Molecular phylogeny of bony fishes,based on the amino acid sequence of the growth hormone

Bony fishes (Osteichthyes) comprise over 22,000 species, about half of all vertebrate species. In order to investigate the phylogenetic relationships within this vertebrate class, we have studied the only protein whose primary structure is known in a rather large (27) number of fish species belonging to seven orders—the growth hormone. The phylogeny obtained using the maximum parsimony method based on amino acid sequences represents the first molecular phylogeny of teleostean fishes based on an extensive set of data. This phylogeny agrees remarkably well with the generally accepted phylogeny based on morphological characters and paleontological data. Correspondence to: G. Bernardi     

A phylogeny of sparoid fishes (Perciformes, Percoidei) based on morphology

 The putative percoid superfamily Sparoidea includes the Nemipteridae, Lethrinidae, Sparidae, and Centracanthidae. Although a rigorous cladistic analysis has never been attempted, two hypotheses regarding relationships among these families have been proposed. One early noncladistic hypothesis considered the Sparidae to be intermediate between the more primitive Nemipteridae and the more derived Lethrinidae. A later nonformal phylogenetic treatment provided evidence for a close relationship between Sparidae and Centranthidae and suggested a closer affinity between the Nemipteridae and Lethrinidae. We examine 54 osteological, ligament, and squamation characters in representatives of all 45 genera of these families and 4 outgroup taxa. The results of our cladistic analysis are congruent with a cladistic interpretation of the earlier hypothesis, with strong support for the phyletic sequence Nemipteridae, Lethrinidae, Sparidae plus Centracanthidae, with placement of centracanthids unresolved with respect to sparid genera. Received: May 21, 2001 / Revised: October 26, 2001 / Accepted: November 19, 2001     

The skin surface of bony fishes

The cuticle described by light microscopists on the skin of various fishes, has been studied by electron microscopy in several species of teleost and in two other actinopterygian fish. Thelcuticle consists of an external coating layer, probably of mucopolysaccharide, which is secreted from the surface epidermal cells, not from the goblet mucous cells. The thickness of the cuticle is commonly of the order of 1 μm. It is particularly well developed in Trigla , where its thickness may vary from a fraction of a micron up to 50 μm, on different parts of the same individual. The cuticle has been detected in species from diverse orders, and is probably a normal constituent of the skin of all bony fishes. It is frequently lost during histological preparation. The external coat continues over the apertures of taste-buds, and may be continuous with the secretion at the mouths of chloride secreting cells. The secretion of the cuticle is partly from, or through, the outer membrane of the epidermal cells, but there is some evidence that cytoplasmic inclusions in the surface epidermal cells are also involved. There is striking variation in the appearance of these inclusions in electron micrographs of different species, and in some cases in different parts of the same fish, notably in Blennius. Certain of the inclusions are membrane-bounded vesicles whose contents are more electron-dense after staining with phosphotungstic acid than with lead citrate. In other cases, the inclusions are electron-transparent vacuoles. The cuticle is briefly compared with similar structures in other aquatic animals.     

Perspectives on the ecomorphology of bony fishes

Synopsis The field of ecomorphology has a long history with early roots in Europe. In this half of the century the application of ecomorphology to the biology of fishes has developed in the former Soviet Union, Poland and Czechoslovakia, The Netherlands, and in North America. While the specific approaches vary among countries, many North American studies begin by comparing morphological variation with variation in ecological characteristics at the intra or interspecific levels. These initial correlative studies form the ground work for hypotheses that explore the mechanistic underpinnings of the observed ecomorphological associations. Supporting these mechanistic hypotheses are insights from functional studies which demonstrate the limits to potential resource use resulting from a particular morphology; however, the actual resource use is likely to be more limited due to additional constraints provided by internal (e.g., behavior, physiology) and external (e.g., resource abundance, predator distribution) factors. The results from performance studies in the laboratory or field can be used to test specific ecomorphological hypotheses developed from the initial correlational and functional studies. Such studies may, but rarely do, incorporate an ontogenetic analysis of the ecomorphological association to determine their effect on performance. Finally, input from phylogenetic analyses allow an investigator to examine the evolution of specific features and to assess the rates and directionality of character evolution. The structural and ecological diversity of fishes provides a fertile ground to investigate these interactions. The contributions in this volume highlight some of the specific directions for ecomorphological research covering a variety of biological processes in fishes. These include foraging, locomotion, reproduction, respiration, and sensory systems. Running through these papers are new insights into universal ecomorphological issues, i.e., the relationships between form and ecological role and the factors that modify these relationships.     

The phylogeny of parental care in fishes

This paper tests the hypothesis that in the evolution of parental care, taxa of bony fish should only exhibit certain transitional states (where a transition is defined by the occurrence of at least two types of parental care within a genus or family). These are those between no parental care and male care, male care and biparental care, biparental care and female care, and female care and no parental care. A review of the teleost literature reveals 21 transitions. All of these agree with the hypothesized transitions and, in some cases, the direction of evolution is inferred by simple pedigree analysis.     

