Hox genes control homocercal caudal fin development and evolution.

Ancient bony fishes had heterocercal tails, like modern sharks and sturgeons, with asymmetric caudal fins and a vertebral column extending into an elongated upper lobe. Teleost fishes, in contrast, developed a homocercal tail characterized by two separate equal-sized fin lobes and the body axis not extending into the caudal fin. A similar heterocercal-to-homocercal transition occurs during teleost ontogeny, although the underlying genetic and developmental mechanisms for either transition remain unresolved. Here, we investigated the role of hox13 genes in caudal fin formation as these genes control posterior identity in animals. Analysis of expression profiles of zebrafish hox13 paralogs and phenotypes of CRISPR/Cas9-induced mutants showed that double hoxb13a and hoxc13a mutants fail to form a caudal fin. Furthermore, single mutants display heterocercal-like morphologies not seen since Mesozoic fossil teleosteomorphs. Relaxation of functional constraints after the teleost genome duplication may have allowed hox13 duplicates to neo- or subfunctionalize, ultimately contributing to the evolution of a homocercal tail in teleost fishes.

Phylogenetic relationships, origin and historical biogeography of the genus Sprattus (Clupeiformes: Clupeidae).

The genus Sprattus comprises five species of marine pelagic fishes distributed worldwide in antitropical, temperate waters. Their distribution suggests an ancient origin during a cold period of the earth's history. In this study, we evaluated this hypothesis and corroborated the non-monophyly of the genus Sprattus, using a phylogenetic approach based on DNA sequences of five mitochondrial genome regions. Sprattus sprattus is more closely related to members of the genus Clupea than to other Sprattus species. We also investigated the historical biogeography of the genus, with the phylogenetic tree showing two well-supported clades corresponding to the species distribution in each hemisphere. Time-calibrated phylogenetic analyses showed that an ancient divergence between Northern and Southern Hemispheres occurred at 55.8 MYBP, followed by a diversification in the Oligocene epoch in the Northern Hemisphere clade (33.8 MYBP) and a more recent diversification in the Southern Hemisphere clade (34.2 MYBP). Historical biogeography analyses indicated that the most recent common ancestor (MRCA) likely inhabited the Atlantic Ocean in the Southern Hemisphere. These results suggest that the ancestral population of the MRCA diverged in two populations, one was dispersed to the Northern Hemisphere and the other across the Southern Hemisphere. Given that the Eocene was the warmest epoch since the Paleogene, the ancestral populations would have crossed the tropics through deeper cooler waters, as proposed by the isothermal submergence hypothesis. The non-monophyly confirmed for the genus Sprattus indicates that its systematics should be re-evaluated.

From Devo to Evo: patterning, fusion and evolution of the zebrafish terminal vertebra.

BACKGROUND: With more than 30,000 species, teleosts comprise about half of today's living vertebrates, enriched with a wide set of adaptations to all aquatic systems. Their evolution was marked by modifications of their tail, that involved major rearrangements of the metameric organization of the axial skeleton. The most posterior or ural caudal skeleton, primitively included more than 10 vertebrae and, through a series of fusions and losses, became reduced to a single vertebra in modern ostariophysans, one of the largest clades of teleosts. The ontogeny of the ostariophysan Danio rerio recapitulates this process by forming two or three separate vertebrae that become a single vertebra in adults. We characterize the developmental sequence of this change by describing the processes of patterning, fusion and differential growth on each of the constitutive elements that sculpt the adult terminal vertebra. RESULTS: The ontogenetic changes of the terminal vertebra were characterized, highlighting their shared and derived characters in comparison with other teleosts. In zebrafish, there is: i) a loss of the preural centrum 1, ii) the formation of an hourglass-shaped autocentrum only in the anterior but not the posterior border of the compound centrum, iii) the formation of a vestigial posterior centrum that does not form an autocentrum and becomes incorporated beneath the compound centrum during development, and iv) the elongated dorso-posterior process of the compound centrum or pleurostyle appears as an independent element posterior to the compound centrum, before fusing to the ural neural arches and the anterior portion of the compound centrum. CONCLUSIONS: The unique features of the formation of the terminal vertebra in Danio rerio reflect the remarkable changes that occurred during the evolution of teleosts, with potential shared derived characteristics for some of the major lineages of modern teleosts. A new ontogenetic model is proposed to illustrate the development of the terminal vertebra, and the phylogenetic implications for the evolution of caudal skeleton consolidation in ostariophysans are discussed.

