Edgeworth's legacy of cranial muscle development with an analysis of muscles in the ventral gill arch region of batoid fishes (Chondrichthyes: Batoidea).

A series of studies by Edgeworth demonstrated that cranial muscles of gnathostome fishes are embryologically of somitic origin, originating from the mandibular, hyoid, branchial, epibranchial, and hypobranchial muscle plates. Recent experimental studies using quail-chick chimeras support Edgeworth's view on the developmental origin of cranial muscles. One of his findings, the existence of the premyogenic condensation constrictor dorsalis in teleost fishes, has also been confirmed by molecular developmental studies. Therefore, developmental mechanisms for patterning of cranial muscles, as described and implicated by Edgeworth, may serve as structural entities or regulatory phenomena responsible for developmental and evolutionary changes. With Edgeworth's and other studies as background, muscles in the ventral gill arch region of batoid fishes are analyzed and compared with those of other gnathostome fishes. The spiracularis is regarded as homologous at least within batoid fishes, but its status within elasmobranchs remains unclear; developmental modifications of the spiracularis proper are evident in some batoid fishes and in several shark groups. The peculiar ventral extension of the spiracularis in electric rays and some stingrays may represent convergence, probably facilitating ventilation and/or feeding in both groups. The evolutionary origin of the "internus" and "externus" remains uncertain, despite the fact that a variety of forms of the constrictor superficiales ventrales in batoid fishes indicates an actual medio-ventral extension of the "externus." The intermandibularis is probably present only in electric rays. The "X" muscle occurs only in electric rays and is considered to be Edgeworth's intermandibularis profundus. Its association with the adductor mandibular complex in narkinidid and narcinidid electric rays may relate to its functional role in lower jaw movement. Contrary to common belief, in most batoid fishes as well as some sharks, muscles that originate from the branchial muscle plate and extend medially in the ventral gill arches do exist: the medial extension of the interbranchiales in most batoid fishes and some sharks and the "Y" muscle in the pelagic stingrays Myliobatos and Rhinoptera. The latter is another example of the medial extension of the "internus." Whether the interbranchiales and "Y" muscle are homologous within elasmobranchs and whether homologous with the obliques ventrales and/or transversi ventrales of osteichthyan fishes await further research. Four hypobranchial muscles are recognized in batoid fishes: the coracomandibularis, coracohyoideus, coracoarcualis, and coracohyomandibularis. The coracohyoideus is discrete from the coracoarcualis; its complete structural separation from the latter occurs in several groups of batoid fishes.(ABSTRACT TRUNCATED AT 400 WORDS)     

Vertebral column and associated elements in dipnoans and comparison with other fishes: development and homology.

A vertebral column consisting of a persistent notochord and ossified arcocentra is the primitive condition for Gnathostomata; it still persists in primitive actinopterygians and sarcopterygians. Advanced actinopterygians and sarcopterygians develop numerous types of centra that include, among others, the presence of holocentrum, chordacentrum, and autocentrum. The chordacentrum, a mineralization or calcification of the fibrous sheath of the notochord, is only found in actinopterygians, whereas an autocentrum is a synapomorphy of teleosts above Leptolepis coryphaenoides. The chordacentrum, formed by migration of cartilaginous cells from the arches into the fibrous sheath of the notochord and usually covered by a thin calcification, is a unique feature of chondrichthyans. The actinopterygian chordacentrum and the chondrichthyan chordacentrum are not homologous. The postcaudal cartilaginous centrum is only known in postcaudal vertebrae of living dipnoans. The holocentrum is present in certain fossil dipnoans and actinopterygians, where it has been independently acquired. It is formed by proliferation of cartilage cells around the elastica externa of the notochord. These cells later ossify, forming a compact centrum. A vertebral column formed by a persistent notochord without vertebral centra is the primitive pattern for all vertebrates. The formation of centra, which is not homologous among vertebrate groups, is acquired independently in some lineages of placoderms, most advanced actinopterygians, and some dipnoans and rhipidistians. Several series of structures are associated with the vertebral column such as the supraneurals, interhaemals, radials, and ribs. In living dipnoans median neural spine, "supraneural," and dorsal radial result from growth and distal differentiation of one median cartilage into two or three median bones during ontogeny. The median neural spine articulates with the neural arch and fuses with it in the caudal vertebrae early in ontogeny. Two bones differentiate in the anterior abdominal vertebrae, i.e., the proximal neural spine and the distal "supraneural." Three bones differentiate in front of the dorsal fin, i.e., the proximal neural spine, the middle "supraneural", and the distal radial; the same pattern is observed in front of the anal fin (the proximal haemal spine, the middle interhaemal, and the distal radial). Considering that the three dorsal (and also the three ventral) bones originate from growth of only one cartilage, they cannot be serial homologs of the neural spines, or "supraneural." They are linear homologs of the median neural cartilage in living dipnoans. The development of these elements differs within osteichthyans from sarcopterygians to actinopterygians, in which the neural spine originates as a continuation of the basidorsal arcualia and in which the supraneural and radial originate from independent cartilages that appear at different times during early ontogeny. The ribs of living dipnoans are unique in that they are not articulated with parapophyses, like in primitive fossil dipnoans, but a remnant of the ventral arcuale surrounded by a small arcocentrum remains at its base. A true caudal fin is absent in living dipnoans. The postcaudal cartilages extend to the caudal tip of the body separating dorsal and ventral rays (or the camptotrichia). Actinotrichia are present in young dipnoans. They are also known in extant actinistians and actinopterygians. They probably represent the primitive state for teleostomes. In contrast, the camptotrichia are unique for extant dipnoans (and probably Carboniferous and younger dipnoans). Lepidotrichia apparently developed many times among osteichthyans.     

