CHARACTER DIAGNOSIS, FOSSILS AND THE ORIGIN OF TETRAPODS

I. The traditional view of the origin of tetrapod vertebrates is that they are descendants of fossil osteolepiform fish, of which Eusthenopteron is best known. In recent years both that conclusion and the methodology by which it has been reached have been challenged by practitioners of cladistic analysis. Particularly a recent review by Rosen et al. (1981) claims that Dipnoi (lungfish) are the sister-group of the Tetrapoda, that Osteolepiformes is a non-taxon and that Eusthenopteron is more distant from tetrapods than are Dipnoi, coelacanths and probably the fossil Porolepiformes. We attempt to refute all these concludions by use of the same cladistic technique. 2. We accept that all the above-mentioned groups, together with some less well-known taxa, can be united as Sarcopterygii by means of shared derived (apomorph) characters. We also agree that Porolepiformes and Actinistia (coelacanths) can be characterized as valid taxa. The primitive and enigmatic fossil fish Powichthys is accepted as representing the plesiomorph sister-group of true porolepiforms. 3. Only two apomorph features, the course of the jaw adductor muscles and the position of incurrent and excurrent nostrils, appear to unite all the fish, living and fossil, currently regarded as Dipnoi. The characteristic tooth plates and the presence of petrodentine both exclude important primitive fossil forms. 4. Contrary to the opinion of Rosen et al., Osteolepiformes can be characterized — by the arrangement of bones forming the cheek plate, the presence of basal scutes to the fins and by the unjointed radials of the median fins. However, if these are true autapomorphies they exclude any osteolepiform from direct tetrapod ancestry. 5. Tetrapoda is a monophyletic group characterized by ten or more autapomorphies, including the bones of the cheek plate, a stapes and fenestra ovalis, and a series of characters of the appendicular skeleton. 6. Tetrapods have a true choana (internal nostril). We accept that the posterior (excurrent) nostril of Dipnoi is the homologue of the tetrapod choana. However, we assert that the posterior nostril of all bony fish is the homologue of the choana. This assertion would be refuted if any fish showed separate posterior nostril and choana. We reject the claim that this ‘three nostril condition’ occurred in porolepiforms and osteolepiforms. The evidence for a choana in porolepiforms is inadequate. Osteolepiforms had a true choana, characterized as in tetrapods by its relationship to the bones of the palate, but no third nostril. Dipnoans are not choanate. 7. Following cladistic practice, the relationship of the extant taxa is established first. Dipnoi are thus shown to be the living sister-group of tetrapods, but only on ‘soft anatomy’ characters unavailable in fossils. Coelacanths are the living sister-group of the taxon so formed. 8. The relationship of the fossil taxa to the extant sarcopterygians is then considered. The synapomorphy scheme proposed by Rosen et al. is discussed at length. Virtually all the characters they use to exclude close relationship of Eusthenopteron (and hence all osteolepiforms) to tetrapods, in favour of coelacanths and dipnoans, are invalid. 9. A series of synapomorphies uniting osteolepiforms and tetrapods is proposed, including a true choana (hence the taxon Choanata), the histology of the teeth, and a number of characters of the humerus. The recently discovered fossil Youngolepis, which lacks a choana, represents the sister-group of the Choanata, and is not uniquely close to Powichthys. The latter, as a porolepiform (s.l.) is a member of the sister-group to Choanata plus Youngolepis. 10. Our cladistic analysis suggests that all the extinct taxa considered are more closely related to tetrapods than are the Dipnoi. Moreover fossil evidence suggests that Dipnoi, considered as an extant taxon, may not even be the living sister-group of Tetrapoda. Early fossil dipnoans appear to have been marine fish without specific adaptations for air breathing. If so the apparent synapomorphies of Dipnoi and Tetrapoda may be homoplastic — the insistence on grouping extant taxa first would then have yielded an invalid inference.     