Notochordal Signals Establish Phylogenetic Identity of the Teleost Spine.

The spine is a defining feature of the vertebrate body plan. However, broad differences in vertebral structures and morphogenetic strategies occur across vertebrate groups, clouding the homology between their developmental programs. Analysis of a zebrafish mutant, spondo, whose spine is dysmorphic, prompted us to reconstruct paleontological evidence, highlighting specific transitions during teleost spine evolution. Interestingly, the spondo mutant recapitulates characteristics present in basal fishes, not found in extant teleosts. Further analysis of the mutation implicated the teleost-specific notochord protein, Calymmin, as a key regulator of spine patterning in zebrafish. The mutation in cmn results in loss of notochord sheath segmentation, altering osteoblast migration to the developing spine, and increasing sensitivity to somitogenesis defects associated with congenital scoliosis in amniotes. These data suggest that signals from the notochord define the evolutionary identity of the spine and demonstrate how simple shifts in development can revert traits canalized for about 250 million years.

The South American and Australian percichthyids and perciliids. What is new about them?

A study including morphological characters and mitogenomics of South American and Australian fishes previously assigned to Percichthyidae was conducted. Results generated from these different data sets reveal major disagreements concerning the content and interpretation of the so-called percichthyids. A phylogenetic analysis based on 54 morphological characters suggests the existence of two major clades: (1) Percichthyidae, including the South American Percichthys and the Australian taxa Macquaria australasica, Macquaria (= Plectroplites), and Maccullochella; (2) Perciliidae with the South American genus Percilia at the base plus more advanced Australian genera Nannoperca, Nannatherina, Bostockia, and Gadopsis. In contrast, molecular and mitogenomic evidence suggests only one clade (Percichthyidae), with the exclusion of species of Macquaria (= Percalates). Additionally, the results reveal the existence of various taxonomic problems, such as the current interpretation of only one species of Percichthys in Argentina, an interpretation that is not supported by the present study; the existence of cryptic species of Percilia as well as of Gadopsis, Nannoperca, and Macquaria that will increase the diversity of the genera; and the need for an extensive revision of species previously assigned to Percalates versus Macquaria. Disagreements point to the need to develop further research on the so-called percichthyids and perciliids.(AU)

Se realizó un estudio de peces sudamericanos y australianos incluyendo caracteres morfológicos y mitogenómicos, para taxa previamente asignados a la familia Percichthyidae. Los resultados generados de estos conjuntos de datos diferentes revelaron desacuerdos importantes entre el contenido y la interpretación de los así llamados percíctidos. Un análisis filogenético basado en 54 caracteres morfológicos sugiere la existencia de dos clados importantes: (1) La familia Percichthyidae, incluyendo el género sudamericano Percichthys y los taxa australianos Macquaria australasica, Macquaria (= Plectroplites) y Maccullochella. (2) Perciliidae con el género sudamericano Percilia en la base, y géneros australianos más avanzados como Nannoperca, Nannatherina, Bostockia y Gadopsis. En contraste, la evidencia molecular y mitogenómica incluye la mayor parte de los géneros dentro de la familia Percichthyidae, excluyendo a las especies de Macquaria (= Percalates). Adicionalmente, los resultados revelan la existencia de variados problemas taxonómicos, tales como la existencia de una sola especie de Percichthys en Argentina, cuya interpretación no es soportada por este estudio; la existencia de especies crípticas de Percilia, Gadopsis, Nannoperca y Macquaria que aumentarán la diversidad específica del género; y la necesidad de una revisión extensiva de especies previamente asignadas a Percalates versus Macquaria. Los desacuerdos encontrados apuntan a la necesidad de investigar más profundamente sobre los así llamados percíctidos y percíliidos.(AU)

Patterning the spine.

The patterning of the spine of a zebrafish is controlled by the notochord, a rod-like structure that supports and instructs the developing embryo.

