Book reviewFishes of the North-eastern Atlantic and the Mediterranean: Volume II. P.J.P. Whitehead, M.-L. Bauchot, J.-C. Hureau, J. Nielsen, E. Tortonese (Editors). UNESCO Press, Paris, 1986, pp. 510–1007, FF 250, Dfl 105.00, ISBN 92-3-002308-6

A checklist of Recent and fossil catfishes (Order Siluriformes) is presented, summarizing taxonomic literature published through 2005. From 4624 nominal species group names and 810 genus group names, 3093 species are recognized as valid, and are distributed among 478 genera and 36 families. Distributional summaries are provided for each species, and nomenclatural synonymies, including relevant information on all name-bearing types, are included for all taxa. One new name is proposed herein: Clariallabes teugelsi, as a replacement for Clarias (Allabenchelys) dumerili longibarbis David & Poll, 1937, which is preoccupied by Clarias longibarbis Worthington, 1933, but has been treated as a valid species of Clariallabes by Teugels. Acrochordonichthys melanogaster Bleeker, 1854, is designated as type species of Acrochordonichthys Bleeker, 1857, inasmuch as no earlier valid designation has been found. A new genus Pseudobagarius , is proposed for the “ pseudobagarius group” of species formerly placed in Akysis . The status of 228 species group names remains unresolved and 31 names based on otoliths ascribed to catfishes are listed but not placed into the checklist. The current emphasis given to catfish taxonomy at present is likely to result in a dramatic increase in the total number of valid taxa as well as major changes in the membership of some of the higher level taxa recognized here.

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Checklist of catfishes, recent and fossil (Osteichthyes: Siluriformes), and catalogue of siluriform primary types

Because of the genetic and developmental information available, Danio rerio stands out as a vertebrate model system in which significant progress in the areas of development and evolution can be made. Despite its increasing popularity, little research has been done on skeletal development. In this report, we provide developmental information on the structure and composition of the zebrafish skull, pectoral, and pelvic girdle. We describe the sequence of ossification of the skull and paired fins from a large series of cleared and Alizarin red-stained specimens at larval and adult stages. The most commonly followed developmental sequence in Danio rerio is described. Chondrocranial development is noted from Alcian blue-stained specimens. General trends in ossification patterns are examined from developmental, phylogenetic, and functional contexts. No clear pattern in ossification order of dermal versus cartilage bones is evident. Ossification sequence conforms to functional need in a general way, but there are inconsistencies in the details of order. Selected phylogenetic comparisons of ossification sequence within cranial regions are made among Danio rerio, Betta splendens, Oryzias latipes, and Barbus barbus. Greater sequence conservation is apparent between D. rerio and Barbus barbus, the ostariophysans, than among other taxon pairs. Intraspecific variation in ossification order is apparent, most of which involves small adjustments in timing. © 1996 Wiley-Liss, Inc.

Development of the cranium and paired fins in the zebrafish Danio rerio (Ostariophysi, Cyprinidae).