Molecules,fossils, and the origin of tetrapods

Summary Since the discovery of the coelacanth, Latimeria chalumnae, more than 50 years ago, paleontologists and comparative morphologists have debated whether coelacanths or lungfishes, two groups of lobe-finned fishes, are the closest living relatives of land vertebrates (Tetrapoda). Previously, Meyer and Wilson (1990) determined partial DNA sequences from two conservative mitochondrial genes and found support for a close relationship of lungfishes to tetrapods. We present additional DNA sequences from the 12S rRNA mitochondria gene for three species of the two lineages of lungfishes that were not represented in the first study: Protopterus annectens and Protopterus aethiopicus from Africa and Neoceratodus forsteri (kindly provided by B. Hedges and L. Maxson) from Australia. This extended data set tends to group the two lepidosirenid lungfish lineages (Lepidosiren and Protopterus) with Neoceratodus as their sister group. All lungfishes seem to be more closely related to tetrapods than the coelacanth is. This result appears to rule out the possibility that the coelacanth lineage gave rise to land vertebrates. The common ancestor of lungfishes and tetrapods might have possessed multiple morphological traits that are shared by lungfishes and tetrapods [Meyer and Wilson (1990) listed 14 such traits]. Those traits that seem to link Latimeria and tetrapods are arguably due to convergent evolution or reversals and not to common descent. In this way, the molecular tree facilitates an evolutionary interpretation of the morphological differences among the living forms. We recommended that the extinct groups of lobe-finned fishes be placed onto the molecular tree that has lungfishes and not the coelacanth more closely related to tetrapods. The placement of fossils would help to further interpret the sequence of morphological events and innovations associated with the origin of tetrapods but appears to be problematic because the quality of fossils is not always high enough, and differences among paleontologists in the interpretation of the fossils have stood in the way of a consensus opinion for the branching order among lobefinned fishes. Marshall and Schultze (1992) criticized the morphological analysis presented by Meyer and Wilson (1990) and suggest that 13 of the 14 morphological traits that support the sister group relationship of lungfishes and tetrapods are not shared derived characters. Here we present further alternative viewpoints to the ones of Marshall and Schultze (1992) from the paleontological literature. We argue that all available information (paleontological, neontological, and molecular data) and rigorous cladistic methodology should be used when relating fossils and extant taxa in a phylogenetic framework. Offprint requests to: Axel Meyer     

Gene Structure and Amino Acid Sequence of Latimeria chalumnae (Coelacanth) Myelin DM20: Phylogenetic Relation of the Fish

The structure of Latimeria chalumnae (coelacanth) proteolipid protein/DM20 gene excluding exon 1 was determined, and the amino acid sequence of Latimeria DM20 corresponding to exons 2–7 was deduced. The nucleotide sequence of exon 3 suggests that only DM20 isoform is expressed in Latimeria. The structure of proteolipid protein/DM20 gene is well preserved among human, dog, mouse, and Latimeria. Southern blot analysis indicates that Latimeria DM20 gene is a single-copy gene. When the amino acid sequences of DM20 were compared among various species, Latimeria was more similar to tetrapods than other fishes including lungfish, confirming the previous finding by immunoreactivity (Waehneldt and Malotka 1989 J. Neurochem. 52:1941–1943). However, when phylogenetic trees were constructed from the DM20 sequences, lungfish was clearly the closest to tetrapods. Latimeria was situated outside of lungfish by the maximum likelihood method. The apparent similarity of Latimeria DM20 to tetrapod proteolipid protein/DM20 is explained by the slow amino acid substitution rate of Latimeria DM20.     