Cephalic muscle development in the Australian lungfish,Neoceratodus forsteri

Lungfishes are the extant sister group of tetrapods. As such, they are important for the study of evolutionary processes involved in the water to land transition of vertebrates. The evolution of a true neck, that is, the complete separation of the pectoral girdle from the cranium, is one of the most intriguing morphological transitions known among vertebrates. Other salient changes involve new adaptations for terrestrial feeding, which involves both the cranium and its associated musculature. Historically, the cranium has been extensively investigated, but the development of the cranial muscles much less so. Here, we present a detailed study of cephalic muscle development in the Australian lungfish, Neoceratodus forsteri, which is considered to be the sister taxon to all other extant lungfishes. Neoceratodus shows several developmental patterns previously described in other taxa; the tendency of muscles to develop from anterior to posterior, from their region of origin toward insertion, and from lateral to ventral/medial (outside‐in), at least in the branchial arches. The m.protractor pectoralis appears to develop as an extension of the most posterior m.levatores arcuum branchialium, supporting the hypothesis that the m.cucullaris and its derivatives (protractor pectoralis, levatores arcuum branchialium) are branchial muscles. We present a new hypothesis regarding the homology of the ventral branchial arch muscles (subarcualis recti and obliqui, transversi ventrales) in lungfishes and amphibians. Moreover, the morphology and development of the cephalic muscles confirms that extant lungfishes are neotenic and have been strongly influenced via paedomorphosis during their evolutionary history.     

CT scan reconstruction of the palate region of Latimeria chalumnae

Synopsis The palate of Latimeria chalumnae is described based mainly on three-dimensional CT scan reconstruction. It is compared with that of other osteichthyans. The palate of L. chalumnae compares best with that of rhipidistians; it is more advanced than that of actinopterygians in having fewer bones. This tendency toward bone reduction in the palate is even more pronounced in dipnoans. The interpretation of features of the Early Devonian genus Diabolepis determines if authors consider dipnoans or actinistians more closely related to tetrapods. Both groups are only distant relatives of tetrapods.     

Development of cranial muscles in the actinopterygian fish Senegal bichir,Polypterus senegalus Cuvier, 1829

Polypterus senegalus Cuvier, 1829 is one of the most basal living actinopterygian fish and a member of the Actinopterygii. We analyzed the spatial and temporal pattern of cranial muscle development of P. senegalus using whole‐mount immunostaining and serial sectioning. We described the detailed structure of the external gill muscles which divided into dorsal and ventral parts after yolk exhaustion. The pattern of the division is similar to that of urodeles. We suggest that, the external gill muscles of P. senegalus are involved in spreading and folding of the external gill stem and the branches. The fibers of the external gill muscles appear postero‐lateral to the auditory capsule. In addition, the facial nerve passes through the external gills. Therefore, the external gill muscles are probably derived from the m. constrictor hyoideus dorsalis. In contrast to previous studies, we described the mm. interhyoideus and hyohyoideus fibers as independent components in the yolk‐sac larvae. The m. hyohyoideus fibers appear lateral to the edge of the ventral portion of the external gill muscles, which are probably derived from the m. constrictor hyoideus dorsalis. These findings suggest that the m. hyohyoidues is derived from the m. constrictor hyoideus dorsalis in P. senegalus . In other actinopterygians, the m. hyohyoideus is derived from the m. constrictor hyoideus ventralis; therefore, the homology of the m. hyohyoidues of P. senegalus and other actinopterygians remains unclear. J. Morphol. 278:450–463, 2017. © 2017 Wiley Periodicals, Inc.     