This review deals with the following seven aspects of vertebrate skeletogenic and odontogenic tissues. 1. The evolutionary sequence in which the tissues appeared amongst the lower craniate taxa. 2. The topographic association between skeletal (cartilage, bone) and dental (dentine, cement, enamel) tissues in the oldest vertebrates of each major taxon. 3. The separate developmental origin of the exo- and endoskeletons. 4. The neural-crest origin of cranial skeletogenic and odontogenic tissues in extant vertebrates. 5. The neural-crest origin of trunk dermal skeletogenic and odontogenic tissues in extant vertebrates. 6. The developmental processes that control differentiation of skeletogenic and odontogenic tissues in extant vertebrates. 7. Maintenance of developmental interactions regulating skeletogenic/odontogenic differentiation across vertebrate taxa. We derive twelve postulates, eight relating to the earliest vertebrate skeletogenic and odontogenic tissues and four relating to the development of these tissues in extant vertebrates and extrapolate the developmental data back to the evolutionary origin of vertebrate skeletogenic and odontogenic tissues. The conclusions that we draw from this analysis are as follows. 8. The dermal exoskeleton of thelodonts, heterostracans and osteostracans consisted of dentine, attachment tissue (cement or bone), and bone. 9. Cartilage (unmineralized) can be inferred to have been present in heterostracans and osteostracans, and globular mineralized cartilage was present in Eriptychius, an early Middle Ordovician vertebrate unassigned to any established group, but assumed to be a stem agnathan. 10. Enamel and possibly also enameloid was present in some early agnathans of uncertain affinities. The majority of dentine tubercles were bare. 11. The contemporaneous appearance of cellular and acellular bone in heterostracans and osteostracans during the Ordovician provides no clue as to whether one is more primitive than the other. 12. We interpret aspidin as being developmentally related to the odontogenic attachment tissues, either closer to dentine or a form of cement, rather than as derived from bone. 13. Dentine is present in the stratigraphically oldest (Cambrian) assumed vertebrate fossils, at present some only included as Problematica, and is cladistically primitive, relative to bone. 14. The first vertebrate exoskeletal skeletogenic ability was expressed as denticles of dentine. 15. Dentine, the bone of attachment associated with dentine, the basal bone to which dermal denticles are fused and cartilage of the Ordovician agnathan dermal exoskeleton were all derived from the neural crest and not from mesoderm. Therefore the earliest vertebrate skeletogenic/odontogenic tissues were of neural-crest origin.(ABSTRACT TRUNCATED AT 400 WORDS)

Development and evolutionary origins of vertebrate skeletogenic and odontogenic tissues.

The muscles that power vertebrate locomotion are associated with springy tissues, both within muscle and in connective tissue elements such as tendons. These springs share in common the same simple action: they stretch and store elastic strain energy when force is applied to them and recoil to release energy when force decays. Although this elastic action is simple, it serves a diverse set of functions, including metabolic energy conservation, amplification of muscle power output, attenuation of muscle power input, and rapid mechanical feedback that may aid in stability. In recent years, our understanding of the mechanisms and importance of biological springs in locomotion has advanced significantly, and it has been demonstrated that elastic mechanisms are essential for the effective function of the muscle motors that power movement. Here, we review some recent advances in our understanding of elastic mechanisms, with an emphasis on two proposed organizing principles. First, we review the evidence that the various functions of biological springs allow the locomotor system to operate beyond the bounds of intrinsic muscle properties, including metabolic and mechanical characteristics, as well as motor control processes. Second, we propose that an energy-based framework is useful for interpreting the diverse functions of series-elastic springs. In this framework, the direction and timing of the flow of energy between the body, the elastic element and the contracting muscle determine the function served by the elastic mechanism (e.g. energy conservation vs power amplification). We also review recent work demonstrating that structures such as tendons remodel more actively and behave more dynamically than previously assumed.

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Flexible mechanisms: the diverse roles of biological springs in vertebrate movement

