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Amatitlania siquia Schmitter-Soto, 2007. Convict Cichlid; Convicto, Mojarra Rayada, Chogorra, Picaculo, Congo, Burra, Carate, Sargento. Distribution: Global: Middle America; from eastern Honduras to central Costa Rica, Atlantic and Pacific drainages; Costa Rica: LN, RF, SJ, SC, Sa, To, Pa, Ma (Atlantic), Ni, Te, Be, Ba and Ta (Pacific), 0���1060 m.a.s.l., Sec, Pot. Occurrence and conservation status: Nat, DD. References: Bussing (1966: 237, as Cichlasoma nigrofasciatum (G��nther 1867), in part���missidentification; listed, information on distribution), Bussing (1987: 213, as C. nigrofasciatum, in part���missidentification; 1998: 342, as Archocentrus nigorfasciatus, in part���missidentification; brief description, including illustrations and an identification key, information on distribution, with a map, and ecology), Schmitter-Soto (2007: 53; detailed description, including illustrations and an identification key, information on distribution, with a map, and ecology) and Angulo et al. (2013: 1004; listed, including taxonomic information and distributional data)., Published as part of Angulo, Arturo, 2021, New records and range extensions to the Costa Rican freshwater fish fauna, with an updated checklist, pp. 1-72 in Zootaxa 5083 (1) on page 47, DOI: 10.11646/zootaxa.5083.1.1, http://zenodo.org/record/5800630

Positive differential allocation predicts that females will increase their parental investment when mated to high-quality males due to the benefits that these males provide to offspring. Differential allocation predicts trade-offs between current and future reproduction, but investment trade-offs can also occur within a breeding event for species that provide parental care. However, differential allocation studies that have examined investment in both gametic effort and parental care remain rare and have only been conducted in birds. Fish may have more conservative investment strategies due to the large effect that current reproductive investment has on future reproductive potential. We examined gametic effort and parental care in the convict cichlid, Amatitlania siquia, a biparental fish, in response to male size, a proxy for quality. We randomly mated females to either large or small males and measured fecundity and parental behaviour of both parents. Females mated to large males did not increase their fecundity, but did provide more parental care to offspring during the larval stage, which is when male desertion in natural populations is most likely. Male size did not predict the amount of paternal care provided to offspring during the larval stage. Female condition at the end of the breeding event was negatively associated with higher fecundity and level of maternal care provided. The observed pattern of maternal investment suggests that females limit costs to future reproduction by limiting their increased investment to only one component of their reproductive effort. The increase in maternal investment during the larval stage may also serve to stabilize the pair bond and ensure continued male care, which ultimately increases the reproductive success of both parents. These results highlight important taxonomic and parental care pattern differences and broaden our understanding of the factors that influence differential allocation in diverse taxa.

Resumen Registramos nueve especies de peces durante cuatro giras de muestreo (2016 y 2017) en el Río Grande de San Ramón (cuenca del río Tárcoles, 1 060msnm), tres de ellas por primera vez para esta altitud o cuenca. Previamente, Alfaro cultratus (N=95) solo se informaba hasta 300msnm en la vertiente caribeña y en la cuenca del río Tempisque; Amatitlaniasiquia (N=4) solo por debajo de 540msnm para la cuenca del río Tárcoles; y Amatitlaniaseptemfasciata (N=22) desde la vertiente atlántica y por debajo de 600msnm. Abstract “First record of three fish species in Río Grande de San Ramón, Costa Rica”. Four sampling trips (2016 and 2017) in Río Grande de San Ramón (Tárcoles River Basin, 1 060masl) produced nine species, three of them for the first time for this altitude or basin. Previously, Alfaro cultratus (N=95) was reported only up to 300masl at the Caribbean slope and in the Tempisque River basin; Amatitlania siquia (N=4) only under 540masl for the Tárcoles river basin; and Amatitlania septemfasciata (N=22) from the Atlantic slope and below 600masl.

Four sampling trips (2016 and 2017) in Río Grande de San Ramón (Tárcoles River Basin, 1 060masl) produced nine species, three of them for the first time for this altitude or basin. Previously, Alfaro cultratus (N=95) was reported only up to 300masl at the Caribbean slope and in the Tempisque River basin; Amatitlania siquia (N=4) only under 540masl for the Tárcoles river basin; and Amatitlania septemfasciata (N=22) from the Atlantic slope and below 600 masl. Registramos nueve especies de peces durante cuatro giras de muestreo (2016 y 2017) en el Río Grande de San Ramón (cuenca del río Tárcoles, 1 060msnm), tres de ellas por primera vez para esta altitud o cuenca. Previamente, Alfaro cultratus (N=95) solo se informaba hasta 300msnm en la vertiente caribeña y en la cuenca del río Tempisque; Amatitlania siquia (N=4) solo por debajo de 540msnm para la cuenca del río Tárcoles; y Amatitlania septemfasciata (N = 22) desde la vertiente atlántica y por debajo de 600msnm.

11 pages; International audience; In monogamous species that provide biparental care, partners with similar behavioural types generally have a better reproductive success than dissimilar ones. The pattern of assortative mating for behavioural type is thus often interpreted as resulting from a mate choice process. However, an alternative process is also possible when the partners become similar through postpairing adjustments (behavioural convergence). The disentanglement of these two nonexclusive mechanisms is an important evolutionary question. Only sexual selection based on behavioural similarity before pairing can explain the maintenance of the behavioural type variability. In a previous study, we reported the existence of behavioural convergence for aggressiveness between partners in convict cichlids Amatitlania siquia, a monogamous tropical fish. We tested herein the assumption of mate choice based on behavioural similarity for aggressiveness using either short-term binary choice or long-term group level pairing. Pairing was not based on behavioural similarity between potential partners. Choosing a behaviourally compatible partner is a complex task because it requires profiling the potential mates. Individuals would thus achieve higher fitness benefits from choosing a partner on more conspicuous criteria (such as size for instance) and then attempt to increase partner similarity by convergence after pairing. Sexual selection does not appear to drive the maintenance of aggressiveness type variability in convict cichlids.

