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.     

Ontogeny,variation, and homology in Salvelinus alpinus caudal skeleton (Teleostei: Salmonidae)

The ontogeny of the caudal skeleton in the Arctic charr, Salvelinus alpinus was examined using an extensive series of cleared and stained specimens. We demonstrate the presence of skeletal components never reported previously within the Salmonidae. In contrast to the generalized condition for salmonids, seven hypurals (instead of six), and four uroneurals (instead of three) have been found in some specimens. Variation in the number and condition of epurals is documented. New hypotheses are proposed concerning (1) relationships among centra and their associated elements, (2) phylogenetic distribution of caudal characters within the Salmonidae, and (3) homology of caudal components. Using the published phylogenetic hypotheses, we provide evidence, that a seventh hypural and a fourth uroneural are taxic atavism in salmonids. The development of the salmonid homocercal fin is discussed in the light of a polyural scheme based on evidences of a one‐to‐one relationship among ural centra and their associated elements. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Microanatomy of the paired‐fin pads of ostariophysan fishes (Teleostei: Ostariophysi)

Members of the teleost superorder Ostariophysi dominate freshwater habitats on all continents except Antarctica and Australia. Obligate benthic and rheophilic taxa from four different orders of the Ostariophysi (Gonorynchiformes, Cypriniformes, Characiformes, and Siluriformes) frequently exhibit thickened pads of skin along the ventral surface of the anteriormost ray or rays of horizontally orientated paired (pectoral and pelvic) fins. Such paired‐fin pads, though convergent, are externally homogenous across ostariophysan groups (particularly nonsiluriform taxa) and have been considered previously to be the result of epidermal modification. Histological examination of the pectoral and/or pelvic fins of 44 species of ostariophysans (including members of the Gonorynchiforms, Cypriniformes, Characiformes, and Siluriformes) revealed a tremendous and previously unrecognized diversity in the cellular arrangement of the skin layers (epidermis and subdermis) contributing to the paired‐fin pads. Three types of paired‐fin pads (Types 1–3) are identified in nonsiluriform ostariophysan fishes, based on differences in the cellular arrangement of the epidermis and subdermis. The paired‐fin pads of siluriforms may or may not exhibit a deep series of ridges and grooves across the surface. Two distinct patterns of unculus producing cells are identified in the epidermis of the paired‐fin pads of siluriforms, one of which is characterized by distinct bands of keratinization throughout the epidermis and is described in Amphilius platychir (Amphiliidae) for the first time. General histological comparisons between the paired fins of benthic and rheophilic ostariophysan and nonostariophysan percomorph fishes are provided, and the possible function(s) of the paired‐fin pads of ostariophysan fish are discussed. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Sperm of Doradidae (Teleostei: Siluriformes)

Spermatic characteristics were studied in 10 species representing several distinct groups within the catfish family Doradidae. Interestingly, different types of spermatogenesis, spermiogenesis and spermatozoa are correlated with intrafamilial groups previously proposed for Doradidae. Semi-cystic spermatogenesis, modified Type III spermiogenesis, and biflagellate sperm appear to be unique within Doradidae to the subfamily Astrodoradinae. Other doradid species have sperm with a single flagellum, cystic spermatogenesis, and spermiogenesis of Type I (Pterodoras granulosus, Rhinodoras dorbignyi), Type I modified (Oxydoras kneri), or Type III (Trachydoras paraguayensis). Doradids have an external mode of fertilization, and share a few spermatic characteristics, such as cystic spermatogenesis, Type I spermiogenesis and uniflagellate sperm, with its sister group Auchenipteridae, a family exhibiting sperm modifications associated with insemination and internal fertilization. Semi-cystic spermatogenesis and biflagellate spermatozoa are also found in Aspredinidae, and corroborate recent proposals that Aspredinidae and Doradoidea (Doradidae + Auchenipteridae) are sister groups and that Astrodoradinae occupies a basal position within Doradidae. The co-occurrence in various catfish families of semi-cystic spermatogenesis and either biflagellate spermatozoa (Aspredinidae, Cetopsidae, Doradidae, Malapturidae, Nematogenyidae) or uniflagellate sperm with two axonemes (Ariidae) reinforces the suggestion that such characteristics are correlated. Semi-cystic spermatogenesis and biflagellate sperm may represent ancestral conditions for Loricarioidei and Siluroidei of Siluriformes as they occur in putatively basal members of each suborder, Nematogenyidae and Cetopsidae, respectively. However, if semi-cystic spermatogenesis and biflagellate sperm are ancestral for Siluriformes, cystic spermatogenesis and uniflagellate sperm have arisen independently in multiple lineages including Diplomystidae, sister group to Siluroidei.     

