Morphology and Interrelationships of Primitive Actinopterygian Fishes

SYNOPSIS. The concept of the Actinopterygii as a natural groupof fishes was not generally accepted until early in this century.Ever since, the characterization of the group has been blurredby the problem of cladistian (polypterid) relationships. Froma review of the structure of polypterids and actinopts, it isconcluded that Cladistia are the sistergroup of Recent actinopterygians(Actinopteri), the two groups together comprising the Actinopterygii.Recent chondrosteans are more closely related to higher actinopts(Neopterygii) than to cladistians. The extinct Palaeonisciformesappear to be a paraphyletic group, comprising stem-group actinopterygians(e.g., Cheirolepts), stem-group actinopterans (e.g., Moythomasia)and relatives of higher actinopterans (e.g., Pteroniscus)     

Locomotor Patterns in the Evolution of Actinopterygian Fishes

SYNOPSIS. Locomotor adaptations in actinopterygian fishes aredescribed for (a) caudal propulsion, used in cruising and sprintswimming, acceleration, and fast turns and (b) median and pairedfin propulsion used for slow swimming and in precise maneuver.Caudal swimming is subdivided into steady (time independent)and unsteady (time dependent acceleration and turning) locomotion. High power caudal propulsion is the major theme in actinopterygianswimming morphology because of its role in predator evasionand food capture. Non-caudal slow swimming appears to be secondaryand is not exploited before the Acanthopterygii. Optimal morphological requirements for unsteady swimming are(a) large caudal fin and general body area, (b) deep caudalpeduncle, often enhanced by posterior dorsal and anal fins,(c) an anterior stabilizing body mass and\or added mass, (d)flexible body and (e) large ratio of muscle mass to body mass.Optimal morphological requirements for steady swimming are (a)high aspect ratio caudal fin, (b) narrow caudal peduncle, (c)small total caudal area, (d) anterior stabilizing body massand added mass, and (e) a stiff body. Small changes in morphologycan have large effects on performance. Exclusive morphological requirements for steady versus unsteadyswimming are partially overcome using collapsible fins, butcompromises remain necessary. Morphologies favoring unsteadyperformance are a recurring theme in actinopterygian evolution.Successive radiations at chondrostean, halecostome and teleosteanlevels are associated with modifications in the axial and caudalskeleton. Strength of ossified structures probably limited maximum propulsionforces early in actinopterygian evolution, so that specializationsfor fast cruising (carangiform and thunmform modes) followedstructural advances especially in the caudal skeleton. No suchlimits apply to eel-like forms which consequently recur in successiveactinopterygian radiations. Slow swimming using mainly non-caudal propulsion probably firstoccurred among neopterygians in association with reduced andneutral buoyancy. Slow swimming adaptations can add to and extendthe scope of caudal swimming, but specialization is associatedwith reduced caudal swimming performance. Marked exploitationof slow swimming opportunities does not occur prior to the anterodorsallocation of pectoral and pelvic girdles and the vertical rotationof the base of the pectoral fin, as found in the Acanthopterygii.     

Patterns of Evolution in the Feeding Mechanism of Actinopterygian Fishes

SYNOPSIS. Structural and functional patterns in the evolutionof the actinopterygian feeding mechanism are discussed in thecontext of the major monophyletic lineages of ray-finned fishes.A tripartite adductor mandibulae contained in a maxillary-palatoquadratechamber and a single mechanism of mandibular depression mediatedby the obliquus inferioris, sternohyoideus, and hyoid apparatusare primitive features of the Actinopterygii. Halecostome fishesare characterized by having an additional mechanism of mandibulardepression, the levator operculi—opercular series coupling,and a maxilla which swings anteriorly during prey capture. Theseinnovations provide the basis for feeding by inertial suctionwhich is the dominant mode of prey capture throughout the halecostomeradiation. A remarkably consistent kinematic profile occursin all suction-feeding halecostomes. Teleost fishes possessa number of specializations in the front jaws including a geniohyoideusmuscle, loss of the primitive suborbital adductor component,and a mobile premaxilla. Structural innovations in teleost pharyngealjaws include fusion of the dermal tooth plates with endoskeletalgill arch elements, the occurrence of a pharyngeal retractormuscle, and a shift in the origin of the pharyngohyoideus. Thesespecializations relate to increased functional versatility ofthe pharyngeal jaw apparatus as demonstrated by an electromyographicstudy of pharyngeal muscle activity in Esox and Ambloplites.The major feature of the evolution of the actinopterygian feedingmechanism is the increase in structural complexity in both thepharyngeal and front jaws. Structural diversification is a functionof the number of independent biomechanical pathways governingmovement.     

