The function of the suckers of larval net-winged midges (Diptera: Blephariceridae)

1. Larval net-winged midges (Diptera: Blephariceridae) possess six ventral suckers that enable them to inhabit swift streams. Each sucker consists of a suction disc and a cavity with a piston. Large muscles are inserted within the piston, as well as at the base of the suction disc. This structure infers that both attachment and release of the sucker is achieved by vertical movements of the piston.
2. Live observations of blepharicerid larvae revealed that the sucker is indeed attached by an upward movement of the piston, but that the cavity is flooded when the sucker is released. The piston is lowered only at the end of a sucker 'step', expelling water from the cavity.
3. During foraging, the maxilla and the piston of the first sucker are moved synchronously, indicating that the first sucker functions as a holdfast thus facilitating grazing.
4. The adhesive forces, as well as the relative size of blepharicerid suckers, differ amongst species. They are highest in Hapalothrix lugubris and lowest in Liponeura cordata , which correlates with the hydraulic stress to which the larvae of these species are exposed in their preferred habitat. The balance between the efficiency of their retention structure and the hydraulic conditions of their preferred habitat defines a key dimension of their ecological niche.     

Evolutionary optimization of an anatomical suction cup: Lip collagen content and its correlation with flow and substrate in Neotropical suckermouth catfishes (Loricarioidei)

In riverine ecosystems, downstream drag caused by fast-flowing water poses a significant challenge to rheophilic organisms. In neotropical rivers, many members of a diverse radiation of suckermouth catfishes (Loricarioidei) resist drag in part by using modified lips that form an oral suction cup composed of thick flesh. Histological composition and morphology of this cup are interspecifically highly variable. Through an examination of 23 loricarioid species, we determined that the tissue most responsible for lip fleshiness is collagen. We hypothesized that lip collagen content is interspecifically correlated with substrate and flow so that fishes living on rocky substrates in high-flow environments have the largest, most collagenous lips. By mapping the amount and distribution of lip collagen onto a phylogeny and conducting ANOVA tests, we found support for this hypothesis. Moreover, these traits evolved multiple times in correlation with substrate and flow, suggesting they are an effective means for improving suction-based attachment. We hypothesize that collagen functions to reinforce oral suction cups, reducing the likelihood of slipping, buckling, and failure under high-flow, high-drag conditions. Macroevolutionary patterns among loricarioid catfishes suggest that for maximum performance, biomimetic suction cups should vary in material density according to drag and substrate requirements.     

The oral sucker of Gyrinocheilus aymonieri (Teleostei: Cypriniformes)

The sucker was studied in young and mature fish by light microscopy, histochemistry, transmission and scanning electron microscopy, X-ray probe microanalysis, dissection, staining preparations of whole skeletons, and watching the animals in aquaria. The fleshy lips are supported by highly flexible, chondroid tissues, the structure and histochemistry of which differ substantially from those of cartilage. They allow the sucker to evert when the fish attaches to a stone or aquarium wall and are connected to the maxillae, premaxillae and dentaries. Lining the inside of the lips are two horny rasps, each with several regular rows of small hooks. The scraping blades of these hooks are keratinized and point towards the mouth. They increase the coefficient of friction for adhesion and enable the fish to feed on encrusting algae. Between the posterior rasp and the -anterior margin of the mandible are two invaginations of the lower lip that extend the sucker chamber beneath large hollows in the dentaries. The anterior margin itself contacts the outer surface of the maxillary oral valve when the mouth is closed, and isolates the sucker chamber from the rest of the buccal and pharyngeal cavities. Contrary to previous views, it is thought that a true vacuum is produced, and that attached fish spend long periods without taking water in through the mouth. The attachments of the principal jaw muscles are described and their role in sucker action discussed. There are similarities with the jaw mechanism of catostomids.     