Fish and tetrapod communities across a marine to brackish salinity gradient in the Pennsylvanian (early Moscovian) Minto Formation of New Brunswick,Canada, and their palaeoecological and palaeogeographical implications

Euryhaline adaptations in Pennsylvanian vertebrates allowed them to inhabit the marine to freshwater spectrum. This is illustrated by new assemblages of fish and tetrapods from the early Moscovian Minto Formation of New Brunswick, Canada. Fish include chondrichthyans (xenacanthids and the enigmatic Ageleodus), acanthodians (gyracanthids and acanthodiforms), sarcopterygians (rhizodontids, megalichthyids and dipnoans), and actinopterygians (eurynotiforms). Tetrapods include small‐ to medium‐sized, and largely aquatic, stem tetrapods (colosteids) and anthracosaurs (embolomeres). A key finding is that the parautochthonous fossil assemblages are preserved across a salinity gradient, with diversity (measured by the Simpson Index) declining from open marine environments, through brackish embayments, and reaching a nadir in tidal estuaries. Chondrichthyans dominate the entire salinity spectrum (65% of fossils), a distribution that demonstrates a euryhaline mode of life, and one large predatory chondrichthyan, Orthacanthus, may have practised filial cannibalism in coastal nurseries because its heteropolar coprolites contain juvenile xenacanthid teeth. In contrast, other fish communities were more common in open marine settings while tetrapods were more common in coastal brackish waters. While all these faunas were also likely to have been euryhaline, their osmoregulation was, perhaps, less versatile. The demonstration of widespread euryhalinity among fish and aquatic tetrapods explains why Pennsylvanian faunas generally show a cosmopolitan biogeography because taxa were able to disperse via seaways. It also resolves the paradox of enriched strontium isotopic signatures observed in these faunas because organisms would have been, at times, exposed to continental water bodies as well. Therefore, our new findings contribute to the long‐running debate about the ecology of Pennsylvanian fishes and tetrapods.     

The evolution of tenascins and fibronectin

Tenascins are extracellular matrix glycoproteins that act both as integrin ligands and as modifiers of fibronectin-integrin interactions to regulate cell adhesion, migration, proliferation and differentiation. In tetrapods, both tenascins and fibronectin bind to integrins via RGD and LDV-type tripeptide motifs found in exposed loops in their fibronectin-type III domains. We previously showed that tenascins appeared early in the chordate lineage and are represented by single genes in extant cephalochordates and tunicates. Here we have examined the genomes of the coelacanth Latimeria chalumnae, the elephant shark Callorhinchus milii as well as the lampreys Petromyzon marinus and Lethenteron japonicum to learn more about the evolution of the tenascin gene family as well as the timing of the appearance of fibronectin during chordate evolution. The coelacanth has 4 tenascins that are more similar to tetrapod tenascins than are tenascins from ray-finned fishes. In contrast, only 2 tenascins were identified in the elephant shark and the Japanese lamprey L. japonicum. An RGD motif exposed to integrin binding is observed in tenascins from many, but not all, classes of chordates. Tetrapods that lack this RGD motif in tenascin-C have a similar motif in the paralog tenascin-W, suggesting the potential for some overlapping function. A predicted fibronectin with the same domain organization as the fibronectin from tetrapods is found in the sea lamprey P. marinus but not in tunicates, leading us to infer that fibronectin first appeared in vertebrates. The motifs that recognize LDV-type integrin receptors are conserved in fibronectins from a broad spectrum of vertebrates, but the RGD integrin-binding motif may have evolved in gnathostomes.     

On the structure of the lower jaw in dipnoans: with a description of an early Devonian dipnoan from Canada, Melanognathus canadenis gen. et sp.nov.

The lower jaw in Melanognathus gen. nov. and several other Devonian dipnoans is described and compared with that in Neoceratodus. It is a most conservative structure, which apart from the effect of the retrogressive development of the skeleton has hardly changed since Devonian times. A new interpretation of the sensory canal bones is given. With respect to these elements and the structure of the lower jaw as a whole, the dipnoans differ fundamentally from the rhipidistid crossopterygians, tetrapods and aetinopterygians. Several resemblances to the lower jaw in holocephalians are demonstrated. The results confirm the view that the Dipnoi comprise an early specialized and isolated group, perhaps more closely related to the elasmo-branchiomorphs than to the teleostomes and tetrapods.     