Locomotor activity rhythms in cave fishes from Chapada Diamantina, northeastern Brazil (Teleostei: Siluriformes)

Previous studies on the locomotor activity of troglobitic (exclusively subterranean) species have shown that circadian rhythmicity may be reduced in populations evolving in the absence of zeitgebers such as daily cycles of light and temperature; therefore, circadian activity rhythms, although not infradian nor ultradian rhythms, seem to have been selected by external, ecological factors. We studied the locomotor activity of a highly specialized Heptapteridae catfish (undescribed genus and species) from Chapada Diamantina, NE Brazil, compared to another specialized Brazilian troglobitic heptapterid, Taunayia sp. Locomotor activity was continuously measured in the laboratory with an infra-red photocell system. Seven specimens of the new genus were tested, each one during 14 consecutive days according to the following schedule: three days in DD → seven days in LD (12:12 h) → four days in DD. Data were submitted both to fast Fourier transform periodogram followed by Siegel's test of significance and Lombs - Scargle periodogram techniques in order to identify spectral composition of the time series. In general, results were similar to those obtained for Taunayia sp.: (a) for most specimens, absence of significant circadian components in locomotor activity under DD; (b) for all specimens, significant circadian components under LD, with higher levels of activity during the dark phase, as expected for species belonging to nocturnal epigean taxa; (c) for most specimens, no residual oscillations recorded when free-running conditions were reinstalled. Circadian locomotor activity detected under LD may thus be interpreted as a direct, masking effect of the LD cycle. This suggests a pattern for highly specialized troglobitic species, isolated for a long time in the subterranean habitat, with a progressive reduction of circadian time-keeping mechanisms. Our studies also demonstrate the potential of subterranean organisms for investigation of the origin, evolution, functioning and genetics of circadian rhthmicity.     

Phylogeny of the Amphiliidae (Teleostei: Siluriformes)

The freshwater African catfish family Amphiliidae had been reviewed based on the 73 osteological characters with Diplomystidae, 2 Hypsidoridae, Amblycipitidae, Sisoridae, and Bagridae as out-groups. Because the family position of Leptoglanis (Bagridae/Amphiliidae) is under debate, this genus has been taken as an out-group too. Results of the study indicate that: 1) the Amphiliidae is not a monophyletic group and must now be restricted to the genera Amphilius and Paramphilius; the two subfamilies Amphiliinae and Doumeinae are separated by the sisorids Euchiloglanis (with most of the glyptosternid fishes) and Glyptothorax (with most of the non-glyptosternid fishes); 2) no synapomorphies were found for the subfamily Amphiliinae. 3) The five genera of subfamily Doumeinae constitute a monophyletic group, Andersonia being the sister-group of the four other genera; subfamily Doumeinae + Leptoglanis form the family Doumeidae. The glyptosternids no longer belong to the Sisoridae (family restricted to the non-glyptosternids) and represent the new family Glyptosternidae.     

Functional morphology of the caudal skeleton in teleostean fishes

The basic function of the caudal skeleton in teleostean fishes is to support the caudal fin, but its parts contribute to this function in somewhat different ways. The main axis for this support is the upturned terminal end of the vertebral column, which ends at the base of the uppermost principal rays. The uroneural struts just ahead of this axis provide support for it. The parts of the caudal skeleton behind and below this upturned axis, the hypurals and parhypural, not only support the caudal rays but also provide a means for differential movements between the upper and lower parts of the fin base. This basic caudal skeleton varies with the position of the fish in the sequence of teleosten evolution, the way in which the fish uses its caudal fin, and to some extent with the shape of the fin.     

Mitochondrial genome of Silurus asotus (Teleostei: Siluriformes)

The complete mitogenome sequence of the Amur catfish Silurus asotus was determined using long PCRs. The genome was 16,528 bp in length and contained 13 protein-coding genes, 2 rRNA genes, 22 tRNA genes, and 1 control region; the gene composition and order of which was similar to most other vertebrates. The overall base composition of the heavy strand is 30.5% A, 25.8% T, 28.0% C, and 15.8% G, with an AT content of 56.3%. The mtDNA sequence of S. asotus shared 93.6% and 90.6% sequence identity with that of Silurus meridionalis and Silurus glanis. This mitogenome sequence data would play an important role in silurid catfish phylogenetics and siluriform catfish systematics in general.     