The ontogeny of the bony skull of the African catfish, Clarias gariepinus, is studied from initial ossification until a complete skull is formed. The ossification sequence in C. gariepinus seems to be related to the func- tional demands that arise in a developing larva. Early ossification of the opercular bone coincides with the initiation of opercular skin movements. Early ossifications involve several dentulous bones, formed shortly before the transition phase from endogenous to exogenous feeding. The enlarging bran- chiostegal membrane becomes supported by the gradual adding of branchio- stegal rays. Parasphenoid ossification may be related to protection of the brain during prey transport, whereas the several hyoid bones, including the parurohyal, are formed in relation to the increasing loads exerted onto the tendons of the sternohyoideus and consequently onto the hyoid bar. Overall skull reinforcement occurs almost simultaneously, with a whole set of perichon- dral bones arising especially at places of high mechanical load. The suspenso- rium becomes protected against dislocation in an anteroposterior direction through a ligamentous connection, which even becomes partially ossified, forming the sesamoid entopterygoid. Later, the cranial lateral-line system becomes enclosed by a set of gutters, which close, frequently becoming plate-like later in ontogeny. The brain also becomes covered dorsally. Addi- tional dentition (prevomeral tooth plates) formation seems to coincide with formation of the opercular four-bar system, as well as with the time the digestive system becomes completely functional. Eventually, unossified re- gions between the bones become closed off, fortifying and completely covering the skull. J. Morphol. 235:183-237, 1998. r 1998 Wiley-Liss, Inc. The osteology of catfishes has been a basis for several studies, especially for taxonomy and phylogeny. In the present study, the taxonomy follows that of Fink and Fink ('96), in which the Siluriformes, or catfish, repre- sent the sister group of the Gymnotiformes, or knife-fish. Both are grouped within the Siluriphysi, which together with the Char- aciphysi and Cypriniphysi comprise the oto- physan group of the Ostariophysi. An exten- sive literature has been produced on the descriptive cranial osteology of siluriform fish, mostly on adult forms (e.g., McMurrich, 1884; Bhimachar, '33; Merriman, '40; Hubbs and Miller, '60; Tilak, '61, '63a,b, '64, '65a,b, '67, '71; Rastogi, '63; Gauba, '66; '70; Taverne and Aloulou-Triki, '74; Lundberg, '75, '82; Srinivasa Rao and Lakshmi, '84; Vignes and Garcia, '87; de Pinna, '88; Howes and Fumi-

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Ontogeny of the osteocranium in the African catfish, Clarias gariepinus burchell (1822) (Siluriformes: Clariidae): Ossification sequence as a response to functional demands

Suction feeding in Astatotilapia elegans occurs by a series of rapid, coupled movements of various head parts. The lower jaw rotates with respect to the neurocranium through an angle of 62° in less than 15 ms. The power requirements for jaw depression are calculated from a mathematical model and may reach a peak of ±4 watt in a 12-cm-long specimen. Data from the literature on mechanical output of fish muscle suggest that a muscular volume equal to 44% of the volume of the fish (= 45 cm3 ) should be required, if it is premised that movement and muscle shortening are directly coupled. Therefore, we assumed that storage and release of strain energy must be considered. The work demands for depression equal 0.014 J. The amount of energy storable in only three pairs of head ligaments can be estimated between 0.008 J and 0.05 J. The use of strain energy implies initial blocking and subsequent triggering of the movement. The position of the hyalomandibular connection, dorsal to the mandibular-suspensorial articulation, appears to be of crucial importance in balancing the forces of sternohyoideus and the body muscles. Triggering at the onset of jaw depression occurs by the contraction of the levator operculi. The line of action of the hyalomandibular connection is lowered beneath the jaw suspension, which raises the equilibrium of forces. Elastic recoil can occur thereafter.

Model of jaw depression during feeding in Astatotilapia elegans (Teleostei Cichlidae). Mechanisms for energy storage and triggering.

During ontogeny, larval fish have to deal with increasing nutritional and respiratory demands as they grow. As early ontogeny is characterized by an increasing complexity of moving structural elements composing a fish skull, some constraints will have to be met when developing mechanisms, which enable feeding and respiration, arise at a certain developmental stage. This article focuses on the presence/absence of a possible functional response in mouth opening during ontogeny in Clarias gariepinus. Some reflections are given, based on morphological data, as well as related function-analysis data from the literature. Starting shortly after hatching, a total of up to five different mouth opening mechanisms may become functional. Of these, three may remain functional in the adult. As could be expected, the apparatuses that enable these mechanisms show an increase in complexity, as well as a putative improvement in mouth opening capacity. Initially, two consecutive mechanisms may allow a restricted depression of the lower jaw (both passively and actively). Synchronously, two more mechanisms may arise, which involve the coupling of the hyoid depression to the mouth opening. At about 11 mm SL a fifth mechanism becomes established, better known as the opercular mouth opening mechanism. An overlapping chronology of functionality of the different mechanisms, as well as differences in efficiencies, could be an indication of the absence of a true critical period in C. gariepinus (at least in relation to mouth opening), as well as the possible presence of a shift in feeding type. Finally, the coupling of the chronology of the shift in mouth opening mechanisms and several morphological, behavioral, and physiological changes during ontogeny, related to feeding and respiration, make it possible to distinguish five important phases in the early life history of C. gariepinus.

