The monophyly of the Characidiinae, a Neotropical group of characiform fishes (Teleostei: Ostariophysi)

Although virtually no phylogenetic evidence (in the sense advocated by Hennig, 1966) had been previously presented to support the monophyly of the Characidiinae, and most 'diagnostic' characters used by previous authors were found to be unacceptable in a cladistic classification, i t is still possible to diagnose the Characidiinae in a phylogenetic sense. This study revealed the existence of 13 synapomorphies supporting the monophyly of the group. Several of these synapomorphies, such as the modifications associated with the mesethmoid, the jaw bones, and the ribs of the fifth vertebra, are unique to the Characidiinae, thus providing a solid basis for recognizing the group as a monophyletic unit of characiform fishes. Demonstration of characidiin monophyly provides a solid foundation for further phylogenetic analysis of characidiin interrelationships, and higher level relationships among characiform fishes.     

Spermiogenesis and spermatozoal ultrastructure in Trichomycteridae (Teleostei: Ostariophysi: Siluriformes)

Spadella, M.A., Oliveira, C. and Quagio‐Grassiotto, I. 2009. Spermiogenesis and spermatozoal ultrastructure in Trichomycteridae (Teleostei: Ostariophysi: Siluriformes). —Acta Zoologica (Stockholm) 91 : 373–389. Siluriformes comprises the most diverse and widely distributed ostariophysan group, a fish assemblage that includes about three quarters of the freshwater fish of the world. In this study, the ultrastructural characterization of spermiogenesis and spermatozoa in specimens of Copionodontinae (the sister group to all other trichomycterids), Trichomycterinae (a derived trichomycterid group), and Ituglanis (a genus not assigned to any trichomycterid subfamily) is presented. The comparative analyses of the data show that trichomycterid species share six of seven analyzed spermiogenesis characters, reinforcing the monophyly of the group. Analyses of trichomycterid sperm ultrastructure showed that the species studied share the same character states for nine of seventeen characters analyzed. Copionodon orthiocarinatus and Ituglanis amazonicus each share more ultrastructural characters with species of Trichomycterus than with one another. Regarding the families of Loricarioidea, the species of Trichomycteridae share more characters of spermatogenesis, spermiogenesis, and sperm with representatives of the families Callichthyidae, Loricariidae, and Scoloplacidae than with Nematogenyidae, its hypothesized sister group. With the exception of the family Nematogenyidae, the character similarities observed reinforce the monophyly of the superfamily Loricarioidea.     

Development of mandibular,hyoid and hypobranchial muscles in the zebrafish: homologies and evolution of these muscles within bony fishes and tetrapods

Background  

During vertebrate head evolution, muscle changes accompanied radical modification of the skeleton. Recent studies have suggested that muscles and their innervation evolve less rapidly than cartilage. The freshwater teleostean zebrafish (Danio rerio) is the most studied actinopterygian model organism, and is sometimes taken to represent osteichthyans as a whole, which include bony fishes and tetrapods. Most work concerning zebrafish cranial muscles has focused on larval stages. We set out to describe the later development of zebrafish head muscles and compare muscle homologies across the Osteichthyes.     

Occurrence of biflagellate spermatozoa in Cetopsidae, Aspredinidae, and Nematogenyidae (Teleostei: Ostariophysi: Siluriformes)

In the present study spermiogenesis was investigated in Cetopsis coecutiens (Cetopsidae), and Bunocephalus amazonicus (Aspredinidae), while spermatozoa ultrastructure was investigated in C. coecutiens, B. amazonicus, and Nematogenys inermis (Nematogenyidae). Aspredinidae and Cetopsidae share a spermatogenesis of the semicystic type, and a particular type of spermiogenesis process not reported in any fish group. In the three species analyzed, spermatozoa are biflagellate with flagella having the classical axoneme formulae (9 + 2). The analysis of thirteen characters showed the presence of eight characters shared by Cetopsidae and Aspredinidae, and six characters shared by Cetopsidae and Nematogenyidae, which may suggest that these three families may be more related than actually hypothesized, comprising a very primitive siluriform lineage originated after Diplomystidae.     

