Molecular phylogenetics and evolutionary diversification of labyrinth fishes (Perciformes: Anabantoidei)

Labyrinth fishes (Perciformes: Anabantoidei) are primary freshwater fishes with a disjunct African-Asian distribution that exhibit a wide variety of morphological and behavioral traits. These intrinsic features make them particularly well suited for studying patterns and processes of evolutionary diversification. We reconstructed the first molecular-based phylogenetic hypothesis of anabantoid intrarelationships using both mitochondrial and nuclear nucleotide sequence data to address anabantoid evolution. The mitochondrial data set included the complete cytochrome b, partial 12S rRNA, complete tRNA Val, and partial 16S rRNA genes (3332 bp) of 57 species representing all 19 anabantoid genera. The nuclear data set included the partial RAG1 gene (1494 bp) of 21 representative species. The phylogenetic analyses of a combined (mitochondrial+nuclear) data set recovered almost fully resolved trees at the intrafamily level with different methods of phylogenetic inference. Phylogenetic relationships at this taxonomic level were compared with previous morphology-based hypotheses. In particular, the enigmatic pike-head (Luciocephalus) was confidently placed within the "spiral egg" clade, thus resolving the long-standing controversy on its relative phylogenetic position. The molecular phylogeny was used to study the evolution of the different forms of parental care within the suborder. Our results suggest that the evolution of breeding behavior in anabantoids is highly correlated with phylogeny, and that brood care evolved three times independently from an ancestral free spawning condition without parental care. Ancestral character state reconstructions under maximum parsimony and maximum likelihood further indicated that both bubble nesting and mouthbrooding have evolved recurrently during anabantoid evolution. The new phylogenetic framework was also used to test alternative biogeographic hypotheses that account for the disjunct African-Asian distribution. Molecular divergence time estimates support either a drift vicariance linked to the breakup of Gondwana or Late Mesozoic Early Tertiary dispersal from Africa to Asia or vice versa.     

Some characters of Trichonotidae,with emphasis to those distinguishing it from Creediidae (Perciformes: Trachinoidei)

An osteological study of two species of the family Trichonotidae reveals numerous differences with members of the family Creediidae. Distinguishing characters include the following: trichonotids possess a flat pelvis base (not bowl-shaped) with oblique lateral flanges, a rodlike ectopterygoid that lacks a fanlike base, a dentary shelf that is notched anteriorly, a welldeveloped postmaxillary process, six infraorbitals, pleural ribs that arise from larger epipleural ribs, a hypurapophysis, and a large predorsal bone. Available evidence supports the view that these two families are monophyletic taxa with a marked gap in morphological characters between them. However, similarities in such bones as the ectopterygoid and mesopterygoid suggests that these two families are relatively closely related.     

A New Basal Lystrosaurid Dicynodont from the Upper Permian of South Africa

A new genus and species of late Permian dicynodont, Kwazulusaurus shakai , is described on the basis of a complete skull from the late Permian Dicynodon Assemblage Zone of the South African Beaufort Group. It is an advanced form which shows characters, such as the shape of the snout and the loss of the ectopterygoid, that link it to the early Triassic genus Lystrosaurus. Kwazulusaurus represents the most basal member of the lystrosaurian lineage. The phylogeny of progressive pristerodontian dicynodonts is discussed. It appears possible that the Kannemeyeriiformes and Lystrosauridae do not form a monophylum, as previously assumed. Instead a sister-group relationship between lystrosaurids and dicynodontids plus kannemeyeriiforms is suggested.     