Otomorphs (= otocephalans or ostarioclupeomorphs) revisited

A morphological revision is presented here on the cohort Otomorpha, a clade currently interpreted as the most primitive among the large supercohort Clupeocephala. Otomorpha is a morphologically heterogeneous group represented by clupei forms , alepocephaliforms, and ostariophysans (gonorynchiforms, cypriniforms, characiforms, siluriforms, and gymnoti forms) that inhabit various marine and freshwater environments worldwide. Otomorphs have a long (ca. 145 Ma) and diverse fossil record. They are the largest fish teleostean clade worldwide, as well as the largest of the Neotropical Region. While molecular studies strongly confirm the monophyly of Otomorpha, most potential morphological synapomorphies of the group become homoplastic largely due to the peculiar morphological character states (either losses or transformations) present in alepocephaliforms. The fusion of haemal arches with their respective vertebral centra anterior to preural centrum 2 stands as an unambiguous synapomorphy of the clade. The ankylosis or fusion of the extrascapular and parietal bones, and silvery areas associated with the gas bladder are also interpreted as synapomorphies, although they are homoplastic characters mainly due to secondary losses or further transformations of the morphological features in the alepocephaliforms.(AU)

Se realizó una revisión morfológica de la cohorte Otomorpha la que se interpreta como el grupo más primitivo dentro de la gran supercohorte Clupeocephala. Otomorpha incluye peces con una gran diversidad corporal la que está representada por clupeiformes, alopocefáliformes y ostariofisos (gonorinchiformes, cipriniformes, caraciformes, siluriformes y gimnotiformes), los que habitan diversos ambientes marinos y de aguas continentales del planeta. Otomorfos son el grupo de peces más grande a nivel mundial y al mismo tiempo, el más grande de la Región Neotropical. Mientras estudios moleculares confirman la monofilia de Otomorfa, la mayoría de las sinapomorfías morfológicas del grupo se interpretan como homoplásticas debido fundamentalmente a la naturaleza peculiar de ciertos caracteres morfológicos (ya sea pérdidas o transformación de estados de caracteres) de alepocefaliformes. La fusión de los arcos hemales con sus respectivos centros vertebrales anterior al centro preural 2 es una sinapomorfía de la cohorte. La anquilosis o fusión de los huesos extrascapular y parietal y la presencia de áreas plateadas asociadas con la vejiga natatoria son interpretados como sinapomorfías, independientemente de que son caracteres homoplásticos debido a pérdidas o transformaciones de tales caracteres en los alepocefáliformes.(AU)

Morphological and taxonomic descriptions of a new genus and species of killifishes (Teleostei: Cyprinodontiformes) from the high Andes of northern Chile.

A new genus and species, Pseudorestias lirimensis, is described from the southern part of the Chilean Altiplano. While sharing several characters that clearly align the new species with Orestias, this new fish is characterized by numerous autapomorphies: the Meckel cartilage is a continuous cartilage that broadly expands posteriorly (in large specimens, it keeps its anterior part and is resorbed posteriorly), the basibranchials are fused into one long element, the second pharyngobranchial is not displaced dorsally over pharyngobranchial tooth plate 3+4, but they are aligned, the anterior and posterior ceratohyals are closely articulated keeping a scarce amount of cartilage between both bones and ventral to them, ossified middle and distal dorsal radials are present in females as well as ossified middle and distal anal radials. Pseudorestias lirimensis presents strong sexual dimorphism associated to size. Females are almost twice as large and long than males, neuromast lines are absent in males, a mesethmoid is present in males, squamation on head is reduced in males, and ossified middle and distal radial of dorsal fin are cartilaginous in males. Pseudorestias and Orestias are suggested as the sole members of the tribe Orestiini. A list of characters diagnosing the tribe is provided. The presence of the new genus is interpreted as a possible result of the ecosystem isolation where the fish is living from surrounding basins-as early as possibly from the Miocene-Pliocene times-and its physical and chemical characteristics. Small populations, living conditions, small habitat, and reduced distribution make this species a strong candidate to be considered critically endangered, a situation already established for all other Chilean species living in the Altiplano. There is high probability it will become extinct due to water demands and climate change in the region.

Phylogenetic classification of bony fishes.