Linking cranial kinematics,buccal pressure,and suction feeding performance in largemouth bass

The rate and magnitude of buccal expansion are thought to determine the pattern of water flow and the change in buccal pressure during suction feeding. Feeding events that generate higher flow rates should induce stronger suction pressure and allow predators to draw prey from further away. We tested these expectations by measuring the effects of prey capture kinematics on suction pressure and the effects of the latter on the distance from which prey were drawn-termed suction distance. We simultaneously, but not synchronously, recorded 500-Hz video and buccal pressure from 199 sequences of four largemouth bass, Micropterus salmoides, feeding on goldfish. From the video, we quantified several kinematic variables associated with the head and jaws of the feeding bass that were hypothesized to affect pressure. In a multiple regression, kinematic data accounted for 79.7% of the variation among strikes in minimum pressure. Faster mouth opening and hyoid depression were correlated with lower pressures, a larger area under the pressure curve, and a faster rate of pressure reduction. In contrast, buccal pressure variables explained only 16.5% of the variation in suction distance, and no single pressure variable had a significant relationship with suction distance. Thus, although expected relationships between head kinematics and buccal pressure were confirmed, suction distance was only weakly related to buccal pressure. Three explanations are considered. First, bass may not attempt to maximize the distance from which prey are drawn. Second, the response of prey items to suction-induced flow depends on prey behavior and orientation and is, therefore, subject to considerable variation. Third, previous theoretical work indicates that water velocity decays exponentially with distance from the predator's mouth, indicating that variation among strikes in flow at the mouth opening is compressed away from the mouth. These findings are consistent with other recent data and suggest that suction distance is a poor metric of suction feeding performance.     

Functional morphology of the Andean climbing catfishes (Astroblepidae,Siluriformes): Alternative ways of respiration,adhesion, and locomotion using the mouth

Astroblepidae or “climbing catfishes” encompass a single genus of species living in high altitude rivers in the Andes of South America. They are characterized by a specialized head morphology closely resembling their better known, widely radiated sister family Loricariidae, or armored suckermouth catfishes. Existent data show that even though both families share important traits, there are some striking differences as well. Albeit poorly known, Astroblepus species possess a duplicated gill opening, and have the ability to climb vertical rocks or waterfalls. In this study, morphological and kinematic data are combined to yield insights into the functions of the mobile elements of the astroblepid head, and to compare head morphology and biomechanics with those of Loricariidae. We found that, even though there is substantial similarity in head structure of both families, there are major differences in functionally important structures. These include a different lower lip muscle configuration, an alternative oral valve system, and an incurrent gill opening only found in astroblepids. Kinematic analyses confirm that the astroblepid suckermouth, freed from its inhalatory function, offers advantages for climbing in the high‐altitude environment, and is used alternately with the extremely mobile pelvic girdle, in a crawling, nonundulatory motion. J. Morphol. 274:1164–1179, 2013. © 2013 Wiley Periodicals, Inc.     

Functional morphology of the feeding mechanism in aquatic ambystomatid salamanders

This study addresses four questions in vertebrate functional morphology through a study of aquatic prey capture in ambystomatid salamanders: (1) How does the feeding mechanism of aquatic salamanders function as a biomechanical system? (2) How similar are the biomechanics of suction feeding in aquatic salamanders and ray-finned fishes? (3) What quantitative relationship does information extracted from electromyograms of striated muscles bear to kinematic patterns and animal performance? and (4) What are the major structural and functional patterns in the evolution of the lower vertebrate skull? During prey capture, larval ambystomatid salamanders display a kinematic pattern similar to that of other lower vertebrates, with peak gape occurring prior to both peak hyoid depression and peak cranial elevation. The depressor mandibulae, rectus cervicis, epaxialis, hypaxialis, and branchiohyoideus muscles are all active for 40–60 msec during the strike and overlap considerably in activity. The two divisions of the adductor mandibulae are active in a continuous burst for 110–130 msec, and the intermandibularis posterior and coracomandibularis are active in a double burst pattern. The antagonistic depressor mandibulae and adductor mandibulae internus become active within 0.2 msec of each other, but the two muscles show very different spike and amplitude patterns during their respective activity periods. Coefficients of variation for kinematic and most electromyographic recordings reach a minimum within a 10 msec time period, just after the mouth starts to open. Pressure within the buccal cavity during the strike reaches a minimum of ?25 mmHg, and minimum pressure occurs synchronously with maximum gill bar adduction. The gill bars (bearing gill rakers that interlock with rakers of adjacent arches) clearly function as a resistance within the oral cavity and restrict posterior water influx during mouth opening, creating a unidirectional flow during feeding. Durations of electromyographic activity alone are poor predictors of kinematic patterns. Analyses of spike amplitude explain an additional fraction of the variance in jaw kinematics, whereas the product of spike number and amplitude is the best statistical predictor of kinematic response variables. Larval ambystomatid salamanders retain the two primitive biomechanical systems for opening and closing the mouth present in nontetrapod vertebrates: elevation of the head by the epaxialis and depression of the mandible by the hyoid apparatus.     