Symposium Summary: Evolutionary Patterns in Actinopterygian Fishes

SYNOPSIS. The actinopterygian fishes are an exemplary cladefor the study of structural and functional evolutionary patterns.With over half of all vertebrate species, ray-finned fisheshave diversified into a wide variety of habitats, and considerableprogress has been made over the last fifteen years in understandingthe genealogical relationships of actinopterygians. This symposiumhas contributed to our understanding of phylogenetic patternsin actinopterygians and to knowledge of the major structuraland functional patterns in locomotor, auditory, trophic, andneural systems. A number of key areas for future research havebeen identified. (1) The relationships of "palaeonisciform"fishes, (2) the study of trends in feeding and locomotor systemswithin a phylogenetic context, (3) the identification of primitivepatterns of pharyngeal jaw movement and steady and unsteadylocomotor patterns in actinopterygians, (4) the homologies,identification, and functional significance of neural pathwaysin the telencephalon, and (5) the comparative study of form-functionrelations in the auditory system. The study of teleost fishbiology has proceeded at the expense of data on primitive actinopterygians(e.g., Polypterus, Polyodon, Aapenser, Lepisosteus, Amia) whichare especially important in the analysis of structural and functionalpatterns in ray-finned fishes.     

An Overview of the Organization of the Brain of Actinopterygian Fishes

SYNOPSIS. The brain of actinopterygian fishes can be subdividedinto five principal parts, rhombencephalon, cerebellum, mesencephalon,diencephalon and telencephalon, each of which contains a numberof separate morphological entities: nuclei, areas or zones.Analysis of the origin and termination of the cranial nervesand their components reveals that many of the morphologicalentities distinguished in the actinopterygian brain can be interpretedin terms of elementary sensory and motor functions. Experimentalanatomical and physiological studies on the fiber connectionsof the entities thus defined have led to a functional interpretationof many other parts of the brain. Thus, the central circuitryrelated to such sensory functions as hydrodynamic pressoreception,electroreception, vision, gustation and olfaction are well-known,and the same holds true for the motor systems related to feedingand locomotion. However, there are still many regions in theactinopterygian brain the functional significance of which ispoorly understood, and it should be emphasized that most ofour knowledge on the organization of the brain of this grouprests merely on observations in teleosts. One of the most interestingaspects of actinopterygian neurobiology is that the telencephalonin this group develops in a way which differs fundamentallyfrom that found in all other craniates, and that the telencephalonshows a marked progressive differentiation in the series: polypterids—chondrosteans—halecomorphs—teleosts.     

Microstructure and Growth of the Dermal Skeleton in Fossil Actinopterygian Fishes: Birgeria and Scanilepis

The dermal bones of Birgeria and Scanilepis contain numerous odontodes of consecutive generations, each consisting of dentine and a single ganoin layer; superimposition of series of ganoin layers, generally encountered in the scales of the palaeonisciforms. does not occur in any of these bones. In Birgeria , some odontodes near to the dentition resemble jaw-teeth proper in possessing an apical portion of acrodin; furthermore, nothing indicates the existence here of nerve-sac groups like those of sturgeons. The histology of the dermal skeleton in Scanilepis does not support the assumption that this form is more closely related to Polypterus than other palaeonisciforms. Remarks are given on the structure of acrodin and ganoin as revealed by SEM study.     

Microstructure and Growth of the Dermal Skeleton in Fossil Actinopterygian Fishes: Boreosomus, Plegmolepis and Gyrolepis

The odontodes of some of the palatal dermal bones in Boreosomus piveteaui Nielsen tend to form odontocomplexes, e.g. symmetrical areal ones where some degree of superimposition may occur between ganoin layers belonging to the component odontodes, and asymmetrical areal ones where this is quite insignificant. In the areal odontocomplexes of the dermal bones in Plegmolepis sp., the degree of overlap between the corresponding layers is somewhat more pronounced. Finally, in the areal odontocomplexes of the dermal bones in Gyrolepis cf. albertii Agassiz, we have a more advanced stage of phyletic specialization where each of the ganoin layers of the component odontodes lies directly superimposed on the preceding one throughout the extent of the latter. At the same time, the effect of phyletic dentine reduction is clearly noticeable here by the development of "extra" ganoin layers no more retaining their original connections with that hard tissue (also developed in the scales of Plegmolepis sp.). Remarks are i.a. given on the characters by which Acropholis and Plegmolepis are said to be distinguishable from each other.     