We present evidence suggesting that some male convict cichlids (Amatitlania siquia) in Lake Xiloa, Nicaragua engage in alternative reproductive tactics (ARTs). These putative ART males were smaller than typical parental males, displayed coloration similar to breeding females, and possessed enlarged gonads. Gonadosomatic indices in these males were 16-fold higher than parental males and three-fold higher than females. This phenotype is consistent with the ‘sneaker’ or ‘satellite’ phenotype reported in other species. A survey of 282 fish in 2014 found three males (1% of all fish, 2% of males) with these characteristics. This finding is significant because convict cichlids are frequently studied in laboratory and field in the context of monogamy and biparental care.

9 pages; International audience; In monogamous species, partner compatibility is a key factor influencing pairing and reproductive success. In pairs with biparental care, studies have mostly focused on behavioral compatibility because it is likely to encourage the coordination of parental care within pairs, leading to a better reproductive success. Behavior modulation, throughout the breeding season and as a function of the social context, is under the regulatory feedback control of endocrine mechanisms. From this link, the idea of hormonal partner compatibility as a key component of pair cohesion and maintenance has recently emerged. Here, we investigated the link between partner behavioral assortment and their hormonal response to the pairing context. We formed assortative and disassortative pairs of convict cichlids based on their behavioral type (proactive or reactive) and took hormone and fitness measurements. Testosterone, 11-ketotestosterone, 17β-estradiol and cortisol levels were measured from fish-holding water before and after pair formation. We found no relationship between the behavioral type of individuals and their pre-pairing hormone levels. Only the level of cortisol was affected by the partner but independently of its behavioral type. Reproductive success was not affected by the level of hormonal similarity within pairs, but we found that the variation in 11-ketotestosterone similarity between the isolated context and the pairing context was related to spawning size, and the variation in cortisol similarity to the number of fry. Behavioral compatibility does not reflect hormonal compatibility in the convict cichlid, but the relationship between reproductive success and the flexibility of hormonal similarity between partners suggests hormonal adjustment within pairs in this species.

In humans, affective states are a key component in pair-bonding, particularly in the early stage of a relationship. Pairing with a high-quality partner elicits positive affective states which, in turn, validate and reinforce the mate choice. Affective states thus strongly affect pair stability and future reproductive success. We propose generalizing the link between affective states and pair-bonding to encompass other monogamous species exhibiting biparental care, chiefly where the reproductive success of the pair critically depends on the coordination between partners. The convict cichlid Amatitlania siquia is a monogamous fish species that forms long-lasting pairs with strong cooperation between parents for parental care. In this species, we showed that females paired with their non-preferred male had lower reproductive success than those paired with their preferred male. We then transposed the judgement bias paradigm, previously used in other animal species, to assess objectively affective states in fishes. Females that were assigned their non-preferred partner exhibited pessimistic bias, which indicates a negative affective state. By contrast, females that were assigned their preferred partner did not exhibit changes in their affective state. Our results highlight that the influence of pair-bonding on affective states is not human-specific and can also be observed in non-human species.

Many monogamous species demonstrate size-assortative mating patterns within natural populations. To better understand the role of intersexual selection in this process, we examined the effect of male preference for female body size in the convict cichlid (Amatitlania siquia). We provided males with a choice between females that differed in size, relative to each other and in relation to the focal male. Based on previous work, we expected males to prefer the largest available female mates across all treatments. Surprisingly, males spent more time near the smaller of two available females, but only when the other female was larger than the male. Additionally, males spent little time with either of two potential female mates when both females were larger than the male. We hypothesized that while males might prefer the largest of available females, female behavior might limit males from acting on this preference. To test this, males were force paired with a smaller or larger female. Pair formation only occurred when the female was smaller than the male, and females that were larger than their male counterparts showed significantly more aggression when compared to smaller females. Together, these data suggest that in the absence of intrasexual competition, male mate preference for large females in convict cichlids might be limited by female aggression.

Female convict cichlids (Amatitlania siquia) exhibit bright orange ventral coloration that males lack. The behavioral implications of this color are poorly understood, particularly in naturally occurring populations where female coloration could play a role in the expression of territorial nest-guarding behaviors. In this field experiment, monogamous breeding pairs of convict cichlids were presented with 3D printed model conspecific intruders of three body sizes (small, medium, and large) exhibiting three orange patch sizes (large, small, or none) to observe how territorial aggression varied as a function of intruder size and female coloration. Individuals occupying breeding pairs that were defending hatched offspring were significantly more aggressive toward intruders with small and large amounts of orange than toward models lacking orange, indicating that color is an important context-dependent elicitor of aggression in this species. Males were significantly more aggressive toward the intruder than females, and male aggression was strongly influenced by their size relative to the intruder. When males were smaller than the intruder, they performed significantly more aggressive acts than when they were the same size or larger than the intruder; this trend persisted across three putative populations in Lake Xiloa, Nicaragua. A potential explanation for these findings is that the orange color functions as a signal of individual quality or breeding readiness and that breeding pairs increase aggression to repel intruders that pose the greatest threat to pair bond and nest maintenance. One or both sexes of many animal species possess brightly colored features that might communicate information about overall heath or reproductive status. In convict cichlid fish, males and females establish pair bonds and jointly defend their nest and offspring. Single females exhibit striking orange coloration that males and breeding females lack. Our field-based experiment provided evidence that more colorful females, which likely pose a threat to pair bond stability and nest maintenance, incite more aggression from breeding pairs than drab females. Our study suggests that color provides salient information about, perhaps, female quality or readiness to breed in natural populations, and adds to a growing body of research that seeks to understand the varied roles that colorful ornaments play in animal communication.