Characters of the female gonad and the phylogeny of Platyhelminthes

The female gonad of Platyhelminthes shows a wide variability in its anatomical and cellular organization and submicroscopic structure, and some characteristics may be useful for elucidating the questioned monophyly of some taxa and reconstructing the phylogenetic tree of the group. The morphological characters of the female gonad have been subdivided into three levels: macroscopic, microscopic and ultrastructural, and some examples with potential phylogenetic value are presented for each level. This revised version was published online in July 2006 with corrections to the Cover Date.     

Molecular phylogeny of Mugilidae fishes revised

Systematics derived from morphological characters often does not correspond with the evolutionary processes underlying the divergence within a group of organisms. In the family Mugilidae (Teleostei) morphological similarities have resulted in inconsistencies between taxonomy and phylogeny among its species, and particularly for the genera Mugil, Liza and Chelon where both intrageneric and intergeneric phylogenetic clarifications are needed. To address these issues, the direct sequencing of the mitochondrial region that encodes Phenylalanine (69 bp), 12S rRNA (842 bp), cytochrome c oxidase subunit I (651 bp) and cytochrome b (702 bp) was carried out. The data reveal that Mugil platanus and Mugil liza represent a continuum of a single species, closely related to but distinct from Mugil cephalus which itself appears to comprise a grouping of multiple and closely related species. This species complex was genetically distinct from Mugil curema, which, based on three clearly diverged species identified in this study along the Atlantic coast of the Americas, requires extensive taxonomic revision throughout its world-wide distribution. Unlike the monophyly supported within Mugil, relationships within Liza are paraphyletic, and a taxonomic revision of the genera Liza, Chelon and Oedalechilus is needed.     

A bibliography and categorization of bony fishes exhibiting parental care

Parental care occurs in a diversity of fishes, but predominantly among freshwater groups. Among the approximately 422 families of bony fishes (Osteichthyes), 89 are presently known to exhibit parental care. Grouping these families into eight categories, based on the sex of the care-giver(s), reveals male parental care is as common or more common than female parental care. Although unusual among vertebrates, parental care by males alone is very common among bony fishes. Lists of families, the forms of parental care exhibited, the modes of fertilization, the environments in which reproduction occurs, and the sources of documentation are presented. An extensive bibliography and index are provided.     

Ontogeny and phylogeny of tooth attachment modes in actinopterygian fishes

There are four major tooth attachment modes in actinopterygians. Type 1 mode is characterized by complete ankylosis of the tooth to the attachment bone; it is the primitive attachment mode for actinopterygians. In Type 2 mode there is a ring of collagen between the tooth base and the bone. In Type 3 mode mineralization extends near or to the bone at the anterior tooth border, and there is a relatively large collagen area on the posterior surface of the tooth; Type 3 teeth are hinged with an anterior axis of rotation. Type 4 teeth also have a relatively large posterior collagen area, but there is no collagenous connection between the anterior basal tooth border and the attachment bone; Type 4 teeth are hinged, with a posterior axis of rotation. Types 2, 3, and 4 attachment modes appear to result from retardation of mineralization and resemble, with some modifications, ontogenetic stages in the development of Type 1 mode; they are considered to be paedomorphic features. Attachment modes 2, 3, and 4 are each associated with a major evolutionary lineage within the Teleostei. The degree to which paedomorphosis has been a factor in teleostean evolution is discussed.     

Evolution of asynchronous motor activity in paired muscles: effects of ecology, morphology, and phylogeny

Many studies of feeding behavior have implanted electrodes unilaterally(in muscles on only one side of the head) to determine the basicmotor patterns of muscles controlling the jaws. However, bilateralimplantation has the potential to achieve a more comprehensiveunderstanding of modification of the motor activity that maybe occurring between the left and right sides of the head. Inparticular, complex processing of prey is often characterizedby bilaterally asynchronous and even unilateral activation ofthe jaw musculature. In this study, we bilaterally implant feedingmuscles in species from four orders of elasmobranchs (Squaliformes,Orectolobiformes, Carcharhiniformes, Rajoidea) in order to characterizethe effects of type of prey, feeding behavior, and phylogenyon the degree of asynchronous muscle activation. Electrodeswere implanted in three of the jaw adductors, two divisionsof the quadratomandibularis and the preorbitalis, as well asin a cranial elevator in sharks, the epaxialis. The asynchronyof feeding events (measured as the degree to which activityof members of a muscle pair is out of phase) was compared acrossspecies for capture versus processing and simple versus complexprey, then interpreted in the contexts of phylogeny, morphology,and ecology to clarify determinants of asynchronous activity.Whereas capture and processing of prey were characterized bystatistically similar degrees of asynchrony for data pooledacross species, events involving complex prey were more asynchronousthan were those involving simple prey. The two trophic generalists,Squalus acanthias and Leucoraja erinacea, modulated the degreeof asynchrony according to type of prey, whereas the two behavioralspecialists, Chiloscyllium plagiosum and Mustelus canis, activatedthe cranial muscles synchronously regardless of type of prey.These differences in jaw muscle activity would not have beendetected with unilateral implantation. Therefore, we advocatebilateral implantation in studies of cranial muscle functionin fishes, particularly when investigating behaviors associatedwith processing complex prey. Incorporating this methodologywill provide a more detailed understanding of the coordinationand evolution of paired-muscle function in the feeding apparatusrelative to behavioral and ecological performance.     