Homology of the Fifth Epibranchial and Accessory Elements of the Ceratobranchials among Gnathostomes: Insights from the Development of Ostariophysans

Epibranchials are among the main dorsal elements of the gill basket in jawed vertebrates (Gnathostomata). Among extant fishes, chondrichthyans most resemble the putative ancestral condition as all branchial arches possess every serially homologous piece. In osteichthyans, a primitive rod-like epibranchial 5, articulated to ceratobranchial 5, is absent. Instead, epibranchial 5 of many actinopterygians is here identified as an accessory element attached to ceratobranchial 4. Differences in shape and attachment of epibranchial 5 in chondrichthyans and actinopterygians raised suspicions about their homology, prompting us to conduct a detailed study of the morphology and development of the branchial basket of three ostariophysans (Prochilodus argenteus, Characiformes; Lophiosilurus alexandri and Pseudoplatystoma corruscans, Siluriformes). Results were interpreted within a phylogenetic context of major gnathostome lineages. Developmental series strongly suggest that the so-called epibranchial 5 of actinopterygians does not belong to the epal series because it shares the same chondroblastic layer with ceratobranchial 4 and its ontogenetic emergence is considerably late. This neomorphic structure is called accessory element of ceratobranchial 4. Its distribution among gnathostomes indicates it is a teleost synapomorphy, occurring homoplastically in Polypteriformes, whereas the loss of the true epibranchial 5 is an osteichthyan synapomorphy. The origin of the accessory element of ceratobranchial 4 appears to have occurred twice in osteichthyans, but it may have a single origin; in this case, the accessory element of ceratobranchial 4 would represent a remnant of a series of elements distally attached to ceratobranchials 1–4, a condition totally or partially retained in basal actinopterygians. Situations wherein a structure is lost while a similar neomorphic element is present may lead to erroneous homology assessments; these can be avoided by detailed morphological and ontogenetic investigations interpreted in the light of well-supported phylogenetic hypotheses.     

Evolution of the facial musculature in basal ray-finned fishes

Background

The facial musculature is a remarkable anatomical complex involved in vital activities of fishes, such as food capture and gill ventilation. The evolution of the facial muscles is largely unknown in most major fish lineages, such as the Actinopterygii. This megadiverse group includes all ray-finned fishes and comprises approximately half of the living vertebrate species. The Polypteriformes, Acipenseriformes, Lepisosteiformes, Amiiformes, Elopiformes, and Hiodontiformes occupy basal positions in the actinopterygian phylogeny and a comparative study of their facial musculature is crucial for understanding the cranial evolution of bony fishes (Osteichthyes) as a whole.

Results

The facial musculature of basal actinopterygians is revised, redescribed, and analyzed under an evolutionary perspective. We identified twenty main muscle components ontogenetically and evolutionarily derived from three primordial muscles. Homologies of these components are clarified and serve as basis for the proposition of a standardized and unifying myological terminology for all ray-finned fishes. The evolutionary changes in the facial musculature are optimized on the osteichthyan tree and several new synapomorphies are identified for its largest clades, including the Actinopterygii, Neopterygii, and Teleostei. Myological data alone ambiguously support the monophyly of the Holostei. A newly identified specialization constitutes the first unequivocal morphological synapomorphy for the Elopiformes. The myological survey additionally allowed a reinterpretation of the homologies of ossifications in the upper jaw of acipenseriforms.

Conclusions

The facial musculature proved to be extremely informative for the higher-level phylogeny of bony fishes. These muscles have undergone remarkable changes during the early radiation of ray-finned fishes, with significant implications for the knowledge of the musculoskeletal evolution of both derived actinopterygians and lobe-finned fishes (Sarcopterygii).