Interspecific patterns of space use in mobile animals have been the subject of considerable study, with allometric slopes often varying between 0.63 and 1.36. Both the slopes and intercepts of these relationships have been typically related to the energetic requirements of the focal animal, in relation to the amount of energy available in the environment. However, other explanations for these patterns are possible, including the ability of the focal animal to move about and/or defend a home range. To provide insight into this issue, we investigated the allometry of patch defence in convict cichlids, Amatitlania siquia , in a laboratory setting while controlling for energetic requirements. By manipulating the distance between two patches of food (0, 20, 40, 60, 80 and 95 cm), we estimated the maximum diameter of an ephemeral territory for 15 dominant fish that differed in body size by almost 40-fold. As expected, dominant fish were less able to defend and monopolize the food as the distance between the patches increased. Furthermore, larger fish were able to defend larger territories than smaller fish. The allometric slope of 1.18 in our study was remarkably similar to the interspecific slopes of home range or territory size in the literature for a broad range of vertebrate taxa. While there are likely multiple causes for the interspecific patterns of space use, our results suggest that the mobility of the defender is just as likely an explanation as energetic requirements.

This study tested if convict cichlids (Amatitlania siquia) with prior predator experience could socially transfer their predator knowledge to their offspring. Prior to reproduction, pairs were assigned to either an experimental or control associative learning treatment, and given novel predator odour from a wolf cichlid (Parachromis dovii) paired with either alarm cue or water respectively. We hypothesized that upon re-exposure to the odour, experimental pairs would socially transfer their acquired predator knowledge to their naïve offspring, which in turn would exhibit antipredator behaviour. While fry from both treatment groups did not show a difference in shoal formation, there is some evidence to suggest a trend in the experimental group regarding parental ability to transfer information to offspring and influence shoal behaviour. When tested individually, experimental fry decreased their activity more than control fry in the presence of the predator odour. This study provides evidence of a unique form of predator learning between parents and offspring, suggesting that biparental care may also serve to provide offspring with information about the habitat into which they are born.

Successful territory defence is a prerequisite for reproduction across many taxa, and often highly sensitive to the actions of territorial neighbours. Nevertheless, to date, assessments of the significance of the behaviour of heterospecific neighbours have been infrequent and taxonomically restricted. In this field study, I examined the importance of both heterospecific and conspecific neighbours in a biparental fish, the convict cichlid, Amatitlania siquia. This was done by assessing the colonisation rates of vacant territories, the rates of aggression by the territory holders, and the overall rates of aggression towards intruders, in treatments that controlled the proximity of both neighbour types. Convict cichlid pairs colonised vacant nesting resources (territory locations) at similar rates independent of the proximity of heterospecific (moga, Hypsophrys nicaraguensis) or conspecific neighbours. However, a model of sympatric cichlid intruder was subjected to considerably higher overall levels of aggression when mogas were nearby. In contrast, the proximity of conspecifics did not have a significant effect on the overall aggression towards the intruder. These results suggest that previously demonstrated higher survival of convict cichlid broods in close proximity of mogas may be driven by aggression towards shared enemies. No conclusive evidence was found regarding whether mogas also influence convict cichlids’ investment into anti-intruder aggression: the results show a marginally non-significant trend, and a moderately large effect size, to the direction of a lower investment in mogas’, but not conspecifics’, proximity. More generally, heterospecific neighbours may provide protective benefits in a wider range of ecological settings than commonly considered. Electronic supplementary material The online version of this article (10.1007/s00442-019-04483-0) contains supplementary material, which is available to authorized users.

Resumen La laguna Hule es un cráter de origen volcánico que se ha inundado con el tiempo, dando como resultado una gran masa de agua, sin estudios previos sobre su comunidad de peces. Muestreamos durante cinco meses la laguna y una quebrada (Pata de Gallo) con redes, gancho y línea. Los especímenes capturados se identificaron in situ al nivel taxonómico posible más bajo. La laguna tenía especies que normalmente se encuentran cerca de la vegetación acuática: Amatitlania siquia, Amphilophus alfari, Astyanax aeneus, Cynodonichthys sp., Parachromis dovii, Phallichthys amates y Poecilia gillii (dominante: A. aeneus, P. gillii y P. amates). La quebrada estaba dominada por Brachyrhaphis olomina pero también tenía Rhamdia laticauda. Los índices de baja diversidad (1,48 en la laguna y 0,26 en la quebrada) son característicos de los ecosistemas lacustres. Las hembras fueron, como se esperaba, más grandes y abundantes en P. amates y B. olomina; mientras que en P. gilli, algunos machos eran más grandes. El área se ve afectada por la contaminación, la introducción de especies invasoras, la pesca incontrolada y el uso de equipo destructivo de pesca. Proponemos la demarcación del estado de protección del sitio y la organización de un grupo comunitario para proteger a esta comunidad de peces. Abstract Laguna Hule is a volcanic origin crater which has been flooded over time, resulting in a deep mass of water with no previous studies about its fish community. For five months we sampled it and a draining stream (Pata de Gallo) with nets, and hook and line. The captured specimens were identified in situ to the possible lowest taxonomic level. The lagoon mostly had species normally found near aquatic vegetation: Amatitlania siquia, Amphilophus alfari, Astyanax aeneus, Cynodonichthys sp., Parachromis dovii, Phallichthys amates and Poecilia gillii (dominant: A. aeneus, P. gillii and P. amates). The stream was dominated by Brachyrhaphis olomina but also had Rhamdia laticauda. The low diversity indices (1,48 in the lagoon and 0,26 in the stream) are characteristic of lacustrine ecosystems. Females were, as expected, larger and more abundant in P. amates and B. olomina; whereas in P. gilli, some males were larger. The area is affected by pollution, introduction of invasive species, uncontrolled fishing and use of destructive fishing gear. We propose demarcation of site protection status and organizing a community group to protect this fish community.