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Ontogenetic shift in mouth opening mechanisms in a catfish (Clariidae, Siluriformes): A response to increasing functional demands

Within clariid fishes several cranial morphologies can be discerned. Especially within anguilliform representatives an increase in the degree of hypertrophy of the jaw adductors occurs. The hypertrophy of the jaw adductors and skeletal modifications in the cranial elements have been linked to increased bite force. The functional significance of this supposed increase in bite force remains obscure. In this study, biomechanical modeling of the cranial apparatus in four clariid representatives showing a gradual increase in the hypertrophy of the jaw adductors (Clarias gariepinus, Clariallabes melas, Channallabes apus, and Gymnallabes typus) is used to investigate whether bite force actually increased. Static bite modeling shows that the apparent hypertrophy results in an increase in bite force. For a given head size, the largest bite forces are predicted for C. apus, the lowest ones for C. gariepinus, and intermediate values are calculated for the other species. In addition, also in absolute measures differences in bite force remain, with C. apus biting distinctly harder than C. gariepinus despite its smaller head size. This indicates that the hypertrophy of the jaw adductors is more than just a correlated response to the decrease in absolute head size. Further studies investigating the ecological relevance of this performance difference are needed.

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Bite performance in clariid fishes with hypertrophied jaw adductors as deduced by bite modeling.

We compare the cranial morphology of four fish species with an increasing anguilliformism in the following order: Clarias gariepinus, Clariallabes melas, Gymnallabes typus, and Channallabes apus. The main anatomical-morphological disparities are the stepwise reduction of the skull roof along with the relative enlargement of the external jaw muscles, which occurred in each of them. Gymnallabes typus and C. apus lack a bony protection to cover the jaw muscles. The neurocranial bones of C. gariepinus, however, form a closed, broad roof, whereas the width of the neurocranium in C. melas is intermediate. Several features of the clariid heads, such as the size of the mouth and the bands of small teeth, may be regarded as adaptations for manipulating large food particles, which are even more pronounced in anguilliform clariids. The jaw musculature of G. typus is hypertrophied and attached on a higher coronoid process of the lower jaw, causing a larger adductive force. The hyomandibula interdigitates more strongly with the neurocranium and its dentition with longer teeth is posteriorly extended, closer to the lower jaw articulation. The anguilliform clariids also have their cranial muscles modified to enable a wider gape. The adductor mandibulae and the levator operculi extend more posteriorly, and the anterior attachment site of the protractor hyoidei dorsalis shifts toward the sagittal plane of the head. A phylogenetic analysis of the Clariidae, which is in progress, could check the validity of Boulenger's hypothesis that predecessors of the primitive fishes, such as Heterobranchus and most Clarias, would have evolved into progressively anguilliform clariids. J. Morphol. 240:169-194, 1999. © 1999 Wiley-Liss, Inc.

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Comparative study on the cranial morphology of Gymnallabes typus (Siluriformes: Clariidae) and their less anguilliform relatives, Clariallabes melas and Clarias gariepinus.

Walter Verraes

Papers

Ontogeny of the osteocranium in the African catfish, Clarias gariepinus burchell (1822) (Siluriformes: Clariidae): Ossification sequence as a response to functional demands

Model of jaw depression during feeding in Astatotilapia elegans (Teleostei Cichlidae). Mechanisms for energy storage and triggering.

Ontogenetic shift in mouth opening mechanisms in a catfish (Clariidae, Siluriformes): A response to increasing functional demands

Bite performance in clariid fishes with hypertrophied jaw adductors as deduced by bite modeling.

Comparative study on the cranial morphology of Gymnallabes typus (Siluriformes: Clariidae) and their less anguilliform relatives, Clariallabes melas and Clarias gariepinus.