Relationship among coelacanths,lungfishes, and tetrapods: A phylogenetic analysis based on mitochondrial cytochrome oxidase I gene sequences

To clarify the relationship among coelacanths, lungfishes, and tetrapods, the amino acid sequences deduced from the nucleotide sequences of mitochondrial cytochrome oxidase subunit I (COI) genes were compared. The phylogenetic tree of these animals, including the coelacanth Latimeria chalumnae and the lungfish Lepidosiren paradoxa, was inferred by several methods. These analyses consistently indicate a coelacanth/lungfish clade, to which little attention has been paid by previous authors with the exception of some morphologists. Overall evidence of other mitochondrial genes reported previously and the results of this study equally support the coelacanth/lungfish and lungfish/tetrapod clades, ruling out the coelacanth/tetrapod clade.Correspondence to: K. Watanabe 0592     

The First Virtual Cranial Endocast of a Lungfish (Sarcopterygii: Dipnoi)

Lungfish, or dipnoans, have a history spanning over 400 million years and are the closest living sister taxon to the tetrapods. Most Devonian lungfish had heavily ossified endoskeletons, whereas most Mesozoic and Cenozoic lungfish had largely cartilaginous endoskeletons and are usually known only from isolated tooth plates or disarticulated bone fragments. There is thus a substantial temporal and evolutionary gap in our understanding of lungfish endoskeletal morphology, between the diverse and highly variable Devonian forms on the one hand and the three extant genera on the other. Here we present a virtual cranial endocast of Rhinodipterus kimberleyensis, from the Late Devonian Gogo Formation of Australia, one of the most derived fossil dipnoans with a well-ossified braincase. This endocast, generated from a Computed Microtomography (µCT) scan of the skull, is the first virtual endocast of any lungfish published, and only the third fossil dipnoan endocast to be illustrated in its entirety. Key features include long olfactory canals, a telencephalic cavity with a moderate degree of ventral expansion, large suparaotic cavities, and moderately enlarged utricular recesses. It has numerous similarities to the endocasts of Chirodipterus wildungensis and Griphognathus whitei, and to a lesser degree to ''Chirodipterus'' australis and Dipnorhynchus sussmilchi. Among extant lungfish, it consistently resembles Neoceratodus more closely than Lepidosiren and Protopterus. Several trends in the evolution of the brains and labyrinth regions in dipnoans, such as the expansions of the utricular recess and telencephalic regions over time, are identified and discussed.     

Distribution patterns of the paraneuronal endocrine cells in the skin, gills and the airways of fishes as determined by immunohistochemical and histological methods

Summary The neuro-endocrine cells of fish skin and respiratory surfaces, and their bioactive secretion as far as is known, are reviewed, and compared with similar elements in tetrapods, particularly amphibians. In the skin of teleost fish, immunohistochemistry has shown that Merkel cells react for serotonin, neuron-specific enolase and enkephalins. The pharmacology is not established in dipnoans or lampreys. In some teleosts, neuromasts react for substance P and leu-enkephalins; substance P is also reported from some ampullary organs (electroreceptors). Taste buds of teleosts may react for enkephalin and substance P. Basal cells of taste buds react for serotonin and neuron-specific enolase. Some unicellular skin glands of teleosts express bioactive compounds, including serotonin and some peptides; this ectopic expression is paralleled in amphibian skin glands. The dipnoan Protopterus has innervated pulmonary neuro-endocrine cells in the pneumatic duct region with dense-cored vesicles. In Polypterus and Amia the lungs have serotonin-positive neuro-endocrine cells that are apparently not innervated. In fish gills, a closed type of neuro-endocrine cell reacts for serotonin, an open type for enkephalins and some calcium-binding proteins (calbindin, calmodulin and S-100 protein). The functions of neuro-endocrine cells in fishes await investigation, but it is assumed they are regulatory.     