BACKGROUND: Fish classifications, as those of most other taxonomic groups, are being transformed drastically as new molecular phylogenies provide support for natural groups that were unanticipated by previous studies. A brief review of the main criteria used by ichthyologists to define their classifications during the last 50 years, however, reveals slow progress towards using an explicit phylogenetic framework. Instead, the trend has been to rely, in varying degrees, on deep-rooted anatomical concepts and authority, often mixing taxa with explicit phylogenetic support with arbitrary groupings. Two leading sources in ichthyology frequently used for fish classifications (JS Nelson's volumes of Fishes of the World and W. Eschmeyer's Catalog of Fishes) fail to adopt a global phylogenetic framework despite much recent progress made towards the resolution of the fish Tree of Life. The first explicit phylogenetic classification of bony fishes was published in 2013, based on a comprehensive molecular phylogeny ( www.deepfin.org ). We here update the first version of that classification by incorporating the most recent phylogenetic results. RESULTS: The updated classification presented here is based on phylogenies inferred using molecular and genomic data for nearly 2000 fishes. A total of 72 orders (and 79 suborders) are recognized in this version, compared with 66 orders in version 1. The phylogeny resolves placement of 410 families, or ~80% of the total of 514 families of bony fishes currently recognized. The ordinal status of 30 percomorph families included in this study, however, remains uncertain (incertae sedis in the series Carangaria, Ovalentaria, or Eupercaria). Comments to support taxonomic decisions and comparisons with conflicting taxonomic groups proposed by others are presented. We also highlight cases were morphological support exist for the groups being classified. CONCLUSIONS: This version of the phylogenetic classification of bony fishes is substantially improved, providing resolution for more taxa than previous versions, based on more densely sampled phylogenetic trees. The classification presented in this study represents, unlike any other, the most up-to-date hypothesis of the Tree of Life of fishes.

Understanding morphological variability in a taxonomic context in Chilean diplomystids (Teleostei: Siluriformes), including the description of a new species.

Following study of the external morphology and its unmatched variability throughout ontogeny and a re-examination of selected morphological characters based on many specimens of diplomystids from Central and South Chile, we revised and emended previous specific diagnoses and consider Diplomystes chilensis, D. nahuelbutaensis, D. camposensis, and Olivaichthys viedmensis (Baker River) to be valid species. Another group, previously identified as Diplomystes sp., D. spec., D. aff. chilensis, and D. cf. chilensis inhabiting rivers between Rapel and Itata Basins is given a new specific name (Diplomystes incognitus) and is diagnosed. An identification key to the Chilean species, including the new species, is presented. All specific diagnoses are based on external morphological characters, such as aspects of the skin, neuromast lines, and main lateral line, and position of the anus and urogenital pore, as well as certain osteological characters to facilitate the identification of these species that previously was based on many internal characters. Diplomystids below 150 mm standard length (SL) share a similar external morphology and body proportions that make identification difficult; however, specimens over 150 mm SL can be diagnosed by the position of the urogenital pore and anus, and a combination of external and internal morphological characters. According to current knowledge, diplomystid species have an allopatric distribution with each species apparently endemic to particular basins in continental Chile and one species (O. viedmensis) known only from one river in the Chilean Patagonia, but distributed extensively in southern Argentina.

Multi-locus phylogenetic analysis reveals the pattern and tempo of bony fish evolution.

Over half of all vertebrates are "fishes", which exhibit enormous diversity in morphology, physiology, behavior, reproductive biology, and ecology. Investigation of fundamental areas of vertebrate biology depend critically on a robust phylogeny of fishes, yet evolutionary relationships among the major actinopterygian and sarcopterygian lineages have not been conclusively resolved. Although a consensus phylogeny of teleosts has been emerging recently, it has been based on analyses of various subsets of actinopterygian taxa, but not on a full sample of all bony fishes. Here we conducted a comprehensive phylogenetic study on a broad taxonomic sample of 61 actinopterygian and sarcopterygian lineages (with a chondrichthyan outgroup) using a molecular data set of 21 independent loci. These data yielded a resolved phylogenetic hypothesis for extant Osteichthyes, including 1) reciprocally monophyletic Sarcopterygii and Actinopterygii, as currently understood, with polypteriforms as the first diverging lineage within Actinopterygii; 2) a monophyletic group containing gars and bowfin (= Holostei) as sister group to teleosts; and 3) the earliest diverging lineage among teleosts being Elopomorpha, rather than Osteoglossomorpha. Relaxed-clock dating analysis employing a set of 24 newly applied fossil calibrations reveals divergence times that are more consistent with paleontological estimates than previous studies. Establishing a new phylogenetic pattern with accurate divergence dates for bony fishes illustrates several areas where the fossil record is incomplete and provides critical new insights on diversification of this important vertebrate group.

The tree of life and a new classification of bony fishes.

