Melanophore death and disappearance produces color metamorphosis in the polychromatic Midas cichlid (Cichlasoma citrinellum)

Summary We describe the histological basis of color metamorphosis in the polychromatic Midas cichlid, Cichlasoma citrinellum. Eight percent of the individuals in a natural population transform from gray with black markings to orange, simultaneously losing their ability to adjust coloration in response to background and social context. This trait is inherited. Light- and electron microscopy revealed that this transformation is a two-step process. First, the melanophores die, then macrophage-like cells remove the debris. As a result of this initial process, the underlying xanthophores become visible, producing the orange coloration. A similar process may occur in individuals that further transform to white, or go directly from gray to white.     

Parental Investment and Division of Labor in the Midas Cichlid (Cichlasoma citrinellum)

The Midas cichlid (Cichlasoma citrinellum) is an aggressive, monogamous fish living in the Great Lakes of Nicaragua. Its breeding success rates are low due to intense competition for breeding sites and high levels of predation on the young. Male Midas cichlids devote a small portion of body weight to gonads and gametes compared with females. Males produce relatively small amounts of sperm perhaps because cichlid fertilization is very efficient and the male has a high certainty of paternity. While pair members invest equal amounts of time in parental care over the course of the breeding cycle, there is a clear division of labor. Males invest more in territorial protection than do females, whereas females provide more nurturance. Both parents are active over the course of the cycle but the male invests more intensely in the early stages of the cycle. About the time the eggs hatch, the burden of care shifts to the female and she continues to invest significantly more than the male over the remainder of the cycle. Females invest much more than do males when tissue investment and parental behavior are combined. Females also assume more of the burden of care as the brood matures. The male is not free to take advantage of this and leave his mate to initiate other broods because two parents are needed to defend the breeding site and brood. Monogamy with biparental care results.     

Mate size and breeding success in a monogamous cichlid fish

Synopsis The patterns of mate size and parental care of a monogamous cichlid fish,Cichlasoma maculicauda, were studied in Gatun Lake, Panama. Males defend territories which serve as courtship and nest sites. Within a population most mates in pairs are of equal size rank. In each pair the male is larger than the female, probably because most mature males are larger than most mature females. Clutch size increases with female body size. Male size affects breeding success in two ways. First, larger males provide nest sites less susceptible to destructive wave action. Second, young of larger males grow faster than young of smaller males. Large males defeat small males in contests for position in feeding areas, and this may provide their young with better feeding conditions. In the laboratory young growth rates increase with food abundance, and at high levels of food surpass those observed in nature. Fast growth of young reduces their vulnerability to predators and should allow parents to breed more often. Young survival rates improve with the size of the parents, so that larger fish raise more offspring at each breeding attempt. These observations suggest why preference for large mates should occur.     

Female Choice Predicts the Best Father in a Biparental Fish,the Midas Cichlid (Cichlasoma citrinellum)

The Midas cichlid is a monogamous, biparental species. It breeds in a highly competitive system where pairs have a low probability of raising fry to independence. Both parents must cooperate to retain the territory and protect the fry from predation. Previous experiments showed that females prefer large, aggressive and sexually experienced males as mates but males do not display any consistent preferences. Here I present the results of two experiments designed to see whether qualities preferred by females correlate with increased success in retaining territories and in providing parental care. Pairs with either large or aggressive males had an advantage in appropriating and holding a breeding territory; reproductive experience conferred no advantage in usurping a territory. Aggressive and reproductively experienced males had an advantage in defending the brood from predators of fry, but size had no effect. Thus, the qualities preferred by females confer advantages both in holding territories and in protecting fry. In contrast, males need not be selective because females, once in possession of a brood, defend it equally well regardless of size, aggressiveness, or reproductive experience. The system is one of mutually enforced monogamy based on female choice; females drive the system because they provide more investment than do males (combining gametes and time) and because this investment is a reliable resource to the male.     

