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.     

Cichlid Fishes from the La Plata Basin. Part IV. Review of the Apistogramma Species, with Description of a New Species (Teleostei, Cichlidae)

Apistogramma inconspicua sp.n. is described from the Rio Candclaria in the Rio Guaporé drainage in Bolivia, and recorded also from Cáceres on the Rio Paraguay in Brazil. The new species is closely related to the Paraguayan species A. commbrae (Regan) and an undescribed species in the Guaporé drainage system. These species share a specialized feature in the colour pattern, viz. an expanded vertical bar on the caudal peduncle which it confluent with the caudal spot. The now presumably complete record of Paraguayan Apistogramma species allows an analytical key to be given. Of the additional species, A. borellii (Regan) and A. trifasciata (Eigenmann & Kennedy) represent two distinct lineages. Apistogramma pleurotaenia (Regan) is probably a La Plata basin species, although no localities are known, and its relationship remain obscure. The assemblage of Paraguayan Apistogramma species reflects the heterogeneity of the Paraguayan cichlid fauna as a whole.     

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.     

Differences in Aggressiveness in the Midas Cichlid Fish (Cichlasoma citrinellum) in Relation to Sex,Reproductive State and the Individual

The Midas cichlid, Cichlasoma citrinellum, is monogamous and biparental. Because of competition for limited spawning sites and intense predation on their young, vigorous defense of their territory is essential. Although both sexes engage in defense, they differ in aggressiveness. The aggressive responses of both sexes were measured by counting the number of bites and bumps each fish directed toward its own mirror image. The size of the fish's genital papilla was also recorded to estimate its reproductive state. Compared with females, males had higher median mirror scores with greater variance. The scores of individual males were also more consistent through time than were those of females. Females close to spawning had the highest mirror scores, whereas male scores were highest early in the reproductive cycle. Selection has apparently favored aggressiveness in both sexes. We argue, however, that differences in aggression are the result of selection acting dissimilarly on the two sexes.     

Species diversity and phylogeography of the Australoheros autrani group (Teleostei,Cichlidae) in eastern Brazil

Cichlid fishes are an important model system in evolutionary biology, primarily because of their exceptional diversity. However, while some cichlid assemblages, such as the ones of the African Great Lakes, have received considerable attention, others are not well studied, including many riverine species. Here, we focus on the Australoheros autrani species group and first report a new record of Australoheros in the upper Paranaíba River drainage, extending the known distribution range of this genus. Through Bayesian inference and maximum likelihood phylogenetic analyses of the mitochondrial cytochrome b gene of these specimens as well as available sequences, we assigned the newly discovered population to Australoheros barbosae. We corroborate the monophyly of the A. autrani species group and the presence of three species in the upper/middle Paraíba do Sul River basin as well as molecular diagnostic characters for each. Finally, we provide evidence for a recent expansion of A. barbosae.

     

Cichlid Fishes from the La Plata Basin. Part V. Description of Aequidens plagiozonatus sp.n. (Teleostei, Cichlidae) from the Paraguay River System

Aequidens plagiozonatus sp.n. is described from 49 specimens collected in the northern Paraguay river system. It is the only known species of Aequidens Eigenmann & Bray in the La Plata basin. Distinguishing characteristics include the slanting vertical bars on the sides, discontinuous horizontal lateral band, immaculate or very sparsely dotted caudal fin and short pectoral fin.     

Comparative ontogenesis and cytomorphology of the nasal organs in some species of cichlid fish (Cichlidae, Teleostei)

This study compares the ontogenesis, morphology and cytology of olfactory organs of several species of Cichlidae (Pisces, Teleostei), from Asia, Africa and America, belonging to the subfamilies Tilapinae and Haplochrominae. Cichlids, in contrast to most other fish families, possess only one naris on each side of the nose that serves as both inlet and outlet opening. The olfactory rosettes are round, 1.8–4.2 mm diameter, and situated directly below the nares. Each rosette consists of a central raphe with attached lamellae whose number increases with age, remains constant in adults and differs in the various species. Onset of organization of the nasal organs from the neuroectoderm begins 14–16 hours after fertilization in bottomspawner cichlids, and 30–53 hours after fertilization in mouthbrooder embryos. A comparative ontogenetic study of embryos and larvae of both ethological types has shown that this difference in timing of morphogenesis also continues in the formation of the nasal pits, the development of the nasal epithelium, and the olfactory nerves. Comparative data are provided for these processes, including LM, TEM and SEM of the stereocilia and kinocilia-bearing cells, as well as of the microvillar cells. The distribution of these cell-types on the olfactory lamellae differs in the various species. The faster development of the nasal organs observed in larvae of bottomspawners as compared to mouthbrooders also matches the development of other vital organs in these two etho-types and seems to be of high adaptive value, providing the bottomspawners at an earlier stage with sensitive organs in the surrounding hostile habitats.     