The tree of life of fishes is in a state of flux because we still lack a comprehensive phylogeny that includes all major groups. The situation is most critical for a large clade of spiny-finned fishes, traditionally referred to as percomorphs, whose uncertain relationships have plagued ichthyologists for over a century. Most of what we know about the higher-level relationships among fish lineages has been based on morphology, but rapid influx of molecular studies is changing many established systematic concepts. We report a comprehensive molecular phylogeny for bony fishes that includes representatives of all major lineages. DNA sequence data for 21 molecular markers (one mitochondrial and 20 nuclear genes) were collected for 1410 bony fish taxa, plus four tetrapod species and two chondrichthyan outgroups (total 1416 terminals). Bony fish diversity is represented by 1093 genera, 369 families, and all traditionally recognized orders. The maximum likelihood tree provides unprecedented resolution and high bootstrap support for most backbone nodes, defining for the first time a global phylogeny of fishes. The general structure of the tree is in agreement with expectations from previous morphological and molecular studies, but significant new clades arise. Most interestingly, the high degree of uncertainty among percomorphs is now resolved into nine well-supported supraordinal groups. The order Perciformes, considered by many a polyphyletic taxonomic waste basket, is defined for the first time as a monophyletic group in the global phylogeny. A new classification that reflects our phylogenetic hypothesis is proposed to facilitate communication about the newly found structure of the tree of life of fishes. Finally, the molecular phylogeny is calibrated using 60 fossil constraints to produce a comprehensive time tree. The new time-calibrated phylogeny will provide the basis for and stimulate new comparative studies to better understand the evolution of the amazing diversity of fishes.

Integrating multi-origin expression data improves the resolution of deep phylogeny of ray-finned fish (Actinopterygii).

The actinopterygians comprise nearly one-half of all extant vertebrate species and are very important for human well-being. However, the phylogenetic relationships among certain groups within the actinopterygians are still uncertain, and debates about these relationships have continued for a long time. Along with the progress achieved in sequencing technologies, phylogenetic analyses based on multi-gene sequences, termed phylogenomic approaches, are becoming increasingly common and often result in well-resolved and highly supported phylogenetic hypotheses. Based on the transcriptome sequences generated in this study and the extensive expression data currently available from public databases, we obtained alignments of 274 orthologue groups for 26 scientifically and commercially important actinopterygians, representing 17 out of 44 orders within the class Actinopterygii. Using these alignments and probabilistic methods, we recovered relationships between basal actinopterygians and teleosts, among teleosts within protacanthopterygians and related lineages, and also within acanthomorphs. These relationships were recovered with high confidence.

Connecting evolutionary morphology to genomics using ontologies: a case study from Cypriniformes including zebrafish.

One focus of developmental biology is to understand how genes regulate development, and therefore examining the phenotypic effects of gene mutation is a major emphasis in studies of zebrafish and other model organisms. Genetic change underlies alterations in evolutionary characters, or phenotype, and morphological phylogenies inferred by comparison of these characters. We will utilize both existing and new ontologies to connect the evolutionary anatomy and image database that is being developed in the Cypriniformes Tree of Life project to the Zebrafish Information Network (HYPERLINK "file://localhost/Library/Local%20Settings/Temp/zfin.org" zfin.org) database. Ontologies are controlled vocabularies that formally represent hierarchical relationships among defined biological concepts. If used to recode the free-form text descriptors of anatomical characters, evolutionary character data can become more easily computed, explored, and mined. A shared ontology for homologous modules of the phenotype must be referenced to connect the growing databases in each area in a way that evolutionary questions can be addressed. We present examples that demonstrate the broad utility of this approach.

Breeding tubercles of Phoxinus (Teleostei: Cyprinidae): Morphology, distribution, and phylogenetic implications.

Studies of the skin with scanning electron microscopy (SEM) reveal a diverse morphology in breeding tubercles among species of Phoxinus. Based mainly on the fine structure of the surface of tubercles, nine morphotypes, coded as letters A-I, occur in Phoxinus. Most of the morphotypes are common to all Phoxinus species, but type E is present only on the dorsum of the head of P. phoxinus, type H on the breast scale of female P. phoxinus, and type I on the pectoral fin in P. erythrogaster. Multicellular breeding tubercles bearing unicellular projections, identified as unculiferous tubercles are found in type H and probably types F and G. The distribution of tubercles on head, body, and fins is described and compared among Phoxinus species. Breeding tubercles in Phoxinus and other minnow genera are compared in order to interpret the phylogenetic implication of the tuberculation in Phoxinus. The deeply embedded breast scales and the breeding tubercles on their apical margins, and a series of tubercles on the apical margins of lateral scales of the caudal peduncle in breeding males of Phoxinus species, are the characters supporting the monophyly of the genus. © 1996 Wiley-Liss, Inc.