Interspecies Insertion Polymorphism Analysis Reveals Recent Activity of Transposable Elements in Extant Coelacanths

Coelacanths are lobe-finned fish represented by two extant species, Latimeria chalumnae in South Africa and Comoros and L. menadoensis in Indonesia. Due to their intermediate phylogenetic position between ray-finned fish and tetrapods in the vertebrate lineage, they are of great interest from an evolutionary point of view. In addition, extant specimens look similar to 300 million-year-old fossils; because of their apparent slowly evolving morphology, coelacanths have been often described as « living fossils ». As an underlying cause of such a morphological stasis, several authors have proposed a slow evolution of the coelacanth genome. Accordingly, sequencing of the L. chalumnae genome has revealed a globally low substitution rate for protein-coding regions compared to other vertebrates. However, genome and gene evolution can also be influenced by transposable elements, which form a major and dynamic part of vertebrate genomes through their ability to move, duplicate and recombine. In this work, we have searched for evidence of transposition activity in coelacanth genomes through the comparative analysis of orthologous genomic regions from both Latimeria species. Comparison of 5.7 Mb (0.2%) of the L. chalumnae genome with orthologous Bacterial Artificial Chromosome clones from L. menadoensis allowed the identification of 27 species-specific transposable element insertions, with a strong relative contribution of CR1 non-LTR retrotransposons. Species-specific homologous recombination between the long terminal repeats of a new coelacanth endogenous retrovirus was also detected. Our analysis suggests that transposon activity is responsible for at least 0.6% of genome divergence between both Latimeria species. Taken together, this study demonstrates that coelacanth genomes are not evolutionary inert: they contain recently active transposable elements, which have significantly contributed to post-speciation genome divergence in Latimeria.     

Structure of the kidney in the coelacanth Latimeria chalumnae with reference to osmoregulation

The morphology of the nephrons of the coelacanth Latimeria chalumnae was investigated by light microscopy. Each nephron is composed of a large renal corpuscle with well‐vascularized glomerulus, non‐ciliated neck segment, proximal convoluted tubule divided into distinct first and second segments, non‐ciliated intermediate segment, distal tubule, collecting tubule and collecting duct. The parietal layer of the Bowman's capsule of the renal corpuscle is composed of low cuboidal cells. The short non‐ciliated neck segment is lined by cuboidal epithelium. The first and second proximal segments display a prominent brush border and contain amorphous material in their lumen. The second proximal segment differs from the first segment in having taller columnar epithelium and a relatively narrow lumen. The intermediate segment is lined by non‐ciliated columnar epithelium and its lumen appears empty. The distal tubule is narrow in diameter and its cuboidal epithelium is devoid of intercalated cells. A unique feature of L. chalumnae is having binucleate cells in the tubule and collecting duct epithelium. The renal arteries have poorly developed tunica media and its cells contain granular material. The structure of L. chalumnae nephrons correlates well with their osmoregulatory function and resembles those of euryhaline teleosts.     

A new genus of Devonian tetrapod from North‐East Greenland,with new information on the lower jaw of Ichthyostega

Abstract: A new genus and species of Devonian tetrapod has been identified from material collected in 1947 from the southern slope of Mt. Celsius, Ymer Ø, North‐East Greenland. The specimen preserves both lower jaws, partial palate, premaxillae and maxillae, with a natural mould of parts of the shoulder girdle. The new taxon, Ymeria denticulata, shows differences in dentition, skull ornament and lateral line expression from both Acanthostega and Ichthyostega, but it shows a closer resemblance to the latter. A cladistic analysis not only suggests that Ymeria lies adjacent to Ichthyostega on the tetrapod stem, but also reveals substantial topological instability. As the third genus and the fifth species of tetrapod identified from North‐East Greenland, it demonstrates the high diversity of Devonian tetrapods in that region.     