During their lives animals constantly need to make decisions that influence their survival and their reproductive success. The objective of this thesis was to evaluate the role of two variability sources in decision-making — personality and emotional states — in a sexual context in a monogamous fish, the convict cichlid. Our work about personality highlights its importance on pair compatibility and reproductive success. We also observe that reactive individuals adjusted their behaviour to their partner and thus increased their reproductive fitness. This result supports the hypothesis that reactive individuals are more flexible and thus compensate for their weak competitiveness.Personality traits are thus to be considered as co-existing alternative strategies. The maintenance of this inter-individual variability can be results from their equal success on the long run. Emotional states are source of intra-individual variability that has been largely underestimate. However they may represent an internal information used by individuals to make choice. We developed a protocol of cognitive judgment bias test in fish to evaluate their emotional states, and we observed a strong relationship with the presence/absence of the preferred partner. These results suggest that emotions is a key component of pair-bonding and then question how animals use them to make decision. Do they choose a partner based on their emotional states in an adaptive way? Through these different studies, we have shown that behavioural variability is not randomly distributed. On the contrary, it is linked to different strategies and it can influence reproductive success. It is thus crucial to take this parameter into account in order to understand the evolution of decision processes., Au cours de leur vie, les animaux doivent sans cesse prendre des décisions qui impactent leur survie et leur succès reproducteur. L’objectif de cette thèse était d’étudier le rôle de deux sources de variabilité dans la prise de décision — la personnalité et l’état émotionnel — en contexte sexuel chez un poisson monogame, le cichlidé zébré. Nos travaux sur la personnalité ont mis en évidence son importance sur la compatibilité comportementale et le succès reproducteur des partenaires. De plus, les individus les plus réactifs se sont ajustés à leur partenaire et ont augmenté leur succès reproducteur. Ceci supporte l’hypothèse d’une plus grande flexibilité de ces individus qui compenseraient ainsi leur faible compétitivité. Les différents profils de personnalité peuvent ainsi être considérés comme des stratégies alternatives. Le maintien de la variabilité interindividuelle pourrait ainsi s’expliquer par le succès équivalent de ces stratégies. Les émotions sont, elles, une cause de variabilité intra-individuelle souvent négligée qui pourrait pourtant représenter une information interne utilisée par l’individu pour prendre ses décisions. En transposant le test cognitif du biais de jugement aux poissons, nous avons pu observer un lien étroit entre leur état émotionnel et la présence/absence du partenaire. Ces résultats suggèrent un rôle des émotions dans les processus d’appariement et interrogent sur leur utilisation dans les prises de décision. Les individus se basent-ils sur leurs émotions pour choisir un partenaire ? Est-ce adaptatif ? À travers l’étude de ces sources de variabilité comportementale, nous avons observé que, loin d’être distribuée aléatoirement, la variabilité était liée à des stratégies décisionnelles et pouvait influencer le succès reproducteur des individus. Il serait donc crucial de la prendre en compte pour appréhender l’évolution des processus décisionnels.

Defense of a territory is a primary factor determining the fitness of territorial individuals. Thus, territorial animals use a variety of means to lower the cost of territorial defense. Although landmarks are known to lower the defense costs of territories, little is known about how they could affect fundamental properties of territories, which could ultimately affect the demography, population regulation, individual behavior, and spatial ecology of a territorial species. This field study examined the effects of natural landmarks on territory size, shape, and defensive costs in Amatitlania siquia, a little-studied member of the convict cichlid group native to Costa Rica and Nicaragua. We investigated the territorial behavior of 103 breeding pairs in a caldera lake in Nicaragua. During 10-min focal observations, we recorded the locations at which residents chased intruders and then used these locations to outline the territory boundaries. Landmarks affected both size and shape of A. siquia territories. Territory size decreased as the percentage of the boundary at landmarks increased. Territories with

Mobile young under parental care have a high potential for intermixing with other broods, which potentially increases the costs to the foster parents. Here, we examined for the first time the genetic composition of wild-caught broods of the convict cichlid (Amatitlania siquia), a socially monogamous biparental fish, for evidence of brood mixing and adoption. Our microsatellite genotyping data revealed that 79% of broods contained adopted young. Moreover, 25% of broods contained adopted sibsets likely arising from extra-pair matings, a phenomenon hitherto not documented for this species. Overall, adopted foreign fry and host fry in mixed broods were generally different in body length, as would be expected if they have different parents. However, fry from possible extra-pair matings were similar in body length to host fry, suggesting that they are of similar age. Our results are important because they reveal a very high prevalence and degree of brood mixing, and indicate that social monogamy does not necessarily lead to genetic monogamy in the convict cichlid in nature. These findings raise questions about potential brood-mixing mechanisms and the reproductive ecology (especially opportunities for polygamy in nature) of this important model species in the study of animal behaviour.

We examined ionoregulatory characteristics of two species of characiform fish and two species of cichlids that are not native to the Rio Negro, an extremely ion-poor, acidic tributary of the Amazon River, in order to gain insight into the origin of the specializations possessed by Rio Negro fish. The two characiform species examined, Congo tetras (Phenacogrammus interruptus) and black neon tetras (Hyphessobrycon herbertaxelrodi), had high-affinity/high-capacity transporters that produce high rates of uptake in dilute water. Na+ uptake for both was pH insensitive (down to pH 3.5). Exposure to 100 μmol L−1 phenamil had no effect on Na+ uptake in either species, while exposure to 1 mmol L−1 NH4Cl− (high external ammonia [HEA]) slightly stimulated Na+ uptake in Congo tetras. Exposure to “Na+-free” water significantly inhibited Na+ uptake (by 65%–85%) but had no effect on net ammonia flux. The two cichlid species examined, convict cichlids (Amatitlania siquia) and red point cichlids (Archocentrus sp.),...