Olfactory organ of acipenseriformes and comparison with other actinopterygians: Patterns of diversity.

The position and structure of the olfactory organ and its openings vary among actinopterygians. The anterior nasal opening is a simple perforation in the skin in many extant actinopterygians (e.g., acipenseriforms, lepisosteids, and primitive Recent teleosts) and represents the primitive condition. Polypterids and Amia each exhibit a derived condition, in which the anterior nasal opening extends into a tube. The olfactory organ is relatively far away from the anterior end of the elongate rostrum in acipenseriforms, whereas the olfactory organs are closer to the anterior end of the snout in extant actinopterygians (e.g., polypterids, lepisosteids, and amiids). In adults, olfactory organs are cuplike structures in most actinopterygians, but these organs are tubelike in polypterids. Among extant actinopterygians, a nasal diverticulum is present only in polypterids. Teleosts have accessory nasal sacs, but chondrosteans, polypterids, lepisosteids, and amiids lack them. The olfactory rosette is formed by primary folds or lamellae that may be placed anterior, lateral, posterior, and/or medial to the axis of the organ. Large acipenserids have 20-32 lamellae, polyodontids have 13-18 lamellae, lepisosteids have 8-10 lamellae, and Amia may have over 100. In teleosts, the number of lamellae varies from none or a few to over 200. Secondary lamellae are present in acipenseriforms, lepisosteids, and some advanced teleosts; secondary lamellae are interpreted as independently acquired in these lineages. Secondary lamellae are absent in Amia and primitive teleosts such as Elops and Hiodon. Tertiary lamellae are present in Acipenser oxyrhynchus. The arrangement of the primary lamellae in relation to the axis of the organ results in at least 11 patterns of the olfactory rosette in actinopterygians. Lamellae that are enclosed in a tubelike sac and that have an anteromedial diverticulum are specializations of polypterids. Primary lamellae anterior, lateral, and posterior to an elongate axis are characteristic of lepisosteids. The presence of primary lamellae lateral, medial, and posterior to an elongate olfactory axis is a synapomorphy of Halecomorpha (Amia plus teleosts). The absence of secondary lamellae is a synapomorphy of Halecomorpha. © 1994 Wiley-Liss, Inc.

Jaws and teeth of american cichlids (Pisces: Labroidei).

The morphology of the upper, lower, and pharyngeal jaws is very similar among American cichlids. Common conditions are: (1) the presence of a premaxillary dentigerous arm shorter than the ascending arm (exceptions are Astronotus, Cichla, and Crenicichla semifasciata), (2) a narrow coulter area; in contrast, a broad coulter area is found in the Crenicichline Group, in certain chaetobranchines, and in Apistogramma, (3) the mandibular sensory canal exists to the skin through five or six simple pores; in contrast, it exits through numerous small pores that increase in number during ontogeny in the Chaetobranchine Group, certain crenicichlines, such as Cichla, Crenicichla lepidota, Crenicichla proteus, and Crenicichla vittata, and certain genera of the Cichlasomine Group A, such as Caquetaia, Petenia, Neetroplus, and "Cichlasoma," and (4) the premaxilla and dentary of American cichlids commonly bear unicuspid, conical teeth with a few exceptions such as Neetroplus (with scraping blade teeth) and "Cichlasoma" facetum, "C." cyanoguttatum, "C." guttulatum, and "C." spilurum (with bicuspid [hooked] teeth). In contrast to the near uniformity of the upper and lower jaws, the upper and lower pharyngeal jaws present a great diversity of tooth shapes. At least seven types are found in American cichlids; usually, several types exist on a single tooth plate, but the combination of tooth types differs among some genera. The pharyngobranchial 4 tooth plate has significant evolutionary transformations in labroids. The caudal margin of the pharyngobranchial 4 tooth plate bears the frayed zone in cichlids and embiotocids. The presence of a broad frayed zone bearing one to seven concavities represents a synapomorphy for the family Cichlidae, whereas a deep, narrow frayed zone is a synapomorphy of Embiotocidae. The absence of the frayed zone is a synapomorphy of Pomacentridae, whereas the loss of the pharyngobranchial 4 is a synapomorphy of Labridae. © 1993 Wiley-Liss, Inc.

Reevaluation of the caudal skeleton of certain actinopterygian fishes: III. Salmonidae. Homologization of caudal skeletal structures.