Mate choice and size assortative pairing by the cichlid fishes of Lake Jiloá, Nicaragua

Size-assortative mating occurs for all six substratum-spawning cichlids studied in Lake Jiloá, Nicaragua. Mate quality is clearly a dominant consideration for both sexes, as larger pairs had disproportionately more young surviving beyond 3 weeks. Mating with the largest available mate should generally be advantageous to both sexes, since the success of a pair is dependent on their ability to take and defend a territory. A model involving mate discrimination by both partners is developed and contrasted to models involving single sex discrimination with intrasexual competition.     

Morphological variability in the Cuatro Cienegas cichlid, Cichlasoma minckleyi

The endemic cichlids of the Cuatro Cienegas Basin, Coahuila, Mexico are currently grouped in a single polymorphic species, Cichlasoma minckleyi . Two morphs of C. minckleyi were distinguished largely by features of the trophic apparatus, especially the pharyngeal dentition. Variation in body shape, based upon analysis of a set of linear measures, was continuous and did not allow recognition of discrete morphs. Individuals raised in the laboratory on several different diets indicated that trophic morphology had an important genetic component. Individuals raised in the laboratory, however, did not differentiate to the degree seen in comparably sized individuals collected at Cuatro Cienegas. This may be because snails used as food in the experiments were not as hard as endemic snails and indicated that some aspects of trophic morphology were also dependent upon environmental cues.     

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.     

Mate familiarity and social learning in a monogamous lizard

Oecologia - Social learning is thought to be advantageous as it allows an animal to gather information quickly without engaging in costly trial-and-error learning. However, animals should be...     

Inbreeding depression in the convict cichlid, Cichlasoma nigrofasciatum (Baird and Girard)

Convict cichlids, Cichlasoma nigrofasciatum , inbred for four and five generations exhibited moderate and severe morphological deformities. Deformed fishes were characterized by a more vertically sloping forehead, irregular jaw, hyoid, opercula and fin structure, reduced growth and irregular swimming behaviour. Inbred fish (F₄, F₅) did not differ from F₁s in their age of maturation or the average size of their first broods. Brood survival was 18.7% in F₅ inbred fish, compared to 75.5% in F₁s.     

Mate fidelity and breeding site tenacity in a monogamous sandpiper, the black turnstone

We examined the relationship between mate fidelity and breeding site tenacity during a 5-year study of the black turnstone, Arenaria melanocephala, a socially monogamous sandpiper breeding in subArctic Alaska. We tested the predictions of several hypotheses regarding the incidence of divorce and the benefits of fidelity to mate and breeding site. Interannual return rates to the breeding grounds (88% for males, 79% for females) were among the highest yet recorded for any scolopacid sandpiper, and 88% of returning birds nested on their previous year's territory. The annual divorce rate was only 11%, and mate fidelity was significantly linked to fidelity to territory but independent of sex and year. Males arrived in spring significantly earlier than their mates and interannual fidelity was influenced by the relative timing of arrival of pair members. Reunited pairs had significantly higher fledging success than new pairs formed after death or divorce. The incidence of divorce was unrelated to reproductive success the previous year, although birds nested significantly further away after failure than after a successful nesting attempt. Sightings of marked individuals suggested that members of pairs do not winter together, and breeding site tenacity provides a mechanism through which pair members can reunite. We reject the 'incompatibility' hypothesis for divorce in turnstones, and our data contradict predictions of the 'better option' hypothesis. Alternatively, we propose the 'bet-hedging' hypothesis to explain the occurrence of divorce, which transpires when an individual pairs with a new mate to avoid the cost of waiting for a previous mate to return. Such costs can include remaining unmated, if the former mate has died, or experiencing lower reproductive success because of delayed breeding. Copyright 2000 The Association for the Study of Animal Behaviour.     