The structure and stratigraphy of<Emphasis Type="Italic">Speonesydrion</Emphasis> from New South Wales,Australia, and the dentition of primitive dipnoans

New material ofSpeonesydrion iani, an Early Devonian dipnoan from New South Wales, has provided additional Information on the dentition and jaws. Two new partial palates have been found, and X-rays of the parasphenoid shows that the structure is well preserved. The palatal teeth are well worn even in partly grown material, and they do not originate at a growth point, but at a thickening of the palate. More mandibles have been collected, and thin sections have been prepared to allow a discussion of their histology. On the mandible the teeth are clear, and they are much more defined than they are on the palate. The dental heel is variably developed, and grows in phases by thickening of the dentine at the contact with the bone. Dentine forms on the bone at the base of the heel, partly by Solution of the bone and the addition of dentine from the pulp canals, but also by direct growth from the pulp canals dorsal to the bone. In the latter case the dentine and bone are in contact, and the two tissues intermingle. The teeth are also formed on a thickened bone and consist of dentine capped with enamel making a crest. Dentine and bone are related as in the heel. We conclude that the teeth inSpeonesydrion are not homologous with the teeth in other dipnoans, and are formed by a different process involving the aggregation of denticles.     

The Changing State of Surfactant Lipids: New Insights from Ancient Animals

SYNOPSIS. Pulmonary surfactant is a mixture of phospholipids(including disaturated phospholipids), cholesterol and proteinslining the air-liquid interface within the lung. Surfactantacts to reduce surface tension, thereby increasing lungcomplianceand also preventing edema. The saccular lungs, or other gas-holdingstructures, of nonmammals have 7–70% more surfactant/cm²of surface than lungs of mammals. Nonmammalian surfactant actsas an antiglue that decreases the inflation pressures of collapsedlungs by reducing the adherence of apposing epithelial surfaces.The autonomic nervous system appears to be the primary systemcontrolling release of surfactant in nonmammals. The lipid compositionis highly conserved within the vertebrates, except that surfactantof teleost fish is dominated by cholesterol whereas tetrapodsurfactant consists primarily of disaturated phospholipids (DSP).The dipnoan Neoceratodus forsteri demonstrates a "fish-type"surfactant profile while the other derived dipnoans demonstratea surfactant profile similar to that of tetrapods. Homologyof the surfactant protein SP-A within the vertebrates pointsto a single evolutionaryorigin for the system and indicatesthat fish surfactant is a "protosurfactant". Amongst the tetrapods,the relative proportions of DSP and cholesterol vary in responseto lung structure, habitat, and body temperature (T_b) but notin relation to phytogeny. The cholesterol content of surfactantis elevated in species with simple saccular lungs, in aquaticspecies, and in species with low T_b. The DSP content is highestin complex lungs, particularly ofaquatic species or specieswith high T_b. The cholesterol content of surfactant also increasesin response to acute decreases in Tb in lizards and torpid marsupials,presumably to maintain fluidity of the lipid mixture.     

Skeletal Morphogenesis of Microbrachis and Hyloplesion (Tetrapoda: Lepospondyli), and Implications for the Developmental Patterns of Extinct,Early Tetrapods

The ontogeny of extant amphibians often is used as a model for that of extinct early tetrapods, despite evidence for a spectrum of developmental modes in temnospondyls and a paucity of ontogenetic data for lepospondyls. I describe the skeletal morphogenesis of the extinct lepospondyls Microbrachis pelikani and Hyloplesion longicostatum using the largest samples examined for either taxon. Nearly all known specimens were re-examined, allowing for substantial anatomical revisions that affect the scoring of characters commonly used in phylogenetic analyses of early tetrapods. The palate of H. longicostatum is re-interpreted and suggested to be more similar to that of M. pelikani, especially in the nature of the contact between the pterygoids. Both taxa possess lateral lines, and M. pelikani additionally exhibits branchial plates. However, early and rapid ossification of the postcranial skeleton, including a well-developed pubis and ossified epipodials, suggests that neither taxon metamorphosed nor were they neotenic in the sense of branchiosaurids and salamanders. Morphogenetic patterns in the foot suggest that digit 5 was developmentally delayed and the final digit to ossify in M. pelikani and H. longicostatum. Overall patterns of postcranial ossification may indicate postaxial dominance in limb and digit formation, but also more developmental variation in early tetrapods than has been appreciated. The phylogenetic position and developmental patterns of M. pelikani and H. longicostatum are congruent with the hypothesis that early tetrapods lacked metamorphosis ancestrally and that stem-amniotes exhibited derived features of development, such as rapid and complete ossification of the skeleton, potentially prior to the evolution of the amniotic egg.     