Development and diversity of the suspensorium of trichomycterids and comparison with loricarioids (Teleostei: Siluriformes)

Studies of ontogenetic series of trichomycterids and other catfishes reveal that the suspensorium of siluroids is highly specialized; several synapomorphies separate siluroids from other teleosts. In siluroids, the palatoquadrate is divided into pars autopalatina and pars pterygoquadrata and both are usually connected by the autopaiatine-metapterygoid ligament. The pterygoquadrate is broadly joined to the dorsal limb of the hyoid arch, forming a cartilaginous hyomandibular-symplectic-pterygoquadrate plate in early ontogeny. This produces a special alignment of the hyomandibula and quadrate which is characteristic of siluroids. A symplectic bone is absent. The interhyal is absent in trichomycterids and astroblepids. Dorsal and ventral limbs of the hyoid arch are connected by a ligament. A rudimentary interhyal and this ligament are present in primitive siluroids such as diplomystids and nematogenyids as well as loricariids. The metapterygoid arises as an anterior ossification of the pars pterygoquadrata in siluroids. The formation and position of the metapterygoid exhibit two patterns: (1) the metapterygoid develops as an ossification of a cartilaginous projection positioned between the future hyomandibula and quadrate in primitive catfishes (e.g., Diplomystes) as well as in Nematogenys, callichthyids, loricariids, and astroblepids; (2) the metapterygoid arises as an ossification of the cartilaginous projection (pterygoid process) positioned just above the articular facet of the quadrate for the lower jaw. An ossified anterior chondral pterygoid process of the complex quadrate is present in trichomycterids, whereas the process is absent (simple quadrate) in catfishes such as diplomystids, nematogenyids, callichthyids, and loricariids. The anterior membranous process of the quadrate of Astroblepus is non-homologous with the chondral pterygoid process of trichomycterids; both structures arose independently within the loricarioids. Despite topological relationships, the origin and development of bones reveal the presence of a chondral hyomandibula which develops a large meinbranous outgrowth during ontogeny and a chondral metapterygoid in trichomycterids. The presence of a compound hyomandibula + metapterygoid or a compound metapterygoid + ectopterygoid + entopterygoid have no developmental support in trichomycterines or other siluroids. The “entopterygoid” of Nematogenys and Diplomystes arises as an ossification of a ligament. The dermal entopterygoid of other ostariophysans and the “entopterygoid” are homologous. An ectopterygoid or tendon bone “ectopterygoid” is absent in loricarioids. The suspensorium is an important structural system which has significant evolutionary transformations which characterize loricarioid subgroups; however, no character, of the suspensorium supports the monophyly of the loricarioids.     

Morphometries of the ectopterygoid in advanced snakes (Colubroidea): a concordance of shape and phylogeny

A principal components analysis was performed on 20 measurements of the ectopterygoids of 85 colubroid snakes. This sample encompassed all the shape variation previously recognized in the ectopterygoid of colubroids. Simple proportions which correlate with the first two principal component axes are: relative ectopterygoid length with the first and five measures of absolute or relative head shape with the second. Using these simple proportions colubroid ectopterygoids can be sorted into four shape classes each concordant with a taxonomic grouping: relatively long with broad notched head—Crotalinac; relatively long with narrow, flat head—Viperinae; relatively short with broad, notched head—Colubridae; and relatively short with narrow, flat head—Elapidae. This concordance has an error of about 10⁰₀. We propose that these shape classes reflect phylogenetically old and conservative functional differences in the palatomaxillary complexes of the four taxonomic groupings. Other aspects of ectopterygoid shape are described and provisional character state phylogenies for some aspects of the ectopterygoid are presented. Finally, the bearing of our data on the systematics of aparallactines, micrurines, sea snakes, Azemiops and certain other genera are discussed.     

A 3-D geometric morphometric study of intraspecific variation in the ontogeny of the temporal bone in modern Homo sapiens