Functional morphology of the feeding apparatus,feeding constraints,and suction performance in the nurse shark Ginglymostoma cirratum

The nurse shark, Ginglymostoma cirratum, is an obligate suction feeder that preys on benthic invertebrates and fish. Its cranial morphology exhibits a suite of structural and functional modifications that facilitate this mode of prey capture. During suction‐feeding, subambient pressure is generated by the ventral expansion of the hyoid apparatus and the floor of its buccopharyngeal cavity. As in suction‐feeding bony fishes, the nurse shark exhibits expansive, compressive, and recovery kinematic phases that produce posterior‐directed water flow through the buccopharyngeal cavity. However, there is generally neither a preparatory phase nor cranial elevation. Suction is generated by the rapid depression of the buccopharyngeal floor by the coracoarcualis, coracohyoideus, and coracobranchiales muscles. Because the hyoid arch of G. cirratum is loosely connected to the mandible, contraction of the rectus cervicis muscle group can greatly depress the floor of the buccopharyngeal cavity below the depressed mandible, resulting in large volumetric expansion. Suction pressures in the nurse shark vary greatly, but include the greatest subambient pressures reported for an aquatic‐feeding vertebrate. Maximum suction pressure does not appear to be related to shark size, but is correlated with the rate of buccopharyngeal expansion. As in suction‐feeding bony fishes, suction in the nurse shark is only effective within approximately 3 cm in front of the mouth. The foraging behavior of this shark is most likely constrained to ambushing or stalking due to the exponential decay of effective suction in front of the mouth. Prey capture may be facilitated by foraging within reef confines and close to the substrate, which can enhance the effective suction distance, or by foraging at night when it can more closely approach prey. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Using the whole body as a sucker: Combining respiration and feeding with an attached lifestyle in hill stream loaches (Balitoridae,Cypriniformes)

Small fishes living in fast‐flowing rivers face a harsh environment as they can easily be swept away by the rapid currents. To survive such circumstances, teleosts evolved a wide variety of attachment mechanisms, based on friction, negative pressure or both. Balitorinae (Balitoridae, Cypriniformes) are exceptional in using their whole body as an adhesive apparatus. We investigated the morphological adaptations of Balitorinae by studying the osteology and myology of four species (Beaufortia leveretti, Sewellia lineolata, Pseudogastromyzon myersi, and Gastromyzon punctulatus) using clearing and staining, serial cross‐sections and CT‐scanning. A kinematic analysis was performed to study the respiration and feeding mechanisms and to identify key structures in these mechanisms. Our research showed that the whole body of Balitorinae acts as a suction disc, with friction‐enhancing structures (unculi) on the thickened anterior rays of the paired fins. The abruptly rising head profile, supported by the extremely enlarged lacrimal bone and the flat ventral body surface facilitate effective substrate attachment. During attachment, the pelvic girdle is pulled anterodorsally, suggesting the formation of a negative pressure underneath the body. Detachment by water inflow underneath the body is prevented by three mechanisms. 1) Barbels control the water inflow by detachment and reattachment to the substrate. 2) Most water present underneath the body is removed during inspiration. 3) Excess water is regularly removed by movements of the posterior pectoral fin rays. The balitorine body is thus modified as such that it allows effective attachment, while not impairing respiration. Comparison with other teleosts living in similar environments shows that most species use more locally concentrated modifications of the paired fins and/or the mouth for attachment. The high diversity in teleostean adhesive apparatuses and associated myological modifications suggest a substantial functional convergent evolution, without necessarily highly convergent anatomical adaptations. J. Morphol. 275:1066–1079, 2014. © 2014 Wiley Periodicals, Inc.     