Relationships among major lineages of characid fishes (Teleostei: Ostariophysi: Characiformes), based on molecular sequence data

The family Characidae is a group of freshwater bony fishes that exhibits high species-level diversity and whose members inhabit parts of Texas, Mexico, and Central and South America. Thus far, morphological data have been of limited use in discerning relationships among subfamilies and incertae sedis genera of the family Characidae. In this study, DNA sequence data from GenBank were combined with new sequences for analyses under Bayesian and parsimony schemes. Sequences fell into four gene partitions, with three genes in the mitochondrial subset (12S, 16S, COI genes) and one gene in the nuclear subset (RAG2 gene). Inferred Bayesian and parsimony-based phylogenies reject the monophyly of certain groups (e.g., Astyanax, Hyphessobrycon, and Bryconamericus), do not reject the monophyly of others (e.g., Cheirodontinae and “clade A” of previous authors), and present new sister-group hypotheses (e.g., Brittanichthys sister to Paracheirodon). Sister to clade A is a lineage referred to herein as “clade B” which includes Exodon and exemplars from Cheirodontinae (the most basal lineage within clade B), Aphyocharacinae, Tetragonopterinae, and Characinae (excluding Gnathocharax). “Clade C” is sister to A + B and contains representatives of large incertae sedis genera (e.g., Hyphessobrycon, Hemigrammus), as well as members of Stethaprioninae. Unless certain other subfamilial names are to be disregarded, the use of Tetragonopterinae should continue to be restricted to species of Tetragonopterus because other genera previously referred to this subfamily grouped in clades A or C, quite distant from Tetragonopterus.     

Interrelationships of the ostariophysan fishes (Teleostei)

The history of ostariophysan classification is summarized and it is noted that traditional concepts of relationships have never been supported by characters found to be unique to the taxa. We present a new hypothesis of relationships among four of the five major ostariophysan lineages: Cypriniformes, Characiformes, Siluroidei, and Gymnotoidei (Otophysi). Cypriniforms are the sister-group of the remaining three (Characiphysi), and characiforms are the sister-group of siluroids plus gymnotoids (Siluriformes). Placement of the Gonorynchiformes as the sister-group of the Otophysi is supported by additional evidence. Each of the five lineages is monophyletic. Analysis was concentrated upon species thought to be the least specialized within each lineage; choices of these species are discussed. Chanos is determined to be a relatively primitive gonorynchiform morphologically and the sister-group of all other Recent members of the order. Opsariichthys and Zacco are found to be morphologically primitive cypriniforms. We propose that a monophyletic group comprising the Citharinidae and Distichodontidae forms the sister-group of all other characiforms. Within the two families, Xenocharax is the least specialized. We suggest that Hepsetus, the erythrinids, and the ctenoluciids are more derived than the distichodontids and citharinids, and may form a monophyletic group within die characiforms. The traditional hypothesis that Diplomystes is the primitive sister-group of all Recent siluroids is substantiated. Our evidence suggests that Sternopygus is the most primitive gymnotoid morphologically; but rather than being the sister-group of all other gymnotoids, it is the primitive sister-group within a lineage called the Sternopygidae by Mago-Leccia. Previous explanations of otophysan distribution have been based on notions of relationships which are unsupported by the evidence presented herein. Our own analysis of relationships serves primarily to make clear the extent of sympatry, and therefore the probability of dispersal, among the major ostariophysan lineages. The extent of sympatry, together with the widespread distribution of ostariophysans, suggests that the group is older than previously supposed, and our hypotheses of relationships among the characiforms implies that many of the extent characiform lineages evolved before the separation of Africa and South America. Further understanding of ostariophysan distribution must await phylogenetic analysis within each of the five major lineages so that distributions linked with vicariance patterns and dispersal events can be sorted out.     

Allometry and developmental integration of body growth in a piranha,Pygocentrus nattereri (Teleostei: Ostariophysi)

Piranhas, like many teleosts, change their diets on both ontogenetic and phylogenetic time scales. Prior studies have suggested that pervasive morphological changes in body form on a phylogenetic time scale may be related to changes in diet, but previous reports have found little shape change in piranhas on an ontogenetic time scale. We re-examine the post-transformational allometry of body form in one piranha, Pygocentrus nattereri (Kner), using the method of thin-plate splines decomposed by their partial warps. We find substantial evidence of allometry, primarily elongation of the mid-body relative to the more anterior and posterior regions, elongation of the postorbital and nape regions relative to the more anterior head and posterior body, and deepening of the head relative to the body. In addition to these pervasive changes throughout the body, there are some that are more localized, especially elongation of the postorbital region relative to eye diameter and snout, and an even more localized elongation of the snout relative to eye diameter. Initial dietary transitions are associated with changes in head and jaw proportions, but rates of shape change decelerate through growth, so that the final transition to a diet increasingly dominated by small whole fish appears associated with change largely in overall body size. © 1995 Wiley-Liss, Inc.     