Evolution of the branchiostegal membrane and restricted gill openings in Actinopterygian fishes

A phylogenetic survey is a powerful approach for investigating the evolutionary history of a morphological characteristic that has evolved numerous times without obvious functional implications. Restricted gill openings, an extreme modification of the branchiostegal membrane, are an example of such a characteristic. We examine the evolution of branchiostegal membrane morphology and highlight convergent evolution of restricted gill openings. We surveyed specimens from 433 families of actinopterygians for branchiostegal membrane morphology and measured head and body dimensions. We inferred a relaxed molecular clock phylogeny with branch length estimates based on nine nuclear genes sampled from 285 species that include all major lineages of Actinopterygii. We calculated marginal state reconstructions of four branchiostegal membrane conditions and found that restricted gill openings have evolved independently in at least 11 major actinopterygian clades, and the total number of independent origins of the trait is likely much higher. A principal component analysis revealed that fishes with restricted gill openings occupy a larger morphospace, as defined by our linear measurements, than do fishes with nonrestricted openings. We used a decision tree analysis of ecological data to determine if restricted gill openings are linked to certain environments. We found that fishes with restricted gill openings repeatedly occur under a variety of ecological conditions, although they are rare in open‐ocean pelagic environments. We also tested seven ratios for their utility in distinguishing between fishes with and without restricted gill openings, and we propose a simple metric for quantifying restricted gill openings (RGO), defined as a ratio of the distance from the ventral midline to the gill opening relative to half the circumference of the head. Functional explanations for this specialized morphology likely differ within each clade, but its repeated evolution indicates a need for a better understanding of diversity of ventilatory morphology among fishes. J. Morphol. 276:681–694, 2015. © 2015 Wiley Periodicals, Inc.     

Functional Morphology of the Heart in Fishes

The systemic heart of fishes consists of four chambers in series,the sinus venosus, atrium, ventricle, and conus or bulbus. Valvesbetween the chambers and contraction of all chambers exceptthe bulbus maintain a unidirectional blood flow through theheart. The heart is composed of typical vertebrate cardiac muscle,although there may be minor differences in the distributionof spontaneously active cells, the rate and nature of spreadof excitatory waves, and the characteristics of resting andaction potentials between different fish and other vertebrates.Cholinergic fibers innervate the heart, except in hagfish whichhave aneural hearts. Fish hearts lack sympathetic innervation.The level of vagal tone varies considerably, and is affectedby many factors. In some fish the heart is essentially aneural(without vagal tone) during exercise and may resemble an isolatedmammalian ventricle with increased venous return causing increasedcardiac output. There are many mechanisms that could increasevenous return in exercising fish. rß-adrenergic receptorshave been located on the hearts of some fish, and changing levelsof catecholamines may play a role in regulating cardiac activity.Changes in cardiac output in fish are normally associated withlarge changes in stroke volume and small cha-nges in heart rate.     

The Evolution of Thyroidal Function in Fishes

Although the thyroid gland evolved from the gut, there is noevidence that thyroxine functions as part of the gastro-intestinalendocrine system nor does it have any major function analogousto the control of glucose by the pancreatic islets. The controlof the thyroid evolved from the pituitary control of the gonadsuggesting that an early role of thyroxine was in reproduction.This idea is supported by the presence of cycles of thyroidactivity associated with reproduction in both elasmobranchsand teleosts. In teleosts thyroxine is necessary for gonadalmaturation. The numerous other effects of thyroxine in teleostsmay have evolved from this maturational effect or have beenadded to it during the course of teleost evolution.     

The Physiology and Evolution of Urea Transport in Fishes

This review summarizes what is currently known about urea transporters in fishes in the context of their physiology and evolution within the vertebrates. The existence of urea transporters has been investigated in red blood cells and hepatocytes of fish as well as in renal and branchial cells. Little is known about urea transport in red blood cells and hepatocytes, in fact, urea transporters are not believed to be present in the erythrocytes of elasmobranchs nor in teleost fish. What little physiological evidence there is for urea transport across fish hepatocytes is not supported by molecular evidence and could be explained by other transporters. In contrast, early findings on elasmobranch renal urea transporters were the impetus for research in other organisms. Urea transport in both the elasmobranch kidney and gill functions to retain urea within the animal against a massive concentration gradient with the environment. Information on branchial and renal urea transporters in teleost fish is recent in comparison but in teleosts urea transporters appear to function for excretion and not retention as in elasmobranchs. The presence of urea transporters in fish that produce a copious amount of urea, such as elasmobranchs and ureotelic teleosts, is reasonable. However, the existence of urea transporters in ammoniotelic fish is curious and could likely be due to their ability to manufacture urea early in life as a means to avoid ammonia toxicity. It is believed that the facilitated diffusion urea transporter (UT) gene family has undergone major evolutionary changes, likely in association with the role of urea transport in the evolution of terrestriality in the vertebrates.     