The signaling function of carotenoid-based orange ventral coloration expressed by female convict cichlids (A. siquia) is still largely unknown. This field study examined the relationship between breeding female A. siquia orange coloration and predation stress, territorial competition, and offspring developmental stage. Female orange coloration decreased significantly with increased behavioral interactions with predators and heterospecific competitors, and the presence of wrigglers, fry, or juveniles was significantly associated with a lack of orange coloration. Multiple alternative explanations for these observations are discussed, including body carotenoid re-allocation and variation in environmental characteristics.

Predation exerts tremendous selection pressure on all organisms. In this study, we exposed embryos of convict cichlids (Amatitlania siquia) twice daily to one of the following: (1) chemical alarm cues of damaged conspecifics + odour of a novel predator (Polypterus endlicheri), (2) chemical alarm cues of damaged conspecifics + water or (3) blank water. No chemical cues were presented after the eggs hatched. When the larvae were 9 days old (mean total length = 5.7 mm), they were exposed to either predator odour or water. Those larvae that had been conditioned as embryos on alarm cues + predator odour showed a significant reduction in activity (i.e. anti-predator behavioural response) to predator odour relative to the other treatments. This is the first demonstration of acquired predator recognition by fish embryos.

The cues associated with social familiarity and genetic relatedness and how they interact to influence the formation of social associations among individuals, and thus group composition and dynamics, is poorly understood. Here, we investigated the concurrent effects of social familiarity and kinship on social affiliations in free-swimming convict cichlid fish young or ‘fry’ (Amatitlania siquia) by pitting the cues of social familiarity and kinship against each other in a four-way choice apparatus. Individual focal fish were given a simultaneous choice to associate (‘shoal’) with conspecifics that were either socially familiar and kin (full sibs), socially unfamiliar and kin, socially familiar and not kin, or socially unfamiliar and not kin. Stimulus shoal preference differed depending on the body length of the focal fish; smaller fry exhibited no preference, whereas larger (more mobile) fry significantly preferred to associate with familiar kin. In the convict cichlid system, where brood mixing occurs in the wild, a preference to associate with familiar kin may confer fitness benefits to individuals, especially when fry become more mobile as they grow and encounter predators more often. Our results contribute to further our understanding of the roles of familiarity and kinship in the formation of social associations in the convict cichlid in particular and in animals in general.

The differential allocation hypothesis predicts individuals will increase their reproductive investment when mated to a high quality partner. In many species of fish with biparental care females prefer large males due to the males’ greater ability to raise more offspring to independence. I examined the relationship between mate quality, parental care and number of offspring in a natural population of convict cichlids Amatitlania siquia. The frequency of frontal displays by females was positively correlated with male standard length. Additionally, as males increased in length relative to their mate, females increased the frequency of chases towards predators, while males decreased the number of displays towards brood predators. This trade-off in parental effort within a pair due to mate quality is a key prediction of differential allocation. The number of offspring was correlated with male, but not female, standard length. These results support the differential allocation hypothesis in that females offered more parental care to offspring of a larger male, while their mates decreased the amount of care they provided. Additionally, females benefited in terms of number of offspring by pairing with higher quality mates. Increased female investment may provide an incentive to ensure male care and maintain pair bonding, which could lead to greater reproductive success through increased offspring survival.

The neotropical convict cichlid fish (Amatitlania siquia) is an important model species for the study of animal behaviour. It is therefore surprising that no genetic markers are available for this species. Here, we have optimised four poly- morphic microsatellite loci for use with this fish, where each locus comprises between 9 and 28 alleles. We demonstrate how these markers can be used to genotype fry (young) and for the analyses of genetic structure and prevalence of interfamilial mixing within biparental broods of wild convict cichlids. These microsatellites will facilitate future research, hitherto not possible, on the molecular, behavioural and evolutionary ecology of the convict cichlid and possibly other closely related species.

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.

We studied parental care behaviour in a field population of convict cichlids, Amatitlania siquia, a fish with biparental care of its free-swimming young (fry). We simulated catastrophic brood loss by removing all of the young from 26 broods of various ages. We recorded parental defence behaviour for 5 min preremoval and three consecutive 5 min postremoval intervals. We then returned all of the young and recorded 5 min of parenteoffspring reunion behaviour. Following brood loss, both parents defended the empty territory and searched for missing young. Females spent more time in the empty territory and performed more attacks in defence of the empty territory than did males. Females searched and defended a significantly larger area than males did. During the 15 min fry-absent period, males increasingly spent more time away, whereas females maintained a constant level of investment in searching the brood area. Four of 26 males (15%) engaged in courtship with a new female within 15 min of brood loss. Attacks to repel potential predators from the brood area declined in frequency over the 15 min fry-absent period in both sexes. Standard length (age) of young was positively correlated with parental investment for females but not males. Number of young was positively correlated with parental investment for males but not for females. We conclude that post-brood-loss behaviours are search and recovery tactics that contribute to parental inclusive fitness in the event of incomplete brood predation.

We determine size, shape and location for a territory that is optimal in the sense of minimizing defense costs, when a given proportion of the boundary is landmarked and its primary benefit in terms of fitness is greater ease of detecting intruders across it. Increasing the landmarked proportion of boundary causes the optimal configuration to be smaller and more elongated, and to be located with its center further from the nest, so that the nest is closer to the landmarked boundary. These predictions accord with observations in a recent study of the convict cichlid Amatitlania siquia. Our results thus confirm the consistency of the observed behavior with the hypothesis that A. siquia designs its territory to make intruders easier to spot. Our results also lead us to conjecture that moving the landmark proportionately closer to or further away from the nest would have yielded essentially the same outcome in this study, because the optimal configuration depends only on the angle subtended by the landmark at the nest and hence only on the length of the landmark relative to its distance from the nest, as opposed to its absolute value.