Systematics of Tanganyikan cichlid fishes (Teleostei: Perciformes)

The relationships among 53 genera of Tanganyikan cichlid fishes were analyzed based on internal and external morphological features. Comparison of the morphological cladistic tree with a previously proposed classification showed 5 of 12 tribes to be nonmonophyletic. Sixteen tribes were recognized, the changes in classification being that Trematocarini was treated as a junior synonym of Bathybatini; 5 new tribes were established for each of the following genera, Benthochromis, Boulengerochromis, Ctenochromis benthicola, Cyphotilapia, and Greenwoodochromis; Ctenochromis horei was transferred from Haplochromini to Tropheini; and Gnathochromis pfefferi was transferred from Limnochromini to Tropheini. The revised classification was supported by previously proposed molecular trees.     

Spermiogenesis and spermatozoa ultrastructure in Salminus and Brycon, two primitive genera in Characidae (Teleostei: Ostariophysi: Characiformes)

In Salminus, spermiogenesis is cystic and gives origin to a type I aquasperm. Spermatid differentiation is characterized by chromatin condensed into thick fibres, nuclear rotation, nuclear fossa formation, cytoplasmic channel formation, mitochondrial fusion producing long and ramified mitochondria, and the presence of several membranous concentric rings around the plasma membrane that encircles the cytoplasmic channel. In Salminus and Brycon, spermatozoa are very similar. They exhibit a spherical nucleus and chromatin condensed into fibre clusters, and a deep nuclear fossa. They show a long midpiece with few elongate mitochondria at the initial region and a cytoplasmic channel completely encircled by one or two membranous concentric rings. The flagellar axis is perpendicular to the nucleus and exhibits the classic axoneme (9 + 2). The very strong similarity observed between Salminus and Brycon spermatozoa supports the hypothesis that these subfamilies are likely to have a monophyletic origin.     

Phylogenetic relationships among atheriniform fishes (Teleostei: Atherinomorpha)

We present evidence from adult and larval morphology for the monophyly and relationships of Atheriniformes, using other atherinomorphs, mugilids and acanthomorph fishes as outgroups. Atheriniformes is diagnosed by ten characters (larval: short preanal length, single mid-dorsal row of melanophores; adult: vomerine ventral face concave, long Al muscle tendon to lacrimal, two anterior infraorbital bones, pelvic-rib ligament, pelvic medial plate not extended to anterior end, and second dorsal-fin spine flexible). We recognize six families within the order, the hierarchical relationships among which are: (Atherinopsidae (Notocheiridae (Melanotaeniidae (Atherionidae (Phallostethidae, Atherinidae))))). Other major conclusions include: (1) Atherinopsidae (Menidiinae, Atherinopsinae) is diagnosed by 20 characters (e.g. ethmomaxillary ligament attached to palatine dorsal process, ventral postcleithrum with two dorsal rami); (2) Melanotaeniidae (Bedotiinae (Melanotaeniinae (Telmatherinini, Pseudomugilini))) is diagnosed by six characters (e.g. absence of second dorsal-fin spine, sexual dimorphism in body colour and median-fin development, greater body depth); (3) Dentatherina is in Phallostethidae; (4) Atherinidae (Atherinomorinae (Craterocephalinae, Atherininae)) is diagnosed by three characters (lacrimal notch, ventral postcleithrum between first and second pleural ribs, pelvic ventral spine); (5) Atherinidae and Phallostethidae form the Atherinoidea clade diagnosed by seven characters (e.g. interopercle dorsal process absent, dorsal wings of urohyal absent, ventral postcleithrum laminar, pelvic medial plate extended to anterior end, presence of anal plate). Bedotia, Rhodes , and melanotaeniines are shown to be derived within atheriniforms rather than the plesiomorphic sister groups to a paraphyletic 'atherinoid' group. We also demonstrate that groups traditionally placed in Atherinidae (Menidiinae, Atherininae, Atherioninae, etc.) comprise a paraphyletic assemblage.     

Determining Management Units in Southeastern Brazil: The Case of Astyanax bimaculatus (Linnaeus, 1758) (Teleostei: Ostariophysi: Characidae)

Characterization, management and protection of biodiversity are the most prominent challenges in conservation biology. Analyses on molecular similarity (Random Amplified Polymorphic DNA-Polymerase Chain Reaction, RAPD-PCR) and morphologic patterns (morphometric and meristic characters) were performed to evaluate the existence of population structuring in the Doce river basin, in a widespread small-sized characin, Astyanax bimaculatus. Three hundred (300) individuals were collected from six locations isolated by waterfalls or dams, in the Doce river basin, Minas Gerais State, Brazil. Genetic differentiation among tributaries was significant (p<0.00001) and accounted for 21% of total variance (p<0.00001). Only the Grande waterfall of the Casca river was an effective factor for differentiation of populations (p<0.00001), and meristic characters were also consistent with molecular data. We concluded that genetic and morphologic variation of this species was not necessarily associated with waterfalls and that molecular and meristic data are effective predictors of population divergence in this basin. The consequences of these findings for the protection and management of aquatic biodiversity were discussed.     