     

Tetrapod teeth from an Upper Ladinian bone bed, Schöningen (Lower Saxony, Germany)

A former clay quarry near Schöningen in Lower Saxony exposes deposits dating from the upper half of the Ladinian. There are several bone beds in this quarry, which differ partly in their faunal composition and in the preservation of the fossils. One of these bone beds contains many morphologically different teeth of mainly terrestrial tetrapods and a variety of remains of actinopterygians, chondrichthyans and dipnoans. The tetrapod teeth are described in this paper. A precise taxonomic determination is not possible, but the material appears to contain two temnospondyls, a synapsid and several species of archosaurs. Some tooth morphotypes can not be assigned, even at a high taxonomic level. The tooth assemblage, which is described in this paper, is briefly compared with published and unpublished data on teeth from the Middle and Upper Keuper of Central Europe.     

The anatomical components of the cardiac outflow tract of chondrichthyans and actinopterygians

The outflow tract of the fish heart is the segment interposed between the ventricle and the ventral aorta. It holds the valves that prevent blood backflow from the gill vasculature to the ventricle. The anatomical composition, histological structure and evolutionary changes in the fish cardiac outflow tract have been under discussion for nearly two centuries and are still subject to debate. This paper offers a brief historical review of the main conceptions about the cardiac outflow tract components of chondrichthyans (cartilaginous fish) and actinopterygians (ray‐finned fish) which have been put forward since the beginning of the nineteenth century up to the current day. We focus on the evolutionary origin of the outflow tract components and the changes to which they have been subject in the major extant groups of chondrichthyans and actinopterygians. In addition, an attempt is made to infer the primitive anatomical design of the heart of the gnathostomes (jawed vertebrates). Finally, several areas of further investigation are suggested. Recent work on fish heart morphology has shown that the cardiac outflow tract of chondrichthyans does not consist exclusively of the myocardial conus arteriosus as classically thought. A conus arteriosus and a bulbus arteriosus, devoid of myocardium and mainly composed of elastin and smooth muscle, are usually present in cartilaginous and ray‐finned fish. This is consistent with the suggestion that both components coexisted from the onset of the gnathostome radiation. There is evidence that the conus arteriosus appeared in the agnathans. By contrast, the evolutionary origin of the bulbus is still unclear. It is almost certain that in all fish, both the conus and bulbus develop from the embryonic second heart field. We suggest herein that the primitive anatomical heart of the jawed vertebrates consisted of a sinus venosus containing the pacemaker tissue, an atrium possessing trabeculated myocardium, an atrioventricular region with compact myocardium which supported the atrioventricular valves, a ventricle composed of mixed myocardium, and an outflow tract consisting of a conus arteriosus, with compact myocardium in its wall and valves at its luminal side, and a non‐myocardial bulbus arteriosus that connected the conus with the ventral aorta. Chondrichthyans have retained this basic anatomical design of the heart. In actinopterygians, the heart has been subject to notable changes during evolution. Among them, the following two should be highlighted: (i) a decrease in size of the conus in combination with a remarkable development of the bulbus, especially in teleosts; and (ii) loss of the myocardial compact layer of the ventricle in many teleost species.     

Gill microcirculation of the air-breathing climbing perch, Anabas testudineus (Bloch):relationships with the accessory respiratory organs and systemic circulation

The general macrocirculation and branchial microcirculation of the air-breathing climbing perch, Anabas testudineus, was examined by light and scanning electron microscopy of vascular corrosion replicas. The ventral aorta arises from the heart as a short vessel that immediately bifurcates into a dorsal and a ventral branch. The ventral branch distributes blood to gill arches 1 and 2, the dorsal branch to arches 3 and 4. The vascular organization of arches 1 and 2 is similar to that described for aquatic breathing teleosts. The respiratory lamellae are well developed but lack a continuous inner marginal channel. The filaments contain an extensive nutritive and interlamellar network; the latter traverses the filament between, but in register with, the inner lamellar margins. Numerous small, tortuous vessels arise from the efferent filamental and branchial arteries and anastomose with each other to form the nutrient supply for the filament, adductor muscles, and arch supportive tissues. The efferent branchial arteries of arches 1 and 2 supply the accessory air-breathing organs. Arches 3 and 4 are modified to serve primarily as large-bore shunts between the dorsal branch of the ventral aorta and the dorsal aorta. In many filaments from arches 3 and 4, the respiratory lamellae are condensed and have only 1-3 large channels. In some instances in arch 4, shunt vessels arise from the afferent branchial artery and connect directly with the efferent filamental artery. The filamental nutrient and interlamellar systems are poorly developed or absent. The respiratory and systemic pathways in Anabas are arranged in parallel. Blood flows from the ventral branch of the ventral aorta, through gill arches 1 and 2, into the accessory respiratory organs, and then returns to the heart. Blood, after entering the dorsal branch of the ventral aorta, passes through gill arches 3 and 4 and proceeds to the systemic circulation. This arrangement optimizes oxygen delivery to the tissues and minimizes intravascular pressure in the branchial and air-breathing organs. The efficiency of this system is limited by the mixing of respiratory and systemic venous blood at the heart.     