On the mechanism of respiration in the bullfrog,Rana catesbeiana: A reassessment

The mechanism of respiration in the bullfrog has been analyzed by means of pressure recordings from the buccal cavity, the lungs and the abdominal cavity, by cinematography and cinefluorography, and by electromyography of buccal, laryngeal and abdominal muscles. Gas flow was investigated by putting frogs in atmospheres of changing argon and nitrogen content and monitoring the concentration of the nostril efflux. Three kinds of cyclical phenomena were found. (1) Oscillatory cycles consist of rhythmical raising and lowering of the floor of the mouth, with open nares. They have a definite respiratory function in introducing fresh air into the buccal cavity. (2) Ventilatory cycles involve opening and closing of the glottis and nares and renewal of a portion of the pulmonary gas. More muscles are involved and the pattern of muscular activity is more complex than in the oscillatory cycles. (3) Inflation cycles consist of a series of ventilation cycles, interrupted by an apneic pause. The intensity of the ventilatory cycles increases before this pause and decreases immediately thereafter. This results in a stepwise increase in pulmonary pressure, to a plateau (coincident with the pause) followed by a sudden or stepwise decrease. The respiratory mechanism depends on the activity of a buccal force pump, which determines pulmonary pressure whose level is always slightly less than the peak pressure values of the ventilation cycles. The elevated pulmonary pressure is responsible for the expulsion of pulmonary gas during the second phase of the next ventilation cycle. This pressure is maintained by the elastic fibers (and the smooth masculature) of the lungs.     

Onset of Buccal Pumping in Catshark Embryos: How Breathing Develops in the Egg Capsule

Respiration in fishes involves buccal pumping, which is characterized by the generation of nearly continuous water flow over the gills because of the rhythmic expansion/compression of the pharyngeal cavity. This mechanism is achieved by the functions of the vascular, skeletal, and muscular systems. However, the process by which the embryo establishes the mechanism remains a mystery. Morphological and kinematical observations on captive cloudy catsharks, Scyliorhinus torazame, have suggested that the embryo starts buccal pumping just before the respiratory slits open on the egg capsule. During the pre-opening period, the embryo acquires oxygen mainly via the external gill filaments. After slit opening, respiration of the embryo involves buccal pumping to pass water over the “internal gills.” The onset of buccal pumping accompanies four morphological changes: (1) regression of the external gill filaments, (2) development of blood vessels within the “internal gills,” (3) completion of the development of hyoid skeletal and muscular elements, and (4) development of the oral valve. A previous study showed that buccal pumping allows the embryo to actively regulate oxygen intake by changing the pumping frequency. Thus, establishment of buccal pumping in the egg capsule is probably important for embryo survival in the unstable oxygen environment of the egg capsule after slit opening.     

Application of the micronucleus test to exfoliated epithelial cells from the oral cavity of beedi smokers, a high-risk group for oral cancer

The primary sites for occurrence of oral cancer include the buccal mucosa, tongue, alveolus, palate, lip and the floor of the mouth. In this study, an attempt was made to estimate the cytogenetic damage in different regions of the oral mucosa in people habituated to smoking beedi,which is one of the major forms of tobacco consumption in India and believed to be a major risk factor for oral cancer. By using the micronucleus assay on exfoliated cells from the buccal mucosa, palate and tongue of beedi smokers, we examined an early cellular response to the effect of beedi smoking. A total number of 50 randomly selected male subjects were included in the study. Case and control groups (smokers and non-smokers, respectively) comprised 25 subjects each. The difference in mean micronucleated cell count between cases and controls was significant (P <0.01) for buccal mucosa and palate, but not for tongue. The correlation between age and micronucleus cell count was weak for both cases (r=0.27) and controls (r=0.36).     

Morphological and ultrastructural aspects of Branchiobdella pentodonta Whit. (Annelida,oligochaeta) suckers

The muscular system in the posterior sucker of Branchiobdella pentodonta Whit. has circular, longitudinal and radial fibers. In the anterior sucker, which has circular and longitudinal fibers, the muscle system is scarce. Concentric fibers are found around the mouth. In both suckers the glandular elements form voluminous complexes secreting mucus for attachment to the substrate. Suckers show neuromuscular junctions and three distinct types of neuroglandular junctions: one with typical neurosecretory granules, one with larger neurosecretory granules produced by cells located at the origin of the segmental nerves, and one with presynaptic vesicles. The second type is peculiar to the posterior sucker. A comparison is made between suckers of Branchiobdella and those of leeches.     

Effects of texture on surface attachment of spawning‐run sea lampreys Petromyzon marinus: a quantitative analysis

Adult sea lamprey Petromyzon marinus attachment strength by the oral sucker was quantified. Surfaces with shallow, rounded discontinuities into which the oral fimbriae could be folded yielded the strongest seal and some fish could control their mouths to improve ‘suction’. Narrow grooves of 1 mm width and 3 mm depth prevented P. marinus from creating a lasting attachment.     