The ontogenetic development of caudal vertebrae and associated skeletal elements of salmonids provides information about sequence of ossification and origin of bones that can be considered as a model for other teleosts. The ossification of elements forming the caudal skeleton follows the same sequence, independent of size and age at first appearance. Dermal bones like principal caudal rays ossify earlier than chondral bones; among dermal bones, the middle principal caudal rays ossify before the ventral and dorsal ones. Among chondral bones, the ventral hypural 1 and parhypural ossify first, followed by hypural 2 and by the ventral spine of preural centrum 2. The ossification of the dorsal chondral elements starts later than that of ventral ones. Three elements participate in the formation of a caudal vertebra: paired basidorsal and basiventral arcocentra, chordacentrum, and autocentrum; appearance of cartilaginous arcocentra precedes that of the mineralized basiventral chordacentrum, and that of the perichordal ossification of the autocentrum. Each ural centrum is mainly formed by arcocentral and chordacentrum. The autocentrum is irregularly present or absent. Some ural centra are formed only by a chordacentrum. This pattern of vertebral formation characterizes basal teleosts and primitive extant teleosts such as elopomorphs, osteoglossomorphs, and salmonids. The diural caudal skeleton is redefined as having two independent ural chordacentra plus their arcocentra, or two ural chordacentra plus their autocentra and arococentra, or only two ural chordacentra. A polyural caudal skeleton is identified by more than two ural centra, variably formed as given for the diural condition. The two ural centra of primitive teleosts may result from early fusion of ural centra 1 and 2 and of ural centra 3 and 4, or 3, 4, and 5 (e.g., elopomorphs), respectively. The two centra may corespond to ural centrum 2 and 4 only (e.g., salmonids). Additionally, ural centra 1 and 3 may be lost during the evolution of teleosts. Additional ural centra form late in ontogeny in advanced salmonids, resulting in a secondary polyural caudal skeleton. The hypural, which is a haemal spine of a ural centrum, results by growth and ossification of a single basiventral ural arococentrum and its haemal spine. The proximal part of the hypural always includes part of the ventral ural arcocentrum. The uroneural is a modification of a ural neural arch, which is demonstrated by a cartilaginous precursor. The stegural of salmonids and esocids originates from only one paired cartilaginous dorsal arcocentrum that grows anteriorly by a perichondral basal ossification and an anterodorsal membranous ossification. The true epurals of teleosts are detached neural spines of preural and ural neural arches as shown by developmental series; they are homologous to the neural spines of anterior vertebrae. Free epurals without any indication of connection with the dorsal arococentra are considered herein as an advanced state of the epural. Caudal distal radials originate from the cartilaginous distal portion of neural and haemal spines of preural and ural (epurals and hypurals) vertebrae. Therefore, they result from distal growth of the cartilaginous spines and hypurals. Cartilaginous plates that support rays are the result of modifications of the plates of connective tissue at the posterior end of hypurals (e.g., between hypurals 2 and 3 in salmonids) and first preural haemal spines, or from the distal growth of cartilaginous spines (e.g., epural plates in Thymallus). Among salmonids, conditions of the caudal skeleton such as the progressive loss of cartilaginous portions of the arcocentra, the progressive fusion between the perichondral ossification of arcocentra and autocentra, the broadening of the neural spines, the enlargement and interdigitation of the stegural, and other features provide evidence that Prosopium and Thymallus are the most primitive, and that Oncorhynchus and Salmo are the most advanced salmonids respectively. This interpretation supports the current hypothesis of phylogenetic relationships of salmonids. © 1992 Wiley-Liss, Inc.

Palatoquadrate and its ossifications: Development and homology within osteichthyans.