Homologies of the branchial arch muscles in Zacco platypus (Teleostei: Cypriniformes: Cyprinidae): Evidence from innervation pattern

Homologies of the branchial arch muscles in the cyprinid Zacco platypus are assessed based on their innervation. Muscles serving the first gill arch are innervated by branches of the glossopharyngeal (IX) nerve and those serving other arches by the vagal (X) nerve. Absence of the levator posterior is confirmed. Five pairs of muscles originating from the cranium and inserted onto the specialized 5th ceratobranchial, all unique to cyprinids, are innervated by the 4th branchial trunks of X, indicating that all pairs are derivatives of the sphincter oesophagi, involving reorganization from intrinsic to extrinsic elements. Homologies of some ventral branchial muscles are also discussed and the criteria for homology improved by clarifying the innervation pattern. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

A three‐dimensional placoderm (stem‐group gnathostome) pharyngeal skeleton and its implications for primitive gnathostome pharyngeal architecture

The pharyngeal skeleton is a key vertebrate anatomical system in debates on the origin of jaws and gnathostome (jawed vertebrate) feeding. Furthermore, it offers considerable potential as a source of phylogenetic data. Well‐preserved examples of pharyngeal skeletons from stem‐group gnathostomes remain poorly known. Here, we describe an articulated, nearly complete pharyngeal skeleton in an Early Devonian placoderm fish, Paraplesiobatis heinrichsi Broili, from Hunsrück Slate of Germany. Using synchrotron light tomography, we resolve and reconstruct the three‐dimensional gill arch architecture of Paraplesiobatis and compare it with other gnathostomes. The preserved pharyngeal skeleton comprises elements of the hyoid arch (probable ceratohyal) and a series of branchial arches. Limited resolution in the tomography scan causes some uncertainty in interpreting the exact number of arches preserved. However, at least four branchial arches are present. The final and penultimate arches are connected as in osteichthyans. A single median basihyal is present as in chondrichthyans. No dorsal (epibranchial or pharyngobranchial) elements are observed. The structure of the pharyngeal skeleton of Paraplesiobatis agrees well with Pseudopetalichthys from the same deposit, allowing an alternative interpretation of the latter taxon. The phylogenetic significance of Paraplesiobatis is considered. A median basihyal is likely an ancestral gnathostome character, probably with some connection to both the hyoid and the first branchial arch pair. Unpaired basibranchial bones may be independently derived in chondrichthyans and osteichthyans.     

The organization of the octavolateralis area in actinopterygian fishes: A new interpretation

The octavolateralis area of actinopterygian fishes can be subdivided into a dorsal lateralis area composed of first-order lateral line nuclei, and a ventral octavus area composed of nuclei receiving first-order input from the eighth nerve. Three patterns of organization of the lateralis area are recognized in the present study. The organization of this area in polypteriforms and chondrosteans is similar to that in chondrichthyans. On the basis of recent studies in chondrichthyans (McCready and Boord, '76; Boord and Campbell, '77; Bodznick and Northcutt, '80), it is hypothesized that this pattern reflects the subdivision of the lateral line system into mechanoreceptive and electroreceptive portions. As petromyzontid agnathans also share this pattern of organization, it is hypothesized that they are elecroreceptive. The lateralis area of holosteans and nonelectroreceptive teleosts exhibits a second organizational pattern that is hypothesized to reflect the loss of the electroreceptive portion of the lateral line system; it is suggested that electroreception was lost sometime between the chondrostean and teleostean radiations. Each group of electroreceptive teleosts is believed to have evolved electroreception independently (Bullock, '74), a situation that is reflected centrally by a third organizational pattern within the lateralis area, which is distinctly different from that of early radiations of electroreceptive fishes. The octavus area of actinopterygians exhibits two patterns of organization–that of polypteriforms, chondrosteans, and holosteans, and that of teleosts. The functional significance of these patterns has yet to be elucidated.     