9 pages; International audience; Interindividual behavioural differences, consistent over time and across situations, are frequently related to differences in reproductive success, susceptibility to disease and competitive ability. The behavioural phenotype of a conspecific is likely to affect the outcome of a social interaction, so an individual must be able to accurately assess behavioural phenotypes in order to optimize behavioural responses during mate choice, cooperative relationships and agonistic interactions. However, behavioural assessment based on dyadic interactions in different contexts can be costly because it may require significant time, increase the risk of predation and often necessitates special cognitive abilities. Different colour cues are associated with different life history strategies in many species, so coloration might also correlate with behavioural phenotype. Thus, conspecifics could use coloration as a single cue for the mediation of social interactions. We investigated the relationship between personality and coloration in the monogamous, biparental convict cichlid. This species has black melanin-based stripes with interindividual variation in the intensity of these stripes. In addition, females have bright orange spots on their abdomens. Using repeatable measures of boldness, exploration and aggressiveness, we were able to define a behavioural syndrome and quantified the behavioural type of each individual. Behavioural types were predicted by colour traits. In particular, proactive individuals (more aggressive, bolder and quicker to resume behaviour after being startled) were more likely to be dark and, among females, have smaller orange areas. Our results support the hypothesis that coloration in the convict cichlid is related to personality. Thus, coloration could play an important role in mediating social interactions, allowing a quick and reliable assessment of conspecifics' behavioural types.

9 pages; International audience; In recent years, considerable research interest in behavioral ecology has focused on characterizing and understanding individual differences in behavior that are consistent over time and across contexts, termed animal “personalities,” and correlations between various behaviors across contexts, termed behavioral syndromes. Although there is some evidence that differences in personality among individuals within populations can be genetically based and adaptive, when and how individual personality differences emerge in a population is not well understood, but of considerable general interest. Here, using juveniles of the convict cichlid (Amatitlania siquia) as a model system, we investigated in the laboratory whether individuals consistently differ in their personalities and whether behavioral syndromes are apparent at an early developmental stage and, if so, whether distinct personality traits are heritable. Under standardized laboratory conditions and using sibling analysis, we quantified interindividual differences in their boldness behavior under potential predation threat and their exploratory activity in a novel environment, 2 ecologically important behaviors, as our focal personality traits and estimated their respective repeatability and heritability. We report for the first time consistent (repeatable) and heritable individual differences in boldness and exploratory behaviors, and a boldness–exploration behavioral syndrome, in young convict cichlids. Bolder fish were more exploratory than relatively timid ones. These results provide novel evidence for the emergence in early life history of consistent individual differences in personality traits and behavioral syndromes in this species and suggest that genetic variation for boldness and exploratory behaviors, and thus potential for selection on these traits, exists in our study population.

In species with extended parental care, mobile dependent young are potentially more vulnerable to predators when they stray and become separated from their parents. We would expect that the likelihood of, and latency time for, a separated young to safely return to its 'family unit' (i.e. parents and brood mates) to be, respectively, inversely and positively related to the initial distance of separation and potentially mediated by its age or body size. Using the biparental convict cichlid fish (Amatitlania siquia), we tested these predictions by capturing individual young and displacing them at varying distances from their family unit in both the field and laboratory. As expected, displaced fish were less likely, and took longer, to return to their family with increasing separation distance from the family unit. The body length of displaced young mediated these relationships and their antipredator behaviour; larger young refuged more than smaller ones and were also less likely to be eaten by predators. These results suggest that selection should favour strong affiliative behaviour in mobile young animals towards their brood mates and protective parents because straying from the family unit leads to increased exposure to predation and a reduced likelihood of returning home with increasing separation distance.

Cichlids are unusual among fishes because they have prolonged care of their young. Convict cichlids,Cichlasoma nigrofasciatum, are monogamous and have biparental care of their young. This species has been studied extensively in the laboratory, however, little is known of their reproductive habits in nature. Four populations of convict cichlids were studied in Costa Rican streams during the long dry season in 1990 and 1991. Basic physical and chemical parameters of the study sites are described. In the study area breeding fish paired size assortatively; large males with large females and small males with small females. A comparison of the size distribution of breeding fish to that of non-breeding fish showed that small males and in some cases, small females were excluded from breeding. This suggests competition among males for nest sites and mates or female-female competition. Further evidence of intrasexual competition in females is provided by the black colour phase adopted by some females. Some of these females were active in courtship and had mature ova. Cave guarding behaviour and reuse of caves show that a ready-made spawning site is an important resource shaping the mating system of this species. This description of basic natural history can serve as a source of testable hypotheses for future research and allow the results from laboratory studies on this and related species to be interpreted within the ecological context of the natural habitat.

This study examines the effect of parent size, fry growth rate, and habitat-related brood predation pressure on two measures of reproductive success (percent brood survival and fry number at independence from their parents) of free-ranging convict cichlids (Cichlasoma nigrofasciatum). Four populations in Costa Rican streams were studied, two in "pool" habitat and two in "stream" habitat. The number of fry that emerged from the spawning site (as free-swimming young) was positively correlated with female standard length (SL) for sites with a relatively wide size range of breeding females. SL of males had no effect on fry number at emergence, nor did size of either parent affect the probability of a brood reaching fry independence. Large parents reared more fry to independence than small parents in stream habitat but not in pool habitat. There was a significant difference among sites in fry number at emergence but not at independence (mean = 27 fry), suggesting a maximum number of fry that two parents can economically defend. Fry growth rate had no effect on either measure of reproductive success. Reproductive success was largely determined by the effect of habitat. Adult Cichlasoma dovii in pool habitat may have reduced mean brood survival from 47% (n = 126 broods) in stream habitat to 14.9% (n = 141 broods) in pool habitat. Predators such as juvenile C. dovii may be more effective in the dim ambient light in pool habitat, overwhelming the effect of parent size on brood defence.