Ciliated Sensory Cells in the Rostral Organ of the Coelacanth Latimeria chalumnae (Smith 1939)

The rostral organ of the coelacanth Latimeria chalumnae contains a sensory epithelium with luminated ciliated sensory cells, which very much resemble electroreceptors from other non-teleost fish. This, as well as the innervation and a dorsal nucleus, indicate that this interesting species is electroreceptive.     

Settlement of seaweeds on coastal structures

Coastal structures are constructed principally to protect the coast line. However, these structures also can act as artificial substrata for seaweeds. In particular, armor blocks, such as tetrapods, prove to be good algal substrata. Our field observations on the vegetation and standing crop of seaweed communities on armor blocks led us to the following conclusions: 1) Ecklonia cavacommunities grew on tetrapods that had been placed to coincide with the maturation period of E. cava. The communities have been maintained for more than four years. 2) The standing crop of seaweeds on an offshore breakwater composed of tetrapods placed seven years before was almost the same as that found under natural conditions. 3) Variations in roughened surfaces (pebbles or scores and grooves 3 or 46 mm wide) did not affect the growth of Sargassum spp. However, a greater number of Ecklonia stolonifera plants were observed attached to roughened surfaces compared to smooth surfaces.     

Stomach contents of Latimeria chalumnae and further notes on its feeding habits

Synopsis Stomach contents of three Latimeria chalumnae dissected in Japan support the hypothesis that the coelacanth is a predominantly nocturnal bottom or near-bottom drift feeder. Prey items identified in this study (Ilyophis brunneus, Cephaloscyllium sufflans and Beryx decadactylus) and those reported previously are mostly benthic or epibenthic dwellers. The drifting of the coelacanth in a headstanding posture would account for the easy capture of prey that moves within or just above the bottom strata at night.     

The fishery of the Comoros,with comments on its possible impact on coelacanth survival

Synopsis The traditional methods of deepsea handline fishing in the Comoros are described. The main target species is the oilfish Ruvettus pretiosus, and the coelacanth Latimeria chalumnae is caught as a bycatch. In recent years motorised dugout canoes as well as outboard- and inboard-powered boats have been introduced into the fishery, and more efficient fishing tackle has become available. The more modern gear is intended for use on pelagic fishes and has added a new dimension to the fishery. Traditional handline fishermen are not considered to be a threat to the coelacanth, but the fishermen equipped with motorised boats and modern tackle would constitute a real threat if they direct their efforts on inshore reefs.     

The relationships of the coelacanth Latimeria chalumnae: evidence from sequences of vertebrate 28S ribosomal RNA genes

Synopsis A subgenomic library created from genomic DNA of Latimeria chalumnae was screened for 28S ribosomal RNA (rRNA) clones. The resulting clone was subcloned into a plasmid vector, and over 2 kb of the 28S rRNA region was sequenced. Sequences of 28S rRNA genes were also obtained for Rhineura floridana (Squamata), Cyprinella lutrensis (Actinopterygii), and Lampetra aepyptera (Petromyzontiformes) by cloning and/or amplification by the polymerase chain reaction. The 28S rDNA sequences were aligned for all the above species as well as for the previously published 28S rDNA sequences of the genera Mus, Rattus, and Homo (Mammalia), Xenopus (Amphibia), and Drosophila (Insecta). Phylogenetic analysis of these species (using both the insect and lamprey sequences for outgroup comparison, or using only the lamprey sequence in the outgroup) produced a single optimal solution: (Outgroup(Cyprinella(Latimeria(Xenopus(Rhineura(Homo(Rattus(Mus)))))))). Bootstrap analysis indicated that the placement of L. chalumnae on this tree was significant at p < 0.01. Previously published alternative hypotheses of relationships of Latimeria require at least 19 additional steps compared to the optimal solution; the rDNA data are sufficient to reject the hypotheses that place Latimeria in groups other than the sarcopterygians.