Structural Organization of the Blood-sinus Systems in Lampreys and Hagfish: Functional and Evolutionary Interpretations

Abstract The head and branchial regions of larval and adult lampreys and hagfish were studied histologically in serial sections. The most remarkable feature in these extant agnathans was the occurrence of large blood-sinuses. In larval lampreys, blood-sinuses are well developed in the velum, an organ that functions to introduce water and accompanying food particles from the mouth into the gill and alimentary regions. The sinuses in the velum may act to transduce the force of contraction of velar muscles to the stroke-like movement of the velum; without these sinuses muscular contractions might simply cause the velum to collapse. In adult lampreys, blood-sinuses are well developed in the peribranchial space that surrounds the branchial (gill) sac and is surrounded by the branchial pouch. It is possible that the force of contractions of the branchial-pouch muscles is transduced effectively to the branchial sac via the peribranchial blood-sinus and facilitates the expiration of water through the external gill pores. If the peribranchial sinus were absent, the muscular contraction might deform the branchial sac in an inappropriate manner. In the hagfish, the blood-sinus system is also well developed in the velum and peribranchial space, although the peribranchial sinus lies outside the muscular branchial pouch. In agnathans, the blood-sinus system may function, at least in part, as a kind of hydrostatic skeleton that transduces the force generated by muscular contraction.     

An exceptionally preserved Late Devonian actinopterygian provides a new model for primitive cranial anatomy in ray-finned fishes

Actinopterygians (ray-finned fishes) are the most diverse living osteichthyan (bony vertebrate) group, with a rich fossil record. However, details of their earliest history during the middle Palaeozoic (Devonian) ‘Age of Fishes'' remains sketchy. This stems from an uneven understanding of anatomy in early actinopterygians, with a few well-known species dominating perceptions of primitive conditions. Here we present an exceptionally preserved ray-finned fish from the Late Devonian (Middle Frasnian, ca 373 Ma) of Pas-de-Calais, northern France. This new genus is represented by a single, three-dimensionally preserved skull. CT scanning reveals the presence of an almost complete braincase along with near-fully articulated mandibular, hyoid and gill arches. The neurocranium differs from the coeval Mimipiscis in displaying a short aortic canal with a distinct posterior notch, long grooves for the lateral dorsal aortae, large vestibular fontanelles and a broad postorbital process. Identification of similar but previously unrecognized features in other Devonian actinopterygians suggests that aspects of braincase anatomy in Mimipiscis are apomorphic, questioning its ubiquity as stand-in for generalized actinopterygian conditions. However, the gill skeleton of the new form broadly corresponds to that of Mimipiscis, and adds to an emerging picture of primitive branchial architecture in crown gnathostomes. The new genus is recovered in a polytomy with Mimiidae and a subset of Devonian and stratigraphically younger actinopterygians, with no support found for a monophyletic grouping of Moythomasia with Mimiidae.     

The development of gill arches and gill blood vessels of the rainbow trout,Salmo gairdneri

Gill development begins on the sixth day of incubation at 10^°C and is complete by 31 days (hatching). Gill arches are formed by fusion and perforation of ectoderm and endoderm across the pharyngeal wall. A primary branchial artery forms within each arch and a second branchial artery forms as a branch from its ventral end. A series of filament loop vessels forms connecting the two arteries and when several are patent a unidirectional blood flow is established via afferent (second) branchial artery, filament loop vessels to efferent (primary) branchial artery. Part of the efferent branchial artery just above its junction with the afferent branchial artery constricts and occludes. It is suggested that this change in the pattern of blood flow is dependent on differences in resistance of the two branchial arteries. A later extension of the gill ventrally is thought not to be homologous with similar regions in elasmobranchs and Acipenser.     

Morphology of central compartment and vasculature of the gills of Lepidosiren paradoxa (Fitzinger)

The four paired gill arches of the South American lungfish Lepidosiren paradoxa contain single branchial arteries directly connecting dorsal and ventral arteries. In gill arches 3 and 4 the branchial arteries also supply looped arlerioles and capillaries to much-reduced gill filaments. Regulation of blood between these routes is thought to be by alteration of vascular resistance. Within the filaments, extensive subepithelial capillary networks and numerous small pumps connect lymphatic vessels in the central connective tissue compartment with venules which, in turn, drain to paired branchial veins.
The features of the endothelium of many of the filament blood vessels suggest extensive transporting, haematolytic and granulopoeitic functions. Large numbers of macrophages pack the connective tissue. Many contain extensive quantities of haemosiderin.     