This study addresses how the human temporal bone develops the population-specific pattern of morphology observed among adults and at what point in ontogeny those patterns arise. Three-dimensional temporal bone shape was captured using 15 landmarks on ontogenetic series of specimens from seven modern human populations. Discriminant function analysis revealed that population-specific temporal bone morphology is evident early in ontogeny, with significant shape differences among many human populations apparent prior to the eruption of the first molar. As early as five years of age, temporal bone shape reflects population history and can be used to reliably sort populations, although those in closer geographic proximity and molecular affinity are more likely to be misclassified. The deviation of cold-adapted populations from this general pattern of congruence between temporal bone morphology and genetic distances, identified in previous work, was confirmed here in adult and subadult specimens, and was revealed to occur earlier in ontogeny than previously recognized. Significant differences exist between the ontogenetic trajectories of some pairs of populations, but not among others, and the angles of these trajectories do not reflect genetic relationships or final adult temporal bone size. Significant intrapopulation differences are evident early in ontogeny, with differences becoming amplified by divergent trajectories in some groups. These findings elucidate how the congruence between adult human temporal bone morphology and population history develops, and reveal that this pattern corresponds closely to that described previously for facial ontogeny.     

Morphological and biochemical variations in the gills of 12 aquatic air-breathing anabantoid fish

All fish species in the Anabantoidei suborder are aquatic air-breathing fish. These species have an accessory air-breathing organ, called the labyrinth organ, in the branchial cavity and can engulf air at the surface of the water to assist in gas exchange. It is therefore necessary to examine the extent of gill modification among anabantoid fish species and the potential trade-offs in their function. The experimental hypothesis that we aimed to test is whether anabantoid fishes have both morphological and functional variations in the gills among different species. We examined the gills of 12 species from three families and nine genera of Anabantoidei. Though the sizes of the fourth gill arch in three species of Trichogaster were reduced significantly, not all anabantoid species had morphological and functional variations in the gills. In these three species, the specific enzyme activity and relative protein abundance of Na(+)/K(+)-ATPase were significantly higher in the anterior gills as compared with the posterior gills and the labyrinth organ. The relative abundance of cytosolic carbonic anhydrase, an indicator of gas exchange, was found to be highest in the labyrinth organ. The phylogenetic distribution of the fourth gill's morphological differentiation suggests that these variations are lineage specific, which may imply a phylogenetic influence on gill morphology in anabantoid species.     

Phylogenetic interpretation of ontogenetic change: sorting out the actual and artefactual in an empirical case study of centrarchid fishes

Hypothesized relationships between ontogenetic and phylogenetic change in morphological characters were empirically tested in centrarchid fishes by comparing observed patterns of character development with patterns of character evolution as inferred from a representative phylogenetic hypothesis. This phylogeny was based on 56–61 morphological characters that were polarized by outgroup comparison. Through these comparisons, evolutionary changes in character ontogeny were categorized in one of eight classes (terminal addition, terminal deletion, terminal substitution, non-terminal addition, non-terminal deletion, non-terminal substitution, ontogenetic reversal and substitution). The relative frequencies of each of these classes provided an empirical basis from which assumptions underlying hypothesized relationships between ontogeny and phylogeny were tested. In order to test hypothesized relationships between ontogeny and phylogeny that involve assumptions about the relative frequencies of terminal change (e.g. the use of ontogeny as a homology criterion), two additional phylogenies were generated in which terminal addition and terminal deletion were maximized and minimized for all characters. Character state change interpreted from these phylogenies thus represents the maxima and minima of the frequency range of terminal addition and terminal deletion for the 8.7 × 10³⁶ trees possible for centrarchids. It was found for these data that terminal change accounts for c. 75% of the character state change. This suggests either that early ontogeny is conserved in evolution or that interpretation and classification of evolutionary changes in ontogeny is biased in part by the way that characters are recognized, delimited and coded. It was found that ontogenetic interpretation is influenced by two levels of homology decision: an initial decision involving delimitation of the character (the ontogenetic sequence), and the subsequent recognition of homologous components of developmental sequences. Recognition of phylogenetic homology among individual components of developmental sequences is necessary for interpretation of evolutionary changes in ontogeny as either terminal or non-terminal. If development is the primary criterion applied in recognizing individual homologies among parts of ontogenetic sequences, the only possible interpretation of phylogenetic differences is that of terminal change. If homologies of the components cannot be ascertained, recognition of the homology of the developmental sequence as a whole will result in the interpretation of evolutionary differences as substitutions. Particularly when the objective of a study is to discover how ontogeny has evolved, criteria in addition to ontogeny must be used to recognize homology. Interpretation is also dependent upon delimitation within an ontogenetic sequence. This is in part a function of the way that an investigator ‘sees’ and codes characters. Binary and multistate characters influence interpretation differently and predictably. The use of ontogeny for determining phylogenetic polarity as previously proposed rests on the assumptions that ancestral ontogenies are conserved and that character evolution occurs predominantly through terminal addition. It was found for these data that terminal addition may comprise a maximum of 51.9% of the total character state change. It is concluded that the ontogenetic criterion is not a reliable indicator of phylogenetic polarity. Process and pattern data are collected simultaneously by those engaged in comparative morphological studies of development. The set of alternative explanatory processes is limited in the process of observing development. These form necessary starting points for the research of developmental biologists. Separating ‘empirical’ results from interpretational influences requires awareness of potential biases in the course of character selection, coding and interpretation. Consideration of the interpretational problems involved in identifying and classifying phylogenetic changes in ontogeny leads to a re-evaluation of the purpose, usefulness and information conveyed by the current classification system. It is recommended that alternative classification schemes be pursued.     