The palatoquadrate and associated dermal bones have significant evolutionary transformations among teleostomes and provide numerous features that characterize teleostomian subgroups. The palatoquadrate forms the upper part of the mandibular arch and is present as a single cartilaginous element in the early ontogeny of teleostomes, except for some advanced teleosts such as siluroids where it is divided into pars autopalatina and pars pterygoquadrata. During ontogeny, the palatoquadrate may ossify as a unit, with a pars autopalatina (absent in Acanthodii), pars quadrata, and pars metapterygoidea in teleostomes (e.g., primitive acanthodians and actinopterygians, onychodonts, and rhipidistians). However, the palatoquadrate may remain cartilaginous (e.g., chondrosteans) or it may ossify as separate elements (e.g., autopalatine, metapterygoid, and quadrate) as occurs in advanced acanthodians, Polypterus and advanced actinopterygians, and advanced actinistians. From the single-unit pattern, separate autopalatine, metapterygoid, and quadrate evolve in parallel in the three teleostomian subgroups. Therefore, it is necessary to distinguish between actinopterygian and actinistian autopalatines and among acanthodian, actinopterygian, and actinistian metapterygoids and quadrates. A palatoquadrate fused with the neurocranium occurs in parallel in dipnoans. There are differences in the timing of ossification of the autopalatine, metapterygoid, and quadrate. The autopalatine ossifies late in ontogeny in Polypterus, Amia, and primitive teleosts (absent in lepisosteids and osteoglossmorphs), whereas both metapterygoid and quadrate ossify early in ontogeny. The early ossification of the autopalatine is characteristic of clupeocephalan teleosts. During ontogeny, tooth plates (not forming a separate dermometapterygoid) fuse with the metapterygoid in actinopterygians. Pars autopalatina, pars metapterygoidea, and pars quadrata are regions at the three corners of the single-unit palatoquadrate present in primitive teleostomes; there are no clear limits among these regions, but they may be identified by their processes, articular facets, and topographical relationships with surrounding bones and the orbit. Autopalatine, metapterygoid, and quadrate are chondral bones, perichondrally ossified. Dermal elements such as dermopalatine(s), entopterygoid, ectopterygoid, and tooth plates may cover the palatoquadrate medially. The predermopalatine that originates in front of pars autopalatina in Cladistia and the "dermopalatine" that lies medial to the ectopterygoid in Ginglymodi are specializations of these groups. A dermopalatine fused with the autopalatine is characteristic of clupeocephalan teleosts. Highly specialized tendon bone pterygoids are found in some teleosts (e.g., siluroids). The presence of both maxilla and lacrimal lateral to the pars autopalatina is synapomorphous of osteichthyans. The eye supported by the bony palatoquadrate is a teleostomian synapomorphy. Dermal elements support the eye in actinopterygians, the entopterygoid in advanced actinopterygians, but the ectopterygoid in lepisosteids. A quadratojugal is a synapomorphy of osteichthyans but exhibits a number of transformations in connection with the vertical pit-line and the preopercular canal; a quadratojugal bearing the vertical pit-line is the primitive condition for osteichthyans. Ontogenetic evidence does not support the homology of the membranous posterior process of the teleostean quadrate with the quadratojugal. The lack of a quadratojugal and the presence of the elongate posterior or posteroventral process of the quadrate is a synapomorphy of teleosts.

The urohyal: Development and homology within osteichthyans.

The formation of the unpaired structure ventral to the basibranchial region, the so-called urohyal, differs within osteichthyans. A cartilaginous preformed, unpaired "urohyal" is present in sarcopterygians. A three-tendon ossification is present in Polypterus. An "urohyal" or urohyal is absent in both Amia and Lepisosteus. The urohyal formed as an unpaired ossification of the tendon of the sternohyoideus muscle is a feature of teleosts. A new structure, the parurohyal, arises as a double ossification of the tendon of the sternohyoideus muscle in siluroids; during ontogeny an anterodorsal crest or cup-like structure derives from the anterior basibranchial region and the tendon bone; therefore, the parurohyal is compound in origin. Judging from their formation and their distribution within osteichthyans the cartilaginous preformed "urohyal" and the teleostean urohyal are nonhomologous, whereas the urohyal and parurohyal are homologous. The urohyal is connected by ligaments with the ventral hypohyals in most teleosts, whereas it articulates with the ventral hypohyals in teleosts such as Anguilla and Chanos. The parurohyal is a synapomorphy of siluroids. The parurohyal in siluroids is articulated with both ventral and dorsal hypohyals, and with the basibranchial region in catfishes such as diplomystids and ictalurids, whereas it articulates only with the ventral hypohyals in other catfishes such as trichomycterines. The passage of the hypobranchial artery through the hypobranchial foramen of the parurohyal is a unique feature of siluroids, like the absence of the basihyal bone. An ossified dorsal hypohyal appears late in ontogeny in Amia, as do tooth plates related to the medial side of the hyoid arch and branchiostegal rays in Amia, and tooth plates on the hyoid arch and branchiostegal rays in Elops (unique features within extant teleosts). Two ossified hypohyals, a synapomorphy of teleosts, are present early in ontogeny. There is intraspecific variation in the onset of ossification of the bones of the head, but the sequence of ossification between bones in a defined structural system is conserved (e.g., branchiostegal rays ossify first, then bones of the hyoid arch).