Ecological and evolutionary consequences of the trophic polymorphism in Cichlasoma citrinellum (Pisces: Cichlidae)

The neotropical cichlid fish Cichlasoma citrinellum is polymorphic in the structure of its pharyngeal jaw apparatus and external morphology. The pharyngeal jaws are either gracile and bear slender, pointed teeth (papilliform) or robust with strong, rounded teeth (molariform). Molariform morphs have a ‘benthic’, and papilliform morphs a ‘limnetic’ body form. Furthermore, this species is also polychromatic, with yellow and black morphs. The molariform morphology of the pharyngeal jaw apparatus adapts the fish for cracking and feeding on snails. Based on analysis of stomach contents, 94% of the molariform morph ate snails whereas only 19%, of the papilliform morph did so. This result suggests that the morphs occupy different ecological niches. The morphology of the pharyngeal jaw apparatus does not correlate significantly with sex, but it does with body colouration (P<0.005). Cichlasoma citrinellum mate assortatively with their own colour; therefore a mating preference for colour may lead to genetic isolation of trophic morphs. The frequency of the molariform morph differs strikingly among populations of five Nicaraguan lakes and its abundance is correlated with the abundance of snails, the fishes' principal prey item. Among populations the frequency of molariform morphs decreases in the dry season. Morphology possibly changes reversibly within particular individuals between seasons. These results suggest that phenotypic plasticity and polymorphisms may be an adaptive characteristic of cichlid fishes. Patterns of intraspecific morphological variation match patterns of interspecific morphological diversification which suggests that universal developmental mechanisms canalize the possible expressions of morphology. The ability to respond morphologically to environmental shifts, in conjunction with genetically determined trophic polymorphisms and sexual selection via mate choice, could be the basis for speciation through intermediate stages of polymorphism of the impressive adaptive radiation of cichlid fishes.     

Mate choice, monogamy and mutational load

The classical Haldane mutational load for dominant mutations is 1 - e-alpha (where alpha = the mutation rate of the genome). This is independent of the fitness of the mutant heterozygotes. As alpha may be between 0.9 and 1.8 this may be too high for low fecundity species to bear. The load may be greatly reduced for mildly deleterious mutations if female choice (for males with few or no mutations) is present in a polygamous species. It is shown here that if females and males prefer partners with few or no mutations then the load may also be reduced in monogamous species. As in the polygamous model the effect is greatest if mutations are mildly deleterious.     

Isotocin regulates paternal care in a monogamous cichlid fish

While the survival value of paternal care is well understood, little is known about its physiological basis. Here we investigate the neuroendocrine contributions to paternal care in the monogamous cichlid, Amatitlania nigrofasciata. We first explored the dynamic range of paternal care in three experimental groups: biparental males (control fathers housed with their mate), single fathers (mate removed), or lone males (mate and offspring removed). We found that control males gradually increase paternal care over time, whereas single fathers increased care immediately after mate removal. Males with offspring present had lower levels of circulating 11-ketotestosterone (11-KT) yet still maintained aggressive displays toward brood predators. To determine what brain regions may contribute to paternal care, we quantified induction of the immediate early gene c-Fos, and found that single fathers have more c-Fos induction in the forebrain area Vv (putative lateral septum homologue), but not in the central pallium (area Dc). While overall preoptic area c-Fos induction was similar between groups, we found that parvocellular preoptic isotocin (IST) neurons in single fathers showed increased c-Fos induction, suggesting IST may facilitate the increase of paternal care after mate removal. To functionally test the role of IST in regulating paternal care, we treated biparental males with an IST receptor antagonist, which blocked paternal care. Our results indicate that isotocin plays a significant role in promoting paternal care, and more broadly suggest that the convergent evolution of paternal care across vertebrates may have recruited similar neuroendocrine mechanisms.     

Mate fidelity in monogamous birds: a re-examination of the Procellariiformes

Mate retention is classically considered advantageous for reproduction in monogamous birds: because of their low fecundity, long-lived species should show the highest year-to-year mate fidelity. However, this hypothesis remains controversial: several studies have found no correlation between mate fidelity and longevity, possibly because they did not control for potential confounding factors on each of these parameters, and one study found a negative correlation in the Procellariiformes (albatrosses and petrels). We re-examined the relations between mate fidelity and longevity, and between mate fidelity and site fidelity, in this group, using our data on 13 species and data from the literature, and after eliminating confounding factors. Procellariiformes are the most long lived of birds despite important interspecific variation in body size, and they show strong mate fidelity and bear high costs of divorce. All species lay only one egg, and the most long lived breed biennially. Because large organisms live longer than small ones and their reproductive effort is lower, we had to control for breeding frequency and body size. Mate fidelity and adult life expectancy were positively correlated, regardless of whether we controlled for these two parameters. We also evaluated whether mate fidelity was related to site fidelity. Biennial albatrosses show high mate fidelity, but low nest fidelity, although they are extremely faithful to a small area around their previous nest. After controlling for body size, adult life expectancy and breeding frequency, we found no correlation between mate fidelity and site fidelity. Because divorce is costly and mate retention advantageous in Procellariiformes, we suggest that mate fidelity does not passively result from site fidelity in these species. Rather, site fidelity would be a means for pairs to reunite, with sites serving as meeting points. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Effects of food supply on the reproduction of the convict cichlid, Cichlasoma nigrofasciatum