Phylogenetic aspects of tooth and jaw structure of the Medaka, Oryzias latipes, and other beloniform fishes

Tooth structure is described for adult male, female, and juvenile Oryzias latipes (Temminck & Schlegel), the Medaka. Adult males have enlarged, unicuspid teeth posteriorly on the premaxilla and dentary. Oral teeth are smaller and more numerous in females, in which no tooth is notably larger than the others. Juveniles have numerous small teeth from about 3 mm SL (standard length) onwards. By about 16 mm SL, males begin to develop the large posterior teeth, as well as other secondary sexual characters. Lower and upper pharyngeal teeth of both males and females are fine, and in numerous even rows.
The large, posterior oral teeth of males are fully-ankylosed to the attachment bone, and, hence, are not depressible. In female Medaka, as in the Halfbeak Dermogenys pusillus van Hasselt, the oral teeth have a ring of unmineralized collagen at the base, and are not depressible. Pharyngeal teeth of Medaka have a ring of unmineralized collagen at the base, and a distinct wedge of collagen absent posteriorly, such that the pharyngeal teeth may be depressed.
Bone in adult Medaka is acellular. Incompletely mineralized teeth, acellular bone, a protrus-ible upper oral jaw, and a mobile branchial apparatus with an interhyal bone, form a complex characteristic of advanced teleosts. The Medaka differs in several ways from the model advanced teleost: absence of an interhyal bone, ascending and articular processes of the premaxilla, and the rostral cartilage, as well as presence of cartilaginous symphyses between the dentaries and premaxillae, all contribute to the fixed or nonprotrusible jaws.
Reduction in the premaxilla is a derived character within beloniform fishes for which an enlarged, beaked outer jaw is considered plesiomorphic.     

Differences between inter- and intraspecific architectonic adaptations to pharyngeal mollusc crushing in cichlid fishes

Because fish heads are densely packed with muscles, ligaments, skeletal elements and other structures, transformations in one structure may influence surrounding structures. Transformations occur during phylogeny, ontogeny and as environmentally induced alterations, i.e. phenotypic plasticity. We describe differences in intra- and interspecific transformations of the pharyngeal jaw apparatus of haplochromine cichlids. Using multivariate clustering techniques we trace possible correlations in transformations of anatomical characters of the pharyngeal jaws and surrounding structures. The intraspecific transformation analysis is based on two environmentally induced morphs of Astatoreochromis alluaudi : a molluscivorous morph with a hypertrophied pharyngeal jaw apparatus and an insectivorous one with a non-hypertrophied apparatus. For the interspecific analysis five other haplochromine species from Lake Victoria with diets ranging from insects to molluscs were investigated. Although ranges in diet are the same, the anatomical ranges differ between A. alluaudi and the species cline. Besides similarities in anatomical changes of the pharyngeal jaw apparatus in the intra- and interspecific cline, differences were also observed. Apparently there are among haplochromines multiple pathways to achieve similar performance. In A. alluaudi architectonic and intrinsic plasticity constraints limit the adaptability of the pharyngeal jaw apparatus. In the species cline, these constraints have been overcome by genetical adaptation.     