The mechanism of breathing in the South Asmerican lungfish, Lepidosiren paradoxa; radiological study

In filling the lungs by means of the buccal "force-pump", air drawn into the buccal cavity by the lowering of the floor of the mouth is forced into the lungs when the mouth is firmly closed and the buccal floor raised. This mechanism depends on the ability of the fish to prevent air being forced out of the mouth through the lips and it is shown that the tongue is pressed against the roof of the mouth to form a seal. Radiologically it has been shown that the hyoid apparatus ("ceratohyal" of some authors) plays a large part in the movements necessary to draw air into the mouth, seal it in and force it into the lungs. Another skeletal element involved is the pectoral girdle which makes considerable movements in association with the filling of the lungs. Expiration of the air from the lungs, which as shown by radiographs may be remarkably complete, is brought about by the elasticity of the lung tissue.     

Aquatic prey capture in ray-finned fishes: a century of progress and new directions

The head of ray-finned fishes is structurally complex and is composed of numerous bony, muscular, and ligamentous elements capable of intricate movement. Nearly two centuries of research have been devoted to understanding the function of this cranial musculoskeletal system during prey capture in the dense and viscous aquatic medium. Most fishes generate some amount of inertial suction to capture prey in water. In this overview we trace the history of functional morphological analyses of suction feeding in ray-finned fishes, with a particular focus on the mechanisms by which suction is generated, and present new data using a novel flow imaging technique that enables quantification of the water flow field into the mouth. We begin with a brief overview of studies of cranial anatomy and then summarize progress on understanding function as new information was brought to light by the application of various forms of technology, including high-speed cinematography and video, pressure, impedance, and bone strain measurement. We also provide data from a new technique, digital particle image velocimetry (DPIV) that allows us to quantify patterns of flow into the mouth. We believe that there are three general areas in which future progress needs to occur. First, quantitative three-dimensional studies of buccal and opercular cavity dimensions during prey capture are needed; sonomicrometry and endoscopy are techniques likely to yield these data. Second, a thorough quantitative analysis of the flow field into the mouth during prey capture is necessary to understand the effect of head movement on water in the vicinity of the prey; three-dimensional DPIV analyses will help to provide these data. Third, a more precise understanding of the fitness effects of structural and functional variables in the head coupled with rigorous statistical analyses will allow us to better understand the evolutionary consequences of intra- and interspecific variation in cranial morphology and function.     

A bridge between original and novel states: ontogeny and function of "suction discs" in the Branchiura (Crustacea)

The emergence of novel structures in the course of evolution faces an explanatory problem, leaving the gap from the ancestral structures difficult to bridge. This difficulty is caused by the lack of intermediate stages. Branchiurans are ectoparasitic crustaceans which use a pair of "suction discs" to attach to their host. These structures are modified first maxillae. During ontogeny, the first maxillae transform from a normal cephalic appendage to the specialized suction disc. However, supposedly ancestral branchiurans lack the suction discs in the adults and the first maxilla remains a normal appendage throughout. We describe the muscular arrangements in the developing first maxillae in Argulus coregoni. The suction discs originate as a fusion of the first and second podomeres. The sucker muscles of the suction discs are homologous to the muscles that insert in the second podomere at the early larval stages. The developmental process of the suction disc can be seen as a "recapitulation" of the evolutionary process. We thus show how the first maxilla can maintain not just the biological role but also a functional continuity during the evolution of the novel structure. From this example it is obvious that the intermediate stages of the emerging novelty, if present in the ontogeny, can help solve at least some of the enigmatic appearances of novel structures.     