Observations on the structure of larval attachment organs in three species of gymnotiforms (Teleostei: Ostariophysi)

Attachment organs are known in certain teleosts but have not been reported in gymnotiforms. This study demonstrates the presence and describes the structure of attachment organs in three species of gymnotiforms: the apteronotids Apteronotus albifrons and A. leptorhynchus , and the rhamphichthyid Rhamphichthys sp. Attachment organs are present only during the first few days after hatching. Organs were studied with light microscopy, SEM, and TEM. In all three species attachment organs are restricted to the head region. They consist of numerous individual attachment cells and modified epithelial cells distributed on the frontal, lateral, and dorsal head area. Only the apical parts of the attachment cells reach the body surface between the epithelium of the epidermis. Attachment cells contain considerable amounts of rough ER and Golgi vesicles filled a the glutinous substance. Epithelial cells in the area in which attachment cells are developed differ in structure from those on the rest of the body and form a functional complex with the attachment cells. In the two Apteronotus species they possess 4–10 μm long projections that are located in the cell centre in A. albifrons , but are slightly shifted to one side of the cells in A. leptorhynchus . Epithelial cells in the organ area of Rhamphichthys sp. lack any projections and resemble cells of the trunk region. However, their pattern of microridges on the outer cell surface is more pronounced and conspicuous. Projections along with modified microridges may aid in distributing and focusing adhesive substances released by attachment cells, and thus may enhance adhesive properties.     

The evolution of pharyngeal bones and teeth in Gobioninae fishes (Teleostei: Cyprinidae) analyzed with phylogenetic comparative methods

The subfamily Gobioninae is a subgroup in the specious fish family Cyprinidae, which bears high diversity in morphological and ecological dimensions and has its most components distributed in East Asia. In this study, the pharyngeal bones and teeth of 39 species belonging to 19 genera of the Gobioninae were examined, with the phylogenetic comparative method (PCM) and correlation methods employed to analyze the character evolution. Three characters on pharyngeal bones (shape of the pharyngeal bones, extension for attachment of the pharyngo-cleithralis internus posterior (PCIP) muscle, and teeth-bearing area) and six characters of pharyngeal teeth (shape of the five teeth in the main row, number of rows of the teeth) were identified and compared. When the character states were mapped on a molecular phylogenetic tree, it was found that, to adapt to different masticatory operations, different Gobioninae species have various morphological types of pharyngeal bones and teeth: some have intermediate pharyngeal bones bearing multiple rows of diverse teeth (conical, coarsely compressed, and compressed), others have broad pharyngeal bones bearing a single row of molar teeth, and still others have narrow pharyngeal bones bearing a single row of extremely compressed teeth. Tests on the phylogenetic signal and evolutionary associations revealed that evolution of the examined characters was all phylogenetically constrained and correlated. Owing to the homoplasy in evolution, it was suggested that the conventional method of using pharyngeal bones and teeth for phylogenetic reconstruction of cyprinid fishes should not be encouraged.     

Homology and evolution of the opercular series in the loricarioid catfishes (Pisces: Siluroidei)

Aspects of the morphology of the opercular region of siluroids (catfishes) were examined to determine the homologies of the opercular bones of loricarioid catfishes, a diverse monophyletic neotropical group. Homology of the component structures was inferred by examination of three criteria: (1) functional relations to the jaw abduction mechanism of primitive halecostome fishes; (2) positional relations to the bones of the suspensorium and the path of the preopercular laterosensory canal; and (3) pattern of development of the preopercular canal and lateral cheek plates of loricarioid catfishes. The majority of siluroids have lost one or more of the preopercular canal exit branches to the skin surface present in most primitive siluroids. A number of specializations of the canal pathway and components of the opercular series have occurred in loricariids. Trichomycterids have the preopercular canal reduced to a short branch from the pterotic. Callichthyids share the presence of a preopercular canal with primitive siluroids, but have lost the communication of the canal between the preopercle and pterotic bones. Callichthyids and scoloplacids share one additional loss of a preopercle exit. The second of two subopercular elements of astroblepids is homologous with the interhyal. Astroblepids and loricariids have lost one additional preopercle exit, the interoperculo-mandibular ligament and the corresponding biomechanical couple for lower jaw abduction. Loricariids have probably lost the interopercle as well. These derived features of loricariids and astroblepids are viewed as specializations for a particular feeding behaviour: scraping algae and detritus from the substratum.     