Studies on the skull of the Indian sisorid catfish, Erethistes pussilus

The skull of Erethistes pussilus is described in detail. The general disposition of the bones corresponds to the siluroid pattern. Among the siluroid fishes, E. pussilus approaches the advanced forms in certain features such as obliteration of myodomic space, edentulous palate, absence of entopterygoids and supratemporals, intimate sutural articulation of posttemporals and complex vertebra with the cranium, diminished cranial cavity and loss of sutural articulation among the palatopterygoquadrate elements. In certain characters like the hyomandibula exclusively supported from the sphenotic, solitary hypohyal on each hyoid cornu, absence of interhyals, reduced orbits, edentulous vomer, small gape of mouth, toothless ectopterygoid and in the small number of branchiostegals, E. pussilus stands specialized alone among the catfishes. A diagnosis of the salient cranial characters of the fish is given and its relationship discussed.     

Studies on the skull of the Indian sisorid catfish, Erethistes pussilus

The skull of Erethistes pussilus is described in detail. The general disposition of the bones corresponds to the siluroid pattern. Among the siluroid fishes, E. pussilus approaches the advanced forms in certain features such as obliteration of myodomic space, edentulous palate, absence of entopterygoids and supratemporals, intimate sutural articulation of posttemporals and complex vertebra with the cranium, diminished cranial cavity and loss of sutural articulation among the palatopterygoquadrate elements. In certain characters like the hyomandibula exclusively supported from the sphenotic, solitary hypohyal on each hyoid cornu, absence of interhyals, reduced orbits, edentulous vomer, small gape of mouth, toothless ectopterygoid and in the small number of branchiostegals, E. pussilus stands specialized alone among the catfishes. A diagnosis of the salient cranial characters of the fish is given and its relationship discussed.     

Shape analysis of the jaws between two minnow species over ontogeny

This study compares sand shiner (Notropis stramineus ) and silverjaw (Ericymba buccata ) minnows, in terms of the morphological shape changes of the upper, lower, and pharyngeal jaws over ontogeny. These two species of minnows initially feed on midge larvae and undergo an ontogenic prey shift. The traditional morphometrics measured—total length, snout‐to‐vent length, eye diameter, premaxilla length, lower jaw length, gape—were regressed onto total length to test for allometry. Digital pictures were processed with tpsDig and further analyzed with MorphoJ utilizing a regular geometric morphometrics procedure using principle component analyses. We examined gut contents for 16 fish of each species. For the silverjaw minnows, we found all jaw variables to exhibit positive allometric growth with increasing total length, while most of the jaw variables for the sand shiner exhibited negative allometric growth with increasing total length. This correlates with an ontogenic prey shift for both species. Sand shiner minnows have been found to be more omnivorous, feeding on algae later in life, while silverjaw minnows undergo a prey shift to larger invertebrates. These species lack oral dentition causing an increased reliance on the pharyngeal apparatus. Principle component analyses revealed elongation of pharyngeal jaw elements in the silverjaw minnows and a relative shortening and bulking of pharyngeal jaws in the sand shiner minnows. The ontogenic dietary shifts observed in these two species provide possible explanation for the morphological changes over ontogeny in jaw elements, which are likely enabling these species to occupy the same habitat with little niche overlap.     