Molecules,morphology, and phylogeny

A complicated, almost conflicting situation exists in contemporary biology, i.e., there are two scientific concepts in existence that solve some phylogenetic problems in a different way. These concepts are evolutionary morphology and molecular biology. Modern biology urgently needs these contradictions to be examined and removed.     

Evolution of ecological traits and wing morphology in Hemileuca (Saturniidae) based on a two-gene phylogeny

We present a molecular phylogeny for the genus Hemileuca (Saturniidae), based on 624 bp of mitochondrial cytochrome oxidase I (COI) and 932 bp of the nuclear gene elongation factor 1 alpha (EF1alpha). Combined analysis of both gene sequences increased resolution and supported most of the phylogenetic relationships suggested by separate analysis of each gene. However, a maximum parsimony (MP) model for just COI sequence from one sample of most taxa produced a phylogeny incongruent with EF1alpha and combined dataset analyses under either MP or ML models. Time of year and time of day during which adult moths fly corresponded strongly with the phylogeny. Although most Hemileuca are diurnal, ancestral Hemileuca probably were nocturnal, fall-flying insects. The two-gene molecular phylogeny suggests that wing morphology is frequently homoplastic. There was no correlation between the primary larval hostplants and phylogenetic placement of taxa. No phylogenetic pattern of specialization was evident for single hostplant families across the genus. Our results suggest that phenological behavioral characters may be more conserved than the wing morphology characters that are more commonly used to infer phylogenetic relationships in Lepidoptera. Inclusion of a molecular component in the re-evaluation of systematic data is likely to alter prior assumptions of phylogenetic relationships in groups where such potentially homoplastic characters have been used.     

Evolution and consequences of endothermy in fishes

Regional endothermy, the conservation of metabolic heat by vascular countercurrent heat exchangers to elevate the temperature of the slow-twitch locomotor muscle, eyes and brain, or viscera, has evolved independently among several fish lineages, including lamnid sharks, billfishes, and tunas. All are large, active, pelagic species with high energy demands that undertake long-distance migrations and move vertically within the water column, thereby encountering a range of water temperatures. After summarizing the occurrence of endothermy among fishes, the evidence for two hypothesized advantages of endothermy in fishes, thermal niche expansion and enhancement of aerobic swimming performance, is analyzed using phylogenetic comparisons between endothermic fishes and their ectothermic relatives. Thermal niche expansion is supported by mapping endothermic characters onto phylogenies and by combining information about the thermal niche of extant species, the fossil record, and paleoceanographic conditions during the time that endothermic fishes radiated. However, it is difficult to show that endothermy was required for niche expansion, and adaptations other than endothermy are necessary for repeated diving below the thermocline. Although the convergent evolution of the ability to elevate slow-twitch, oxidative locomotor muscle temperatures suggests a selective advantage for that trait, comparisons of tunas and their ectothermic sister species (mackerels and bonitos) provide no direct support of the hypothesis that endothermy results in increased aerobic swimming speeds, slow-oxidative muscle power, or energetic efficiency. Endothermy is associated with higher standard metabolic rates, which may result from high aerobic capacities required by these high-performance fishes to conduct many aerobic activities simultaneously. A high standard metabolic rate indicates that the benefits of endothermy may be offset by significant energetic costs.     

Ventricle morphology in pelagic elasmobranch fishes

Ventricle weights of the warm-bodied great white shark, Atlantic shortfin mako, and the common thresher shark (the latter presumed to be warm-bodied) are similar to those of ectothermic blue sharks, sandbar sharks, dusky sharks, tiger sharks and scalloped hammerhead sharks. Ventricle muscularity, as estimated by the ratio of cortical to spongy layer thickness, is almost twice as great in the former three species than in the latter elasmobranchs. Measurements of ventricular volumes suggest that the ventricles of the great white, Atlantic shortfin mako and common thresher sharks are better adapted to respond to demands for increases in cardiac output via increased heartbeat frequency in comparison with ectothermic species of shark.