Intraspecific variation in tolerance of warming in fishes

Intraspecific variation in key traits such as tolerance of warming can have profound effects on ecological and evolutionary processes, notably responses to climate change. The empirical evidence for three primary elements of intraspecific variation in tolerance of warming in fishes is reviewed. The first is purely mechanistic that tolerance varies across life stages and as fishes become mature. The limited evidence indicates strongly that this is the case, possibly because of universal physiological principles. The second is intraspecific variation that is because of phenotypic plasticity, also a mechanistic phenomenon that buffers individuals’ sensitivity to negative impacts of global warming in their lifetime, or to some extent through epigenetic effects over successive generations. Although the evidence for plasticity in tolerance to warming is extensive, more work is required to understand underlying mechanisms and to reveal whether there are general patterns. The third element is intraspecific variation based on heritable genetic differences in tolerance, which underlies local adaptation and may define long-term adaptability of a species in the face of ongoing global change. There is clear evidence of local adaptation and some evidence of heritability of tolerance to warming, but the knowledge base is limited with detailed information for only a few model or emblematic species. There is also strong evidence of structured variation in tolerance of warming within species, which may have ecological and evolutionary significance irrespective of whether it reflects plasticity or adaptation. Although the overwhelming consensus is that having broader intraspecific variation in tolerance should reduce species vulnerability to impacts of global warming, there are no sufficient data on fishes to provide insights into particular mechanisms by which this may occur.     

Ontogeny of the catfish pectoral-fin spine (Teleostei: Siluriformes)

The characteristic and morphologically variable pectoral-fin spine of catfishes (order Siluriformes) has been well-investigated based on later developmental stages (juveniles and adults) but information on the earliest life stages are lacking. Here, we document the ontogeny of pectoral-fin spines in four siluroid (Ictalurus punctatus, Noturus gyrinus, Silurus glanis and Akysis vespa) and two loricarioid catfishes (Corydoras panda and Ancistrus sp.). To further our understanding of pectoral-fin spine development, we also examined adult and juvenile specimens representing 41 of the currently 43 recognized families of catfishes. Development of the pectoral-fin spine is similar in all catfishes and resembles the development of a typical soft fin ray. Fusion between hemitrichia of the anteriormost lepidotrichium occurs proximally first, forming the spine proper, with growth of the spine occurring through the subsequent fusion of developing distal hemitrichial segments that comprise the spurious ray. The variation of pectoral-fin spine morphology observed is largely attributed to the presence/absence of five traits, which either develop as part of the hemitrichial segments that are added to the distal tip of the spine during growth (distal rami, anterior/posterior serrae) or develop independent of these segments (denticuli and odontodes).     

Nomenclature of cartilaginous elements in the caudal skeleton of teleostean fishes

Nomenclature and abbreviations are proposed for the cartilaginous elements of the caudal skeleton of teleostean fishes. These were developed on the basis of examination of 510 species within 198 families of 31 orders and the determination of the positional relationship between these structures and the bony elements. A review of the most important relative literature is also provided.     

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.     

When two equals three: developmental osteology and homology of the caudal skeleton in carangid fishes (Perciformes: Carangidae)

Ontogeny often provides the most compelling evidence for primary homology in evolutionary developmental studies and is critical to interpreting complex structures in a phylogenetic context. As an example of this, we document the ontogenetic development of the caudal skeleton of Caranx crysos by examining a series of cleared and stained larval and postlarval specimens. By studying ontogeny, we are able to more accurately identify some elements of the adult caudal skeleton than is possible when studying the adult stage alone. The presence of two epurals has been used as a synapomorphy of Caranginae (homoplastically present in the scomberoidine genera Scomberoides and Oligoplites). Here we find that three epurals (ep) are present in larvae and small postlarval juveniles (i.e.,<25 mm standard length [SL]) of C. crysos and other carangines, but ep2 never ossifies and does not develop beyond its initial presence. Ep2 was last observed in a 33.6 mm SL specimen as a small nodule of very lightly stained cartilage cells and eventually disappears completely. Therefore, the two epurals present in the adult are ep1 and ep3. In other carangines examined (e.g., Selene, Selar), the rudimentary ep2 ossifies and appears to fuse to the proximal tip of ep1. In these taxa, therefore, the two epurals of the adult appear to be ep1+2 and ep3. We found no indication of three epurals at any stage in the development of Oligoplites (developmental material of Scomberoides was unavailable). We discuss the osteology of the caudal skeleton of carangoid fishes generally and emphasize the power and importance of ontogeny in the identification of primary homology.     