Amatitlania siquia, new species Figures 17, 21 Cichlasoma nigrofasciatum (part. et non G��nther), Bussing 1987: 213 (misidentification). Archocentrus nigrofasciatus (part. et non G��nther), Bussing 1998: 346 (misidentification). Holotype. UMMZ 245585, 61 mm SL (Fig. 21), M. J. Allen, Aug. 10, 1932. A stream flowing out of the forest into R��o Siquia, 7 miles N of Rama, Atlantic Nicaragua. Paratypes. ANSP 88241 (3), 67475 (4), 140692 (1), BMNH 1925.3.6.124-125 (2), FMNH 7733 (1), MHNG 2160.60 (1), UMMZ 145720 (2), 166474 (1), 196939 (1), 196948 (25), 196954 (7), 213938 (6). Diagnosis. Unique autapomorphies (figs. 24e and 6d in Schmitter-Soto, in press): gut simple (S-shaped), but folded ventrorostrally, anal loop and medial loop not touching (vs. simple, not folded); peritoneum only rostrally pigmented (vs. not only rostrally pigmented). Further distinguished from all other Amatitlania as follows: caudal blotch about two-thirds on peduncle and one-third on fin (vs. completely on fin); posterior edge of mesethmoid straight-irregular (vs. rounded); scales from lateral line to base of first dorsal fin ray modally 1.5 (vs. modally 2 or more); caudal vertebrae modally 14 (vs. modally 13). Description. D. XVII -XVIII,8-10; A. IX -X,7-8. Gill rakers bifid in larger specimens, sometimes serrated, or at least distally expanded, compressed. Scales from lateral line to base of first dorsal-fin ray always 1.5; circumpeduncular scales usually 15-17, modally 16; total vertebrae 26-27 (additional meristic data appear in Table 3). Largest specimen examined, 79 mm SL. A deep-bodied Amatitlania, body depth 48-55% of SL, usually greater than 49% of SL (further morphometric data appear in Table 4). Head profile straight to convex. Teeth conical, pointed, slightly or not labiolingually compressed. Upper symphysial teeth abruptly larger than adjacent teeth. Upper lip medially narrow; upper angle of lower lip at corner much rounded, lower angle acute, tapering ventrad. Pectoral fins always reaching caudad beyond 3rd, pelvic fins always beyond 5th anal-fin spine and often to 8th anal-fin spine. Filamentous rays of dorsal fin to distal third of caudal fin. Up to three pored scales continuing lateral line on caudal fin, isolated subsidiary scales on larger specimens. Dorsal- and anal-fin interradial scales arranged in one row, up to six scales long. Gut length may reach 188% of SL. Genital papilla oval, but often medially constricted, peanut-shaped, longer than wide, with a long protuberance on tip; isolated melanophores on base and sides. Suborbital streak present, at least in young; stripe from snout to eye usually well defined. Interorbital bands sometimes not so diffuse. Head bar more intense dorsally. Eyes bluish, greenish, greyish. Bars on side of body ventrally more obsolescent and coalescent than in other species; 4th bar I-shaped. Dots sometimes discernible on bases of soft dorsal and anal fins. Breast olive. Axil of pectoral fin with a dorsal dark spot; base of pectoral definitely white. Caudal blotch two-thirds on fin, more dorsal than ventral to lateral line, never to ventral edge, oval. Distribution. Both coasts of Costa Rica (from tributaries to the Golfo de Nicoya on the Pacific coast and R��o Parismina on the Atlantic) to Nicaragua (including the Great Lakes and the Mosquitian basins); also north to Atlantic Honduras (R��o Yeguar��) (Fig. 17). Etymology. Siquia is the name of the river chosen as type locality. The name means ���avocado��� in the Miskito dialect Ulwa (Salamanca 2002). A noun in apposition., Published as part of Juan J. Schmitter-Soto, 2007, A systematic revision of the genus Archocentrus (Perciformes: Cichlidae), with the description of two new genera and six new species., pp. 1-78 in Zootaxa 1603 on pages 54-56