Morphology of the Hagfish Inner Ear

The inner ear of five species of hagfishes was examined with different light and electron microscopical techniques. In all species, the labyrinth contains a single macula and two cristae, in a single semicircular canal. The macula consists of a horizontal, a middle vertical and a posterior horizontal component. Each component is covered by numerous round statoconia. The ring-shaped cristae have very long kinocilia, but lack a proper cupula. The sensory epithelia show signs of regeneration, indicated by the presence of mitoses and apoptotic hair cells.     

The Evolution of Male and Female Parental Care in Fishes

In this paper we propose an explanation for (a) the predominanceof male care in fishes, and (b) the phylogenies and transitionsthat occur among care states. We also provide a general evolutionarymodel for studying the conditions under which parental careevolves. Our conclusions are as follows: (i) Parental care hasonly one benefit, the increased survivorship of young. It may,however, have three costs: a "mating cost," an "adult survivorshipcost," and a "future fertility cost." (ii) On average, malesand females will derive the same benefit from care. They probablyalso pay the same adult survivorship cost. However, their matingcost and future fertility costs may differ, (iii) A mating costusually applies only to males. However, this cost may be reducedby male territoriality and, in some situations, be entirelyremoved. Under this condition, natural selection on presentreproductive success is equivalent for males and females, (iv)When fecundity accelerates with body size in females, whilemale mating success follows a linear relationship with bodysize, future fertility costs of parental care are greater forfemales than males. Although further tests are needed, a preliminaryanalysis suggests this often may be the case in fishes. Thus,the predominance of male parental care in fishes is not explainedby males deriving greater benefits from care, but by males payingsmaller future costs. Males thus accrue a greater net fitnessadvantage from parental care (see expressions [6] and [12]).(v) The evolution of biparental care from uniparental male caremay occur because male care selects for larger egg sizes andincreased embryo investment by females. This increases the benefitto the female of parental care, (vi) By contrast, uniparentalfemale care may originate from biparental care when males areselected to desert. This occurs when female care creates a matingcost to males. In some cases male desertion may "lock" femalesinto uniparental care. However, in many other cases femalesmay be selected to desert, giving rise to "no care." (vii) Theorigin of uniparental female care from no care is rare in externallyfertilizing fishes. This is because the benefits of care rarelyoutweigh a female's future fertility costs (expression [9]).For internally fertilizing species, however, the benefit ofcare is high whereas the cost is probably low. Most of thesespecies have evolved embryo retention, (viii) When parentalcare begins with male care and moves to biparental care, ouranalysis suggests that care evolution will include cyclicaldynamics. Parental care in some fishes may thus be seen as transitionaland changing through evolutionary time rather than as an evolutionarilystable state. In theory, "no care" may be a phylogeneticallyadvanced state.     

Evolution of the Vertebrate Cytosolic Malate Dehydrogenase Gene Family: Duplication and Divergence in Actinopterygian Fish

Abstract A general correlation between neural expression and negative charge in isozymes suggests charge represents an adaptation to the neural environment. Interestingly, a notable exception exists in teleost fish. Two cytosolic malate dehydrogenase (MDH) isozymes have different spatial expression patterns in certain fishes: one is expressed in all tissues and the second is expressed primarily in the eye and skeletal muscle. While the neural MDH isozyme is negatively charged, the difference in charge between the two isozymes is not as pronounced as that observed in other gene families (e.g., triosephosphate isomerase and lactate dehydrogenase). Most tetrapods express a single cytosolic MDH isozyme, and it has been demonstrated recently that the pair of isozymes found in teleosts results from a gene duplication sometime after the separation of teleosts and tetrapods, although the exact timing of this duplication has not been inferred. Phylogenetic analyses suggest that the duplication of teleost isozymes occurred during the radiation of actinopterygian fish, consistent with the timing of duplication at other loci. Using inferred amino acid sequences, we examine the pattern of change following the duplication and across the rest of the MDH gene tree. Comparison between the MDH gene family and another gene family that shows a larger charge differential among members (triosephosphate isomerase) indicates that the smaller charge difference between MDH isozymes is best explained by greater constraint on amino acid change directly following the duplication, not greater constraint across the entire gene tree. This difference in constraint might result from the wider pattern of expression of the “neural” MDH isozyme.     

