perissodus microlepis,with particular reference to intraspecific brood ‘farming out’

Synopsis One parent was experimentally removed from brooding pairs of Perissodus microlepis in the field. The removal elicited drastic behavioural changes in the remaining parent and young. The remaining parent sometimes showed a sequence of peculiar behaviours which were not observed when guarding the young with its mate. Some left the brooding site with young in their mouths and then put the young under the care of another pair of brooding parents.     

Sex Differences in Retrieval Behavior by the Biparental Convict Cichlid

Biparental species occasionally demonstrate a division of roles in which parents perform sex-typical tasks, with females offering direct care and spending the majority of their time with the offspring while males are more indirect in their care, providing the majority of defense against potential brood predators. To examine the flexibility in the sex-typical roles shown by convict cichlids ( Amatitlania nigrofasciata ), we displaced non-swimming young at three different distances from the nest and then analyzed the retrieval behaviors of each parent. Retrieval of altricial young is a behavior commonly used to measure parental care in mammalian studies, but has rarely been used in other taxa. We found sex differences in retrieval behavior: on average, females retrieved young close to the nest and males retrieved young far from the nest. This difference in parental contribution suggests a division of labor with sex-specific roles. Sex differences may be due to proximity to young and/or apparent risk of offspring predation. Additionally, we found that latency to first retrieval and total time spent retrieving young remained consistent across the various displacement distances, suggesting that retrieval is an essential parental behavior. Additionally, we include observations of wriggler retrieval by parents in a natural population of Costa Rican convict cichlids.     

A new linkage group involved in sex determination in haplotilapiine cichlids

The high diversity of sex chromosomes and sex determination systems among haplotilapiines suggests that this large cichlid clade is a good model for investigating the evolution of genetics of sex determination. Nonetheless, information about sex determination in this clade remains sparse. The present study reports a microsatellite marker that is closely associated with sex in Xenotilapia rotundiventralis from Lake Tanganyika, Africa. This study is the first to suggest the role of linkage group 17 in sex determination in haplotilapiine cichlids.     

Do pinyon jays alter nest placement based on prior experience?

The role of prior experience in nest-site selection by a long-lived corvid, the pinyon jay, Gymnorhinus cyanocephalus, was investigated. The major sources of nest failure were loss to avian predators (Corvus corax and Corvus brachyrhynchos) and abandonment after cold and snowy spring weather. Cold weather favours exposed nesting because solar radiation reduces the energetic costs to nesting females and quickly melts snow in and around the nest. Predation favours cryptic nests. The relative height at which individual jays nested (an index of nest exposure) was compared to their previous nest height and to the fate of that nesting attempt. For the following reasons jays appear to associate nest exposure with the fate of a particular nesting attempt: (1) after nesting in exposed sites, subsequent nests were 27·3% lower (more concealed) in the nest tree following predation, but only 9·7% lower when predation did not occur; (2) concealed nest sites were avoided only after failure due to cold weather; and (3) nest placement following the successful fledging of young did not differ significantly from the previous nest placement. The frequency of nesting in exposed locations dropped from 80% to 55% after individuals suffered their third predatory experience when nesting in exposed locations. Experienced jays nested relatively low throughout the season, which enhances concealment, but nested farther out from the trunk early in the season, which reduces incubation costs. The use of prior experience in nest-site selection is adaptive because sites can include properties associated with past success and exclude those associated with past failure.     

ASPECTS OF THE BREEDING BIOLOGY OF HERRING GULLS LARUS ARGENTATUS IN URBAN COLONIES

Herring Gulls nesting on rooftops in South Shields and Sunderland were observed during 1974–76. These colonies, then around 200 pairs in each town, have expanded rapidly despite culls. Ringing results identified the Forth area as an important source of recruits, and also suggested that females are more likely to breed away from their natal area than males. It is clear from the high breeding success that these towns provide favourable alternatives to more typical breeding sites. The gulls in the towns nest at low densities, other than on flat roofs. Their breeding success, averaging 1.2–1.6 chicks per pair, was higher than that recorded at more typical colonies, and on structurally isolated rooftop sites was almost double that on flat roofs. This high success is attributed to the absence of cannibalism in these small colonies and the lack of territorial aggression at the structurally isolated sites.     

Corticosterone facilitates begging and affects resource allocation in the black-legged kittiwake

Parent black-legged kittiwakes (Rissa tridactyla) and their dependent chicks respond to food shortages by increasing circulatinglevels of corticosterone. To examine the behavioral significanceof corticosterone release, we experimentally increased levelsof circulating corticosterone in parents and chicks up to thelevels observed during food shortages. We found that corticosterone-implantedchicks begged more frequently than sham-implanted controls.Corticosterone-implanted chicks in broods of two begged morefrequently than singletons. Parent kittiwakes then respondedto the increase in corticosterone levels in their chicks byincreasing chick-feeding rates. However, feeding rates werenot different among corticosterone-implanted chicks in broodsof two and singletons. We also found that corticosterone-implantedparents spent more time away from the nest—perhaps foraging—andless time brooding/guarding chicks than sham-implanted controls.Untreated mates of the corticosterone-implanted bird did notcompensate for the change in their partner's behavior; consequently, chicks were left unattended about 20% of the time compared to1% at the control nests. However, corticosterone-implantedparents did not decrease their chick-feeding rates. Our findingssuggest two functional implications of the increased corticosteronesecretion during food shortages in the black-legged kittiwake:it facilitates begging in chicks, and it affects time allocatedby parents to guarding young at the nest. Thus, release of corticosterone might provide a mechanistic link between physiological condition and behavioral interactions among adults and theiryoung.     

Brood‐guarding duration in black‐browed albatrosses Thalassarche melanophris: temporal,geographical and individual variation

In birds, the period spent brooding or guarding young chicks is highly variable, but such variation has seldom been studied. Previous single‐year studies of Antarctic petrels Thalassoica antarctica and grey‐headed albatrosses Thalassarche chrysostoma revealed a pronounced seasonal decline in brood‐guarding duration and gave rise to the ‘synchronisation hypothesis’, which suggests that some of the variation in the length of the brood‐guarding stage is related to predictable seasonal changes in the risk of chick predation. We tested the predictions of this and three other hypotheses in a two‐site, four‐year study of the black‐browed albatross T. melanophris. The existence of a pronounced seasonal decline in brood‐guarding duration was apparent at both sites, and in years of contrasting food availability, providing further support for the ‘synchronisation hypothesis’. Alternative explanations for this pattern are that short brood‐guarding periods for late‐hatched chicks result from a seasonal decline in food availability or from the fact that early nesting birds are of higher individual quality. However, these explanations are at odds with the absence of a seasonal decline in early chick growth or in probability of chick survival. Furthermore, adult quality (measured as past reproductive performance) had a weak and inconsistent effect on the duration of brood‐guarding. Weather changes explained some of the variation in brood‐guarding, but there were no differences between regions of contrasting climates. Individual pairs displayed a degree of inter‐annual consistency in brood‐guarding duration and, at least in some years, longer brood‐guarding resulted in higher fledging probability. We speculate that a higher investment in brood‐guarding increases the cost of reproduction, which counteracts other selective pressures that would otherwise lead to longer brood‐guarding durations.