Structure and development of first-generation teeth in the cichlid Hemichromis bimaculatus (Teleostei, Cichlidae)

In order to build a reference system to assess results of ongoing in vitro experiments on the study of epithelial-mesenchymal interactions during odontogenesis in actinopterygians, we have chosen to study the first-generation teeth of the cichlid Hemichromis bimaculatus from initiation until attachment both at the light and transmission electron microscopical level. Although their development follows the general pattern of teleost tooth formation, first-generation teeth show peculiarities compared with later tooth generations, including their size, bare emergence from the epithelium, absence of dentinal tubules and of nerves and capillaries in the pulp cavity, and organization of the outer dental epithelium. Four developmental stages (a to d) prior to attachment (stage e) have been distinguished. The oral epithelium invaginates into the underlying mesenchyme (stage a) and is later folded to form a bell-shaped dental organ (stage b) without any primordial thickening, or any other morphological indication of imminent invagination. Then, the collagenous enameloid matrix is laid down, most probably by the odontoblasts (early stage c), soon followed by predentine deposition and the beginning of enameloid mineralization (late stage c). With ongoing dentinogenesis, the enameloid matrix matures (stage d), i.e. the organic constituents are removed and the matrix further mineralizes. Finally (stage e), an annular collar of attachment bone is deposited to fix the tooth onto the underlying bone.     

Cytological and morphological ontogenesis and involution of the thymus in cichlid fishes (Cichlidae,Teleostei)

The ontogenesis and involution of thymus in cichlid fishes was studied with the aim of comparing development in the bottom-spawning species Tilapia zillii and T. tholloni, and in the mouth-brooding species Oreochromis auratus, O. niloticus, O. mossambicus, and Sarotherodon galilaeus. For comparison, data are also given on bottom-spawning Cichlasoma spp. from America and mouth-brooding Pseudotropheus auratus and Aulonocara nyassae from Africa. Developmental changes were followed histologically by means of light and electron microscopy of sections, beginning with embryos 24 h after fertilization, up to 14-year-old specimens of O. auratus. In all these fish, the anlagen of the thymus glands begins from the third and fourth gill pouches, and their development shows a high correlation with the pace of general organogenesis, which differs greatly in the bottom-spawning and mouth-brooding cichlids. In juveniles of bottom-spawners of 20–40 mm total length and in mouth-brooders of 40–60 mm total length, three cell types are present in the thymus: thymocytes, with large, dense nuclei; epitheliocytes, with long cell extensions containing bundles of tonofibrils; and reticulocytes, with short, granulated cell extensions. Hassall's corpuscles start to develop in larvae of T. zillii at 20–35 mm total length, and in specimens of 40 mm and more total length the corpuscles are typical, formed by inner and outer rings of epitheliocytes. At 30–45 mm total length, cell debris starts to accumulate in the interior of the corpuscles as an early sign of regression. As involution continues, macrophages accummulate within and around the Hassall's corpuscles. The epitheliocyte rings are eventually completely broken down. Isles of thymocytes persist in tilapias from the age of 1–14 years, but most of the thymus volume is occupied by blood lacunae and pigmented macrophage aggregations. The morphology is similar in the mouth-brooding species Pseudotropheus beginning at 1.5 years of age. © 1995 Wiley-Liss, Inc.     

A new Apistogramma species (Teleostei, Cichlidae) from the Rio Negro in Brazil and Venezuela

Apistogramma diplotaenia sp.n. occurs in the upper and middle Rio Negro. The largest among 186 specimens studied is a 29.2 mm SL male, the largest female is 22.9 mm SL. Distinguishing characters include a unique colour pattern, consisting of two dark bands along the side which converge anteriorly and posteriorly. A comparative osteological study, including other species of Apistogramma and the genera Taeniacara and Apistogrammoides , indicates a progressive forwards loss of infraorbital ossicles and foramina among dwarf cichlids. The long, rod-like interarcual cartilage of Apistogramma, Apistogrammoides and Satanoperca possibly represents a plesiomorphic condition otherwise unknown among cichlids.     