The Evolution of the Tetrapod Middle Ear in the Rhipidistian-Amphibian Transition

From a survey ot the structure of the skull in rhipidistianfishes and early labylinthodont Amphibia and of the mechanismof hearing in these two groups, an account of the evolutionof the tetrapod middle ear is presented. The overall modificationof the otic region of the skull during the rhipidistian-amphibiantransition is analyzed in terms of changes in different organsystems in response to different selective pressures (affecting,for example, the feeding, respiratory, and locomotory mechanisms).These changes are seen to occur in a completely integrated pattern.Considerations of the different requirements for sound receptionunder water and in air, in connection with this correlated progressionof evolutionary change in the otic region of the head, revealthe manner in which the hyomandibular, spiracular diverticulum,and operculum of rhipidistian fishes became modified to formthe stapes, the tympanic cavity, and the outer portion of thetympanum, respectively, of tetrapods.     

Diversity and Evolution of Body Size in Fishes

The diversity of body sizes observed among species of a clade is a combined result of microevolutionary processes (i.e. natural selection and genetic drift) that cause size changes within phylogenetic lineages, and macroevolutionary processes (i.e. speciation and extinction) that affect net rates of diversification among lineages. Here we assess trends of size diversity and evolution in fishes (non-tetrapod craniates), employing paleontological, macroecological, and phylogenetic information. Fishes are well suited to studies of size diversity and evolution, as they are highly diverse, representing more than 50% of all living vertebrate species, and many fish taxa are well represented in the fossil record from throughout the Phanerozoic. Further, the frequency distributions of sizes among fish lineages resemble those of most other animal taxa, in being right-skewed, even on a log scale. Using an approach that measures rates of size evolution (in darwins) within a formal phylogenetic framework, we interpret the shape of size distributions as a balance between the competing forces of diversification, pushing taxa away from ancestral values, and of conservation, drawing taxa closer to a central tendency. Within this context we show how non-directional mechanisms of evolution (i.e. passive diffusion processes) can produce an hitherto unperceived bias to larger size, when size is measured on the conventional log scale. These results demonstrate how the interpretation of macroecological datasets can be enriched from an historical perspective, and document the ways in which macroevolutionary and microevolutionary processes may be decoupled in the production of size diversity.     

Diet and Diversification in the Evolution of Coral Reef Fishes

The disparity in species richness among evolutionary lineages is one of the oldest and most intriguing issues in evolutionary biology. Although geographical factors have been traditionally thought to promote speciation, recent studies have underscored the importance of ecological interactions as one of the main drivers of diversification. Here, we test if differences in species richness of closely related lineages match predictions based on the concept of density-dependent diversification. As radiation progresses, ecological niche-space would become increasingly saturated, resulting in fewer opportunities for speciation. To assess this hypothesis, we tested whether reef fish niche shifts toward usage of low-quality food resources (i.e. relatively low energy/protein per unit mass), such as algae, detritus, sponges and corals are accompanied by rapid net diversification. Using available molecular information, we reconstructed phylogenies of four major reef fish clades (Acanthuroidei, Chaetodontidae, Labridae and Pomacentridae) to estimate the timing of radiations of their subclades. We found that the evolution of species-rich clades was associated with a switch to low quality food in three of the four clades analyzed, which is consistent with a density-dependent model of diversification. We suggest that ecological opportunity may play an important role in understanding the diversification of reef-fish lineages.     

Evolution and Diversification of Antarctic Notothenioid Fishes

Antarctica supported fossil ichthyofaunas during the Devonian,Jurassic, Cretaceous and Eocene/Oligocene. These faunas arenot ancestral to each other, nor are they related to any componentof the modern fauna. About one hundred species of notothenioidsdominate a modern fauna of over 200 species of bottom fishes.This highly endemic perciform suborder is not representedinthe fossil record of Antarctica. Notothenioids may have evolvedin situ on the margins of the Antarctic continent while graduallyadapting to cooling conditions during the Tertiary. Cladisticstudies indicate that notothenioids are a monophyletic group,but a sister group has not been identified among perciform fishes.With relatively few non-notothenioid fishes in Antarctic waters,notothenioids fill ecological roles normally occupied by taxonomicallydiverse fishes in temperate waters. There are six notothenioidfamilies: Bovichtidae, Nototheniidae, Harpagiferidae, Artedidraconidae,Bathydraconidae and Channichthyidae. Aspects of theirbiologyare briefly considered with emphasis on the Nototheniidae, themost speciose family. Evolutionary diversification within thisfamily allows recognition of species which are pelagic, cryopelagic,benthopelagic and benthic.