Cichlasoma nigrofasciatum , the convict cichlid, is a biparental, substrate-spawning, Central American cichlid. Male-female pairs were kept for 4 months on one of three food rations: high–1 g food daily; medium–1 g three times per week; low–1 g per week. The number of spawnings was directly related to the food ration. The fecundity of the females on the low and medium rations declined. At all rations the inter-spawning interval and egg size increased during the experiment. Egg size was positively related to ration and the length of the inter-spawning interval, but the latter was inversely related to ration. Females on the high ration grew in weight while those on the low ration lost weight. Histological examination of the ovaries at the end of the experiment suggested that low rations reduced the proportion of oocytes that become vitellogenic. At the end of the experiment, the total, gonad and liver weight of females at a given length were significantly related to ration. No similar, significant effect was found for the males, although growth in length of the males was related to ration. Ration had no detectable effect on the histological appearance of the testes.     

Mate choice in modern societies

Most research on mate choice in modern societies is based on data that may or may not reflect actual mating behavior (e.g., stated preferences, personal advertisements). In the present study, real-life matings were reported by a large representative sample of men and women (N = 1,133). These data were used to test an evolutionary model in which mate choice is hypothesized to depend on resources potentially contributed to reproduction by each sex. Consistent with the model, it was found that (a) men (but not women) of higher social status acquire more mating partners, suggesting that male status is an important criterion in female choice; (b) women’s (but not men’s) number of partners decreases linearly with age, suggesting that female reproductive potential is an important criterion in male choice; and (c) women (but not men) display a significant relationship between marital dissolution and promiscuity, suggesting that female sexual exclusivity is an important criterion in male choice. These results are discussed in relation to understanding mate choice mechanisms from behavioral data. Daniel Pérusse is an assistant professor in the Department of Anthropology at the Université de Montréal. His research interests include the evolutionary biology of human social and reproductive behavior, sexual selection theory, and biocultural evolution. His current research bears on human socialization processes and psychosocial development from an evolutionary and behavior-genetic perspective. Recent publications include “Cultural and Reproductive Success in Industrial Societies: Testing the Relationship at the Proximate and Ultimate Levels” (Behavioral and Brain Sciences 16(2):267–322, 1993) and “Human Parental Behavior: Evidence for Genetic Influence and Potential Implications for Gene-Culture Theory” with M. C. Neale, A. Heath, and L. J. Eaves (Behavior Genetics, in press).     

Mate choice in Darwin's Finches

Female Geospiza conirostris on Isla Genovesa, Galapagos, pair preferentially with males who have had previous breeding experience. They choose mates on the basis of courtship behaviour and black adult plumage. By mating with experienced black males, they gain a fitness advantage in terms of fledgling production and recruitment of young into the breeding population. Behavioural signs of past breeding experience and black plumage are reliable age- and condition-dependent traits. We suggest that females use conspicuous black plumage to identify old males at a distance, then interactions through courtship to modify initial assessments. Females paired with inferior males may increase the genetic quality of their offspring by extra-pair copulations; results of heritability analysis of morphology are consistent with this suggestion. Females change mates at a frequency of 12–27% per breeding season. They re-pair with males who are generally old, experienced, and hold territories adjacent to the deserted male. Females that re-pair gain a benefit, whereas males who are deserted within a breeding season incur a cost of more than 50% of their future potential production for that season. We conclude that females in choosing males seek reliable indicators of potential parental care, and in addition they may seek indicators of genetic quality.