Am. siquia: UMMZ 245585 (holotype), UMMZ 196948 (25 paratypes, 2 C&S, 16 dig.), UMMZ 196954 (7 paratypes), Nicaragua, R��o Siquia, 7 mi above Rama; ANSP 88241 (3 paratypes), Costa Rica, ���Esperata��� [Esparta]; ANSP 67475 (4 paratypes), Nicaragua, ���great falls��� on R��o Pis Pis; ANSP 140692 (1 paratype), Costa Rica, Puntarenas, R��o Ahogados; FMNH 7733 (1 paratype), Costa Rica, Turrubares; UMMZ 145720 (2 paratypes, 2 dig.), Costa Rica, Talamanca, R��o Bioliri, tributary to R��o Lari; UMMZ 166474 (1 paratype), Costa Rica, Laguna del Misterio; UMMZ 196939 (1 paratype, dig.), Nicaragua, creek tributary of R��o Siquia, 7 mi upstream from Rama; BMNH 1925.3.6.124-125 (2 paratypes), Costa Rica, Talamanca, R��o Bioliri, tributary to R��o Lari; MHNG 2160.60 (1 paratype), Nicaragua, Lake Managua at Momotombo; UMMZ 213938 (6 paratypes), Nicaragua, Rivas, R��o Ca��as Gordas; ANSP 93215 (2), Costa Rica, ���Esperata��� [Esparta]; ANSP 168328 (57), Costa Rica, Alajuela, Lake and R��o Cuarto; BMNH 1929.8.9.20-29 (10), Costa Rica, Liberia; BMNH 1968.1.12.2-4 (3), Nicaragua; BMNH 1968.1.12.32-33 (2), Costa Rica, ���Squirres��� [Siquirres]; UMMZ 165774 (1), Nicaragua, Matagalpa; UMMZ 166472 (2), Costa Rica, Laguna del Misterio; UMMZ 180616 (1), Nicaragua, Lago Managua at Matear��; UMMZ 188120 (3), Honduras, El Para��so, R��o Cato in Valle de Jamastr��n; UMMZ 188144 (6, 4 dig.), Honduras, Moraz��n, Lagunita S of road Tegucigalpa-Danl��; UMMZ 188257 (7), UMMZ 188258 (1), Nicaragua, Zelaya, R��o Huahuash��n at Corozo Camp; UMMZ 188293 (4), Honduras, Moraz��n, R��o Yeguar�� at Tegucigalpa; UMMZ 190191 (18, 2 C&S, 14 dig.), Costa Rica, Guanacaste, 5 km NW of Las Ca��as, R��o Corobic��; UMMZ 190199 (16, 6 dig.), Costa Rica, Guanacaste, R��o Piedras, 12 mi SE of Liberia; UMMZ 196939 (1, dig.), UMMZ 199627 (129), Nicaragua, R��o Kurnog, tributary of Laguna P��hara [Pahara?], SE of Bilwaskarma; UMMZ 199637 (86, 6 dig.), Nicaragua, 6 km E of Waspam, R��o Putkrukira, tributary of R��o Coco; UMMZ 199650 (51, 6 dig.), Nicaragua, 35 km S of Waspam, R��o Likus, tributary to R��o Wawa; UMMZ 199661 (35), Nicaragua, R��o Coco, 300 m upstream of Waspam; UMMZ uncat. (22), Costa Rica, Guanacaste, Tempisque, 2 mi S of Hacienda Tenorio., Published as part of Juan J. Schmitter-Soto, 2007, A systematic revision of the genus Archocentrus (Perciformes: Cichlidae), with the description of two new genera and six new species., pp. 1-78 in Zootaxa 1603 on page 72

Few studies have examined how landmarks affect territories' fundamental characteristics. In this field study, we investigated effects of landmarks on territory size, shape and location in a cichlid fish ( Amatitlania siquia ). We provided cans as breeding sites and used plastic plants as landmarks. During 10 min trials, we recorded locations where residents chased intruders and used those locations to outline and measure the territory. In two experiments, we observed pairs without landmarks and with either a point landmark (one plant) or linear landmark (four plants) placed near the nest can. We alternated which trial occurred first and performed the second trial 24 h after the first. Territories were approximately round without landmarks or with a point landmark but were significantly more elongated when we added a linear landmark. Without landmarks, nests were centrally located; however, with any landmark, pairs set territory boundaries closer to the landmark and thus the nest. Territory size was significantly reduced in the presence of any landmark. This reduction suggests that a smaller territory with well-defined boundaries has greater benefits than a larger territory with less well-defined borders.

established wild populations in the USA (Hawaii), Mexico, Israel, Japan and Australia (Ishikawa and Tachihara 2010; Welcomme 1988). The species is a member of the Cichlidae, a family of fi sh known for prolonged biparental care of offspring (Bernstein 1980; Keenleyside 1991). Due to confusion over taxonomy, the various studies of convict cichlids may not all represent the same species. However, all species are closely related, and therefore the results of behavioural studies are broadly applicable across the group. The convict cichlid group has been extensively studied in the laboratory (Wisenden 1995) but despite signifi cant research, few studies have investigated the life history of Amatitlania nigrofasciata in natural environments. Adults were shown to reach 100 mm standard length (SL) (Kullander 2003) and mature females were found at 32 mm SL in introduced populations in Japan (Ishikawa and Tachihara 2010). This size is similar to that reported for the closely related Amatitlania siquia1, whose males mature at sizes as small as 51 mm SL, and is a species which has received more intensive study (Wisenden 1995). Parental pairs are monogamous, and in the introduced population in Japan, bred throughout the year despite water temperatures of less than 20oC (Ishikawa and Tachihara 2010). Amatitlania siquia excavates small caves under hard substrate and lays its eggs on the roof of these caves (Wisenden 1995). Both sexes care for the young (Noonan 1983) with adult fi sh caring for the brood by shepherding juveniles when outside the cave (Wisenden et al. 1995) and returning them to the cave at night for shelter, where adults stand guard and defend against potential predators (Gagliardi-Seeley and Itzkowitz 2009; Wisenden 1995). The non-native population in Japan exhibits rapid growth in the fi rst The convict cichlid Amatitlania nigrofasciata (Cichlidae): fi rst record of this non-native species in Western Australian waterbodies

The temporal pattern of breeding, the bathymetric distribution of breeding pairs, the microhabitat selection of breeding sites and the sizes of breeding individuals of nine species of cichlid and one eleotrid predator, Gobiomorus dormitor, were examined in Lake Jilod, Nicaragua. For two relatively short periods of the year, during the middle of both wet and dry seasons, competition for breeding sites between these fishes was intense. Over 90% of all territories were lost prior to complet- ing a breeding cycle. Cichlid pairs in shallow water tended to be more successful and breeding sites in these areas were more vigorously contested. The seasonal pattern of breeding was bimodal with a different species complex breeding in the wet season as contrasted to the dry. In the wet season Cichlasoma citrinellum aggressively excluded all but the two smallest species from breeding in rocky areas. In the dry season, when C. citrinellum was not breeding, all other species were able to breed, but they, in turn, were separated within that season by both depth and timing of breeding.