Ontogenesis and functional metamorphosis of the head-kidney in bottomspawner and mouthbrooder cichlid fishes (Cichlidae,Teleostei)

Histological and electron-microscopical studies of the head-kidney ontogenesis in a group of bottomspawner and mouthbrooder cichlid fishes reveal that this organ undergoes three functional and structural stages. At the initial stage, 2–12 days after fertilization, the head-kidney is a filtering organ of two glomeruli and adjacent renal tubules; the second stage, beginning 12–30 days after fertilization, is a functionally mixed one, with the nephron and developing hemopoietic and endocrine, chromaffin, and interrenal cells; during the third stage, which begins 30–60 days after fertilization, the nephron degenerates, and the head-kidney starts functioning solely as a lympho-adrenal organ. With age the interrenal tissue expands, forming fields, especially along the blood vessels. The number of melano-macrophage centers progressively increases within the hemopoietic parts. Data are given on the various cell types, the reticuloendothelial stroma, and the renal part of this organ in various species. The earlier ontogenesis and function of the head-kidney, as thymus and other organs, in bottomspawner larvae compared to the more protected mouthbrooders provide support for earlier free functioning in a dangerous environment. © 1996 Wiley-Liss, Inc.     

Ultrastructural and Phylogenetic Description of Kudoa orbicularis n. sp. (Myxosporea: Multivalvulida): A Parasite Infecting the Muscle of the Fish Chaetobranchopsis orbicularis (Teleostei: Cichlidae) in the Amazon Region

A new myxosporean species is described from the muscle of the Amazonian freshwater fish Chaetobranchopsis orbicularis (Teleostei, Cichlidae), with basis on morphometric, ultrastructural and molecular data. Numerous myxospores were observed within pseudocysts located on the hosts' dorsal and ventral muscles, near the neural spines and neural canal (spinal cord). Mature myxospores quadrangular with rounded ends in apical view, measuring 4.3 (3.6–5.0) μm in length and 5.1 (4.2–5.8) μm in width. The myxospores wall is formed by four symmetric valves. Within, four pyriform polar capsules, 2.1 (1.7–2.6) μm long and 1.3 (0.9–1.7) μm wide, located two by two in opposite sides of the myxospores longitudinal axis, each containing a polar filament forming 2–3 coils. Molecular analysis of the SSU rRNA gene by maximum likelihood, neighbor‐joining and maximum parsimony confirms the parasite as a new member of the genus Kudoa, herein named Kudoa orbicularis n. sp., the second species of its genus reported from the South American freshwater fauna, and the fourth species worldwide known to occur in the freshwater environment. Furthermore, its sequence of the SSU rRNA gene constitutes the first entry of a freshwater Kudoa species in GenBank.     

Rapid adaptation to a novel light environment: The importance of ontogeny and phenotypic plasticity in shaping the visual system of Nicaraguan Midas cichlid fish (Amphilophus citrinellus spp.)

Colonization of novel habitats is typically challenging to organisms. In the initial stage after colonization, approximation to fitness optima in the new environment can occur by selection acting on standing genetic variation, modification of developmental patterns or phenotypic plasticity. Midas cichlids have recently colonized crater Lake Apoyo from great Lake Nicaragua. The photic environment of crater Lake Apoyo is shifted towards shorter wavelengths compared to great Lake Nicaragua and Midas cichlids from both lakes differ in visual sensitivity. We investigated the contribution of ontogeny and phenotypic plasticity in shaping the visual system of Midas cichlids after colonizing this novel photic environment. To this end, we measured cone opsin expression both during development and after experimental exposure to different light treatments. Midas cichlids from both lakes undergo ontogenetic changes in cone opsin expression, but visual sensitivity is consistently shifted towards shorter wavelengths in crater lake fish, which leads to a paedomorphic retention of their visual phenotype. This shift might be mediated by lower levels of thyroid hormone in crater lake Midas cichlids (measured indirectly as dio2 and dio3 gene expression). Exposing fish to different light treatments revealed that cone opsin expression is phenotypically plastic in both species during early development, with short and long wavelength light slowing or accelerating ontogenetic changes, respectively. Notably, this plastic response was maintained into adulthood only in the derived crater lake Midas cichlids. We conclude that the rapid evolution of Midas cichlids’ visual system after colonizing crater Lake Apoyo was mediated by a shift in visual sensitivity during ontogeny and was further aided by phenotypic plasticity during development.     