Morphological convergence of pharyngeal jaw structure in durophagous perciform fish

This study investigated the ecomorphology of pharyngeal jaw structure and durophagy in three families of marine teleosts: the Sciaenidae, Haemulidae and Carangidae. Regressions of the bone and muscle mass of pharyngeal jaws were generated to elucidate the differences associated with eating hard-bodied and soft-bodied prey; within-family comparisons revealed significant differences in masses of bones and muscles involved with processing the former. Generally, the durophagous species − Trachinotus carolinus (Carangidae), Pogonias cromis (Sciaenidae) and Anisotremus surinamensis (Haemulidae) − had heavier and stronger pharyngeal toothplates and larger protractor pectoralis muscles, with masses of these musculoskeletal elements ranging from five times to nearly an order of magnitude larger than those of their soft-prey feeding relatives. Pogonias cromis and T. carolinus demonstrate convergence in the ontogeny and morphological modification of the pharyngeal toothplates and protractor pectoralis muscles that enhance crushing ability. In the Haemulidae, moderate size increases in a few pharyngeal jaw elements (and larger overall body size in A. surinamensis ) are sufficient for durophagy. Morphospace analysis of six species from the three families illustrates the strong functional association between the biomechanical properties of prey and the relative sizes of biting and transport mechanisms.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80 , 147−165.     

REVIEW ON PEIPIAOSTEUS BASED ON NEW MATERIALS OF P. PANI

<正>Based on abundant new materials of Peipiaosteus pani, many features are newly described, they include: rostral bones, circumorbital series, the ectopterygoid process of the maxilla, quadratojugals, branchial toothplates, axial skeleton, fulcra and epiaxial fin-rays on the caudal fin. A few other features such as the hyomandibula and branchiostegal are revised. The ontogenetic and taphonomic characters of Peipiaosteus indicate that it probably assumed a habit of migration like its extant descendants. A review of the differences between P. pani and P. fengnengensis shows that all the differences except the dorsal and anal fin-ray numbers proposed by Bai (1983) are not true or remain uncertain. Through comparison, stichopterus is included in the revised family: Peipiaosteidae. Finally, a review of the phylogeny of Acipenseriformes results in a conclusion of Peipiaosteidae and Acipenseridae as sister groups.     

Alternative life histories of the genusLucania: 2. Early ontogeny ofL. goodei,the bluefin killifish

Synopsis This is the second of three papers devoted to the interpretation of morphological development in altricial and precocial species within the genus Lucania. The focus of this paper was the early life history of the bluefin killifish,Lucania goodei. Reproductively mature specimens were collected in the run below Newport Spring, north of St. Marks National Wildlife Refuge in northwest Florida. These specimens were transported to the laboratory, where they served as brood stock for specimens described in this study. Offspring were reared under controlled conditions and were described according to the theory of saltatory ontogeny, which gives a sampling design based on morphological, rather than chronological, progression. The morphological development of these offspring is described on the basis of detailed illustrations, photomicrographs, and measurements of mensural and meristic characters. This account of early ontogeny, in combination with a corresponding study on the early ontogeny of the rainwater killifish,L. parva (Crawford & Balon 1994a), establishes the empirical basis for an altricial-precocial life history model (Crawford & Balon 1994b).     

The evolution of pharyngeal bones and teeth in Gobioninae fishes (Teleostei: Cyprinidae) analyzed with phylogenetic comparative methods

The subfamily Gobioninae is a subgroup in the specious fish family Cyprinidae, which bears high diversity in morphological and ecological dimensions and has its most components distributed in East Asia. In this study, the pharyngeal bones and teeth of 39 species belonging to 19 genera of the Gobioninae were examined, with the phylogenetic comparative method (PCM) and correlation methods employed to analyze the character evolution. Three characters on pharyngeal bones (shape of the pharyngeal bones, extension for attachment of the pharyngo-cleithralis internus posterior (PCIP) muscle, and teeth-bearing area) and six characters of pharyngeal teeth (shape of the five teeth in the main row, number of rows of the teeth) were identified and compared. When the character states were mapped on a molecular phylogenetic tree, it was found that, to adapt to different masticatory operations, different Gobioninae species have various morphological types of pharyngeal bones and teeth: some have intermediate pharyngeal bones bearing multiple rows of diverse teeth (conical, coarsely compressed, and compressed), others have broad pharyngeal bones bearing a single row of molar teeth, and still others have narrow pharyngeal bones bearing a single row of extremely compressed teeth. Tests on the phylogenetic signal and evolutionary associations revealed that evolution of the examined characters was all phylogenetically constrained and correlated. Owing to the homoplasy in evolution, it was suggested that the conventional method of using pharyngeal bones and teeth for phylogenetic reconstruction of cyprinid fishes should not be encouraged.     