The composition of the caudal skeleton of teleosts (Actinopterygil: Osteichthyes)

The diural caudal skeleton of teleostean actinopterygians develops phylogeneticaily and ontogenetically from a polyural skeleton. The reduction of the polyural anlage to four, three, two or fewer centra in the adult caudal skeleton takes different pathways in different genera (e.g. compare Elops and Albula) and groups of teleosts. As a result, ural centra are not homologous throughout the teleosts. By numbering the ural centra in a homocercal tail in polyural fashion, one can demonstrate these and the following differences. The ventral elements (hypurals) always occur in sequential series, whereas the dorsal elements (epurals and uroneurals) may alter like the ural centra. The number of epurals, five or four in fossil primitive teleosts, is reduced in other primitive and advanced teleosts, but the same epurals are not always lost. The number of uroneurals, seven in fossil teleosts, is reduced in living teleosts, but it has not been demonstrated that the first uroneural is always derived from the neural arch of the same ural centrum. The landmark in the homocercal tail is the preural centrum I which can be identified by (1) bifurcation of the caudal artery and vein in its ventral element, the parhypural, (2) its position directly caudal to the preural centrum (PU2) which supports the lowermost principal caudal ray with its haemal spine, (3) carrying the third hypaxial element ventral to the course of arteria and vena pinnalis, and (4) by carrying the first haemal spine (parhypural) below the dorsal end of the ventral cartilage plate. The study of the development of the vertebral column reveals that teleosts have different patterns of centrum formation. A vertebral centrum is a complete or partial ring of mineralized, cartilaginous or bony material surrounding at least the lateral sides of the notochord. A vertebral centrum may be formed by arcocentrum alone, or arcocentral arcualia and chordacentrum, or arco-, chorda- and autocentrum, or arcocentral arcualia and autocentrum. This preliminary research demonstrates that a detailed ontogenetic interpretation of the vertebral centra and of the caudal skeleton of different teleosts may be useful tools for further interpretations of teleostean interrelationships.     

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.     

Character variation in separate body regions of Gerreidae (Osteichthyes: Teleostei) fishes inferred from geometric morphometrics

Body morphology is a valuable feature for distinguishing teleostean fishes. However, the utility of character variation in separate body regions has yet to be tested. The taxonomy of the Gerreidae family is controversial due to character overlapping among its fish species. This work aims to analyze and compare the body shape variation in three regions, cephalic, trunk, and caudal peduncle, using landmark data and geometric morphometric methods in 17 species and five genera of the family Gerreidae. The pattern of shape variation for the cephalic region consisted of well-defined character states exclusive of each species analyzed. Shape variation in the trunk and caudal peduncle regions does not distinguish all species in this study. This study showed that the dorsal cephalic profile is highly variable among the species, therefore, shape variation in this region is useful for distinguishing Gerreidae species. In contrast, some species within the same genus share similar shape states in the trunk and caudal peduncle regions, with the most shape variation in the dorsal profile and anal fin for the trunk and in the middle of the caudal peduncle.     

Development of the cranium and paired fins in Betta splendens (Teleostei: Percomorpha): Intraspecific variation and interspecific comparisons

The development of the chondrocranium and the relative timing of ossification of the osteocranium is described in the teleost fish Betta splendens from a large series of cleared and differentially stained specimens. General trends in ossification patterns are examined from developmental, phylogenetic, and functional contexts. As in many other vertebrates, dermal bones form before cartilage bones. Ossification sequence conforms to functional need in a very general way, but there are many inconsistencies in the details of order. For example, some bones that are directly involved in feeding ossify no earlier than bones more indirectly involved. Comparisons of ossification sequence within specific cranial regions are made among Betta splendens, Oryzias latipes (Atherinomorpha), and Barbus barbus (Ostariophysi) within a phylogenetic framework. Many evolutionary changes in relative sequence of ossification are evident within regions among these taxa, yet many other sequences are conserved. The logistic difficulty of comparing entire cranial ossification sequences (vs. regional sequences) makes evident the need for new methods for identifying and quantifying sequence changes. Intraspecific variation in order of ossification is described for the first time in teleost fishes. To the extent that ossification sequence varies intraspecifically, conclusions drawn from previous interspecific comparisons are compromised. Understanding the importance of changes in ossification order within and among taxa will require experimental, functional, and evolutionary work. © 1996 Wiley-Liss, Inc.     

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.