Mechanisms of light organ occlusion in flashlight fishes, family Anomalopidae (Teleostei:Beryciformes), and the evolution of the group

The circumtropical, nocturnal, shore-fish family Anomalopidae is characterized by a subocular luminous organ containing symbiotic luminous bacteria. The five known species are placed in four genera, one of which is new. Phthanophaneron is restricted to the eastern Pacific, Kryptophanaron to the western Atlantic, Photoblepharon is Indo-West Pacific in distribution and Anomalops is west Pacific. The symbiotic bacteria emit light continuously, and two superficially different mechanisms of occluding the glowing face of the organ are found. In Photoblepharon a black shutter of elastic skin is drawn up over the face of the organ, whereas in Anomalops the organ is rotated downward, so that only the heavily pigmented back of the organ is exposed. In Phthanophaneron and Kryptophanaron, both rotational and shutter mechanisms are present. Elucidation of the structures and linkages involved in light-organ occlusion reveals that the superficially different mechanisms are based on a common functional complex. In all four genera, the light organ is supported by a cartilaginous cup that articulates anteriorly with a cartilaginous stalk. Motive power for both the shutter and rotational mechanisms is supplied by the adductor mandibulae through a complex biomechanical linkage involving the ethmomaxillary ligament and a ligament unique to anomalopids, the Ligament of Diogenes. The structures involved in shutter erection and organ rotation are illustrated and described in detail for Photoblepharon and Anomalops and are compared with those in the other two forms; a functional hypothesis is advanced. Extrafamilial relationships of the Anomalopidae are discussed, and a hypothesis of the phylogenetic relationships of the four genera is derived from a cladistic analysis involving 19 non-light-organ characters and corroborated by some light-organ characters. Most characters associated with the light-organ complex cannot be polarized by conventional outgroup comparison, and the evolution of the light organ occlusion mechanisms is interpreted in light of the hypothesized phylogeny and a hypothesized ancestral mechanism. We propose that the common ancestor of anomalopids possessed a forced rotational mechanism like that of Phthanophaneron and Kryptophanaron. This was refined to a more efficient flipping rotational mechanism in Anomalops, the sister group of the lineage comprising the other three genera, within which the shutter mechanism was progressively refined. The ostensibly unnecessary complexity of the shutter mechanism is apparently a result of functional-morphological constraints imposed on the system by the pre-existence of a rotational mechanism. A brief zoogeographic scenario is proposed.     

Comparative anatomy of the cheek muscles of Korean cobitid fishes (Ostariophysi: Cypriniformes), with the comments on their phylogenetic relationships

Cheek muscles of 16 species of all Korean cobitid species belong to six genera were examined and described in detail, with comments on their phylogenetic relationships made on the basis of characteristics of cheek muscles. Korean cobitids were divided into two groups, the Cobitis and Misgurnus groups, by five characters related to three cheek muscles (adductor mandibulae, preorbitalis, and retractor arcus palatine). The former group, comprising Cobitis, Iksookimia, Kichulchoia, Koreocobitis, and Niwaella, is defined by four apomorphies, and the latter, including Misgurnus, by a single apomorphy.     

Embryonic growth and trophotaenial development in goodeid fishes (Teleostei: Atheriniformes)

Embryonic growth and trophotaenial development are examined in two species of goodeid fish, Ameca splendens and Goodea atripinnis. During gestation of A. splendens, embryonic dry mass may increase from 0.21 mg at the onset of development to 31.70 mg at term. In G. atripinnis, embryonic dry mass ranges from 0.25 mg at the onset of development to 3.15 mg at term. Increase in mass is primarily due to the uptake of maternally derived nutrients by trophotaeniae, externalized embryonic gut derivatives. Trophotaenial development in both species is divisible into five phases. During the first phase, the anus is formed. The second phase involves dilation of the anus, enlargement of the perianal lips, differentiation of the hindgut absorptive epithelium, and formation of the trophotaenial peduncle. The third phase is characterized by a further marked hypertrophy and lateral expansion of the perianal lips that results in the formation of short trophotaenial processes. During the fourth phase, there is continued outward expansion of the inner mucosal surface of the trophotaenial peduncle that results in its eversion and lobulation. Placental function is established by this phase. Axial elongation and dichotomous branching of trophotaenial processes occurs during the fifth phase. Development of rosette and ribbon trophotaeniae differ in the degree of axial elongation during the fifth and final phase.