Observations on tooth development and implantation in the upper pharyngeal jaws in Astatotilapia elegans (Teleostei,Cichlidae)

Prominent stages in the development of teeth, associated with the upper pharyngeal jaws in early postembryonic stages of the mouth brooding cichlid A statotilapia elegans were studied on semithin sections in relation to changes in the underlying endoskeletal parts and to the formation of the dentigerous bone. Because the pattern of tooth implantation on infrapharyngobranchial III-IV is constant, at least in early postembryonic stages, it is possible to trace the life history of a given tooth by tracing its homologue throughout the ontogenetic series. A probable causal relationship exists between tooth development and erosion of the underlying cartilage. Fully developed, though unerupted teeth, differentiate annular bone of attachment, which, depending on its position, is formed either outside the cartilage or within the previously induced erosion cavities. Attachment bone of adjacent teeth fuses to build up the dentigerous bone, which, as a result, may be situated within the area previously occupied by cartilage. As soon as the tooth has built up its bone of attachment, it may erupt. The collagenous matrix between tooth and attachment bone persists and gives rise to the movable connection between both. Differentiation of teeth on infrapharyngobranchial III-IV, together with enlargement of the dentigerous bone, proceeds from the lateral and the rostral border, where new germs constantly form. The appearance of new germs on infrapharyngobranchial II is more unpredictable.     

A new species and geographical variation in the Telmatochromis temporalis complex (Teleostei, Cichlidae) from Lake Tanganyika

Telmatochromis brachygnathus sp.n. is described from the southern and central parts of Lake Tanganyika. It can be distinguished from the similar T. temporalis mainly by its smaller mouth. Morphological distinct populations were found in both species.     

Scanning electron microscopy of photoreceptor cells in the light- and dark-adapted retina of Haplochromis burtoni (Cichlidae,Teleostei)

Summary The photoreceptor layer in the retina of Haplochromis burtoni (Cichlidae, Teleostei) was studied by scanning electron microscopy. Three types of receptors were identified: rods, single-cones and double-cones. The three-dimensional arrangement of these photoreceptors is described in the light- and dark-adapted retina. The surface of the inner segment of the photoreceptor cells displays fine vertical fissures which give rise to slender processes. These so called calycal processes which are of different lengths in rods and cones, surround the beginning of the smooth-surfaced outer segment. The myoid, the contractile part of the receptor, which is located beneath the ellipsoid, was examined in the single-cones of the dark-adapted retina. It is a slender structure with surface infoldings. The myoid, studied by transmission electron microscopy, contains bundles of parallel myofilaments, which are thought to be contractile.This investigation was supported by grants of the Deutsche Forschungsgemeinschaft (Sonderforschungsbereich 51-E/10)     

Functional microsatellite and possible selective sweep in natural populations of the black-chinned tilapia Sarotherodon melanotheron (Teleostei, Cichlidae)

The Prolactin I (PrlI) gene is a key locus involved in fish osmotic regulation. Two microsatellites, PrlI GT and PrlI AC, are present in the promoter region of this gene in tilapias. One of these microsatellite is associated to PrlI gene expression and growth rate. Aquaria reared individuals homozygous for long microsatellite alleles at the PrlI AC locus expressed less PrlI in fresh water than fishes with other genotypes. To assess the occurrence of selection acting on these microsatellites we carried out a survey of their variability compared with four others microsatellites in natural populations. Samples of two tilapia species were studied, Oreochromis niloticus (6 samples) a typically freshwater species inhabiting osmotically stable environments and Sarotherodon melanotheron (5 samples) a brackish water species obliged and able to adapt very rapidly to any salinity changes. For both species the different populations studied exhibited a high amount of differentiation (all F(st) value were statistically significant) at every loci. Only one locus, PrlI AC, was monomorphic for the same allele in all S. melanotheron populations. We have hypothesized that this outlier locus with its unique allele could have experienced a selective sweep that took place in the early stages of the species origin. S. melanotheron represents an example of adaptation (to fluctuant salinity environment) acting indirectly on gene product through its promoter.