Ontogeny of the Middle-Ear Air-Sinus System in Alligator mississippiensis (Archosauria: Crocodylia)

Modern crocodylians, including Alligator mississippiensis, have a greatly elaborated system of pneumatic sinuses invading the cranium. These sinuses invade nearly all the bones of the chondrocranium and several bony elements of the splanchnocranium, but patterns of postnatal paratympanic sinus development are poorly understood and documented. Much of crocodylomorph—indeed archosaurian—evolution is characterized by the evolution of various paratympanic air sinuses, the homologies of which are poorly understood due in large part to the fact that individual sinuses tend to become confluent in adults, obscuring underlying patterns. This study seeks to explore the ontogeny of these sinuses primarily to clarify the anatomical relations of the individual sinuses before they become confluent and thus to provide the foundation for later studies testing hypotheses of homology across extant and extinct Archosauria. Ontogeny was assessed using computed tomography in a sample of 13 specimens covering an almost 19-fold increase in head size. The paratympanic sinus system comprises two major inflations of evaginated pharyngeal epithelium: the pharyngotympanic sinus, which communicates with the pharynx via the lateral (true) Eustachian tubes and forms the cavum tympanicum proprium, and the median pharyngeal sinus, which communicates with the pharynx via the median pharyngeal tube. Each of these primary inflations gives rise to a number of secondary inflations that further invade the bones of the skull. The primary sinuses and secondary diverticula are well developed in perinatal individuals of Alligator, but during ontogeny the number and relative volumes of the secondary diverticula are reduced. In addition to describing the morphological ontogeny of this sinus system, we provide some preliminary exploratory analyses of sinus function and allometry, rejecting the hypothesis that changes in the volume of the paratympanic sinuses are simply an allometric function of braincase volume, but instead support the hypothesis that these changes may be a function of the acoustic properties of the middle ear.     

Zoogeography and biodiversity of the freshwater fishes of Southeast Asia

The ichthyofauna of the freshwater system of Southeast Asia is extremely diverse. A recent estimate of about 1000 species is probably an understatement. More than 10 new species are being added to the list annually. The distribution pattern of the Southeast Asian freshwater fishes can be divided into five zoogeographic regions. The first one is the Salween basin in Burma, with fishes mainly of the Indian subcontinent origin such as Amblypharyngodon atkinsoni, Bangana almorae and Brachydanio jayarami. The second zoogeographic area is the Mekong, Chao Phraya and Mae Khlong drainages which harbour fishes typical of the mainland of Southeast Asia such as Acanthorhodeus deignani, Barbichthys nitidus and Cirrhinus siamensis. The Malay Peninsula is the third region whose species composition is heavily influenced by the Siamese (such as Homaloptera smithi, Tuberoschistura baenzigeri and Botia beauforti) and Indonesian (such as Botia hymenophysa, Luciocephalus pulcher and Parosphromenus deissneri) elements. The islands of Sumatra, Borneo and Java are the fourth zoogeographic area of fish distribution. These islands show a high degree of endemism, especially in fishes of the family Belontiidae. Finally, the freshwater system of the Philippines is the last zoogeographic region of Southeast Asia. The area is characterized by the presence of closely related species of the cyprinids especially in Lake Lanao. Currently, high diversity of these freshwater fishes is being threatened by land development, such as deforestation, road construction and land expansion for plantation. Recent studies in the Gombak River basin show the extermination of 41 per cent of native fish species from 1969 to 1990. This is probably due to the construction of highways, logging, as well as land clearing for agriculture.