The diencephalon of the primitive bony fish Polypterus in the light of the problem of homology

The diencephalon of Polypterus can be divided into an epithalamus, thalamus, and hypothalamus. The habenulae, the nervous parts of the epithalamus, are comparable to their homologues in other lower vertebrates with respect to sulcal boundaries, cellular structure, and fiber connections. The thalamus of Polypterus is not divisible into a pars dorsalis and pars ventralis by the sulcus medius; rather this sulcus is in the middle of a uniform, laminated cytoarchitectonic field. In this respect Polypterus differs from other species in whom the sulcus medius divides the thalamus into dorsal and ventral parts. There are six migrated nuclei in the thalamus of Polypterus. There is only one circumscribed projection into the thalamus, i.e., the optic tract, but there are numerous diffuse fibers terminating in this region of the brain. The hypothalamus, except for a partially migrated nucleus, has retained the periventricular arrangement of cells. It has large fiber connections with the forebrain and brainstem. The literature on the diencephalon of lower forms has been reviewed with special emphasis on the question of how homologies are established in this brainpart. It appeared that three different criteria, either singly or in combination, have been employed as a clue to identification of structures in the diencephalon. These are, (1) ventricular grooves, (2) nuclear boundaries, and (3) fiber connections. In order to test the practical validity of these criteria the diencephalon of Polypterus was compared to that of five related species, i.e., the actinopterygians Acipenser and Polyodon, the dipnoans Protopterus and Neoceratodus, and the crossopterygian Latimeria. In addition three amphibians, Necturus, Ambystoma and Rana, were involved in our comparative considerations. It was concluded that, within the confines of the diencephalon of the species mentioned, cytoarchitectural differences form the most valid criterion for establishing homologies. The drawback and restrictions connected with the use of ventricular sulci and fiber connections, as a clue to identification have been evaluated and discussed.     

The myth of dorsal ribs in gnathostome vertebrates

Traditionally, two types of rib are distinguished in gnathostomes: dorsal (upper) and ventral (lower, pleural) ribs. They are defined according to their position in the connective tissue system of the body: dorsal ribs develop at the intersection of the serially arranged myosepta with the horizontal septum that separates epaxial from hypaxial musculature, whereas ventral ribs develop at the intersection of myosepta with the peritoneum and usually encircle the body cavity. Distribution of rib types among gnathostomes has traditionally been reported as follows: elasmobranchs have dorsal ribs; all Actinopterygii have only ventral ribs with the exception of polypterids, and two subgroups of teleosts, which supposedly also have dorsal ribs; within Sarcopterygii tetrapods have dorsal ribs, whereas dipnoans have ventral ribs. Here, we report the development of ribs in polypterids, a taxon playing a crucial role in discussions on rib homology. We demonstrate that putative dorsal ribs of polypterids have a unique ontogeny and represent an autapomorphy of this taxon. We discuss previous hypotheses of rib homology and offer a more plausible (i.e. more parsimonious) alternative to the conventional interpretation. We conclude that dorsal ribs do not exist and that ribs of gnathostomes are ventral ribs.     

HEADS AND TAILS: A CHORDATE PHYLOGENY

Abstract— A cladistic analysis of chordates is presented, based on some 320 nested characters. All the principal higher taxa are defined by synapomorphies, including extinct acanthodians and placoderms. The data base draws broadly from adult anatomy (including osteological data for Recent and fossil taxa), embryology, physiology, and biochemistry. A conventional sequence of chordate higher taxa is generated (hemichordates, urochordates, cephalochordates, craniates). Among the craniates, cyclostomes are considered paraphyletic. Gnathostomes are monophyletic, but two fossil "agnathan" groups (galeaspids, osteostracans) are regarded as stem gnathostomes. Chondrichthyans and osteichthyans are monophyletic. New arguments for osteichthyan affinity of acanthodians are presented. The phylogenetic position of placoderms is still problematic, but they can no longer be perceived as stem chondrichthyans or even as "elasmobranchiomorphs." Recent dipnoans and tetrapods are sister groups, but new paleontological discoveries refute many of their supposed osteological synapomorphies, thereby reopening the possibility of a closer relationship between tetrapods and osteolepiform rhipidistians.