Morphological and functional bases of durophagy in the queen triggerfish,Balistes vetula (Pisces,tetraodontiformes)

Tetraodontiform fishes are characterized by jaws specialized for powerful biting and a diet dominated by hard-shelled prey. Strong biting by the oral jaws is an unusual feature among teleosts. We present a functional morphological analysis of the feeding mechanism of a representative tetraodontiform, Balistes vetula. As is typical for the order, long, sharp, strong teeth are mounted on the short, robust jaw bones of B. vetula. The neurocranium and suspensorium are enlarged and strengthened to serve as sites of attachment for the greatly hypertrophied adductor mandibulae muscles. Electromyographic recordings made from 11 cranial muscles during feeding revealed four distinct behaviors in the feeding repertoire of B. vetula. Suction is used effectively to capture soft prey and is associated with a motor pattern similar to that reported for many other teleosts. However, when feeding on hard prey, B. vetula directly bit the prey, exhibiting a motor pattern very different from that of suction feeding. During buccal manipulation, repeated cycles of jaw opening and closing (biting) were coupled with rapid movement of the prey in and out of the mouth. Muscle activity during buccal manipulation was similar to that seen during bite-captures. A blowing behavior was periodically employed during prey handling, as prey were forcefully “spit out” from the mouth, either to reposition them or to separate unwanted material from flesh. The motor pattern used during blowing was distinct from similar behaviors described for other fishes, indicating that this behaviors may be unique to tetraodontiforms. Thus B. vetula combines primitive behaviors and motor patterns (suction feeding and buccal manipulation) with specialized morphology (strong teeth, robust jaws, and hypertrophied adductor muscles) and a novel behavior (blowing) to exploit armored prey such as sea urchins molluscs, and crabs. © 1993 Wiley-Liss, Inc.     

Morphology and function of the feeding apparatus of the lungfish, Lepidosiren paradoxa (Dipnoi)

The feeding mechanism of the South American lungfish, Lepidosiren paradoxa retains many primitive teleostome characteristics. In particular, the process of initial prey capture shares four salient functional features with other primitive vertebrates: 1) prey capture by suction feeding, 2) cranial elevation at the cranio-vertebral joint during the mouth opening phase of the strike, 3) the hyoid apparatus plays a major role in mediating expansion of the oral cavity and is one biomechanical pathway involved in depressing the mandible, and 4) peak hyoid excursion occurs after maximum gape is achieved. Lepidosiren also possesses four key morphological and functional specializations of the feeding mechanism: 1) tooth plates, 2) an enlarged cranial rib serving as a site for the origin of muscles depressing the hyoid apparatus, 3) a depressor mandibulae muscle, apparently not homologous to that of amphibians, and 4) a complex sequence of manipulation and chewing of prey in the oral cavity prior to swallowing. The depressor mandibulae is always active during mouth opening, in contrast to some previous suggestions. Chewing cycles include alternating adduction and transport phases. Between each adduction, food may be transported in or out of the buccal cavity to position it between the tooth plates. The depressor mandibulae muscle is active in a double-burst pattern during chewing, with the larger second burst serving to open the mouth during prey transport. Swallowing is characterized by prolonged activity in the hyoid constrictor musculature and the geniothoracicus. Lepidosiren uses hydraulic transport achieved by movements of the hyoid apparatus to position prey within the oral cavity. This function is analogous to that of the tongue in many tetrapods.     

Development and evolution of adult feeding structures in Caenogastropods: overcoming larval functional constraints

SUMMARY Comparative study of the developing foregut in three species of caenogastropods, including an herbivorous grazer ( Lacuna vincta ) and two carnivores ( Euspira [ Polinices ] lewisii and Nassarius mendicus ), suggests how the specialized adult foregut of a carnivorous neogastropod evolved within a life cycle having a planktotrophic larva. Postmetamorphic feeding structures (buccal cavity and radular sac) in all three species achieve advanced differentiation in the larval stage, permitting juvenile feeding at 3 days postmetamorphosis. Recent phylogenetic hypotheses for the Gastropoda predict that foregut developmental patterns in E. lewisii and N. mendicus are derived, relative to that of L. vincta. In hatching larvae of these three, the anlage of postmetamorphic feeding structures is a small patch of nonciliated cells embedded in the ventral wall of the larval foregut and the patch soon forms an outpocketing. During subsequent morphogenesis, Euspira lewisii and N. mendicus share a developmental novelty that involves semi-isolation of the developing, postmetamorphic buccal cavity and radular sac from the larval foregut and formation of a new, definitive mouth at metamorphosis. Nassarius mendicus , a neogastropod, embellishes this novelty by adding the entire anterior esophagus and valve of Leiblein ( de novo structures) to the semi-isolated buccal cavity. Therefore, a valve and long stretch of muscular anterior esophagus, which are necessary for feeding with a pleurembolic proboscis, are preformed in the larval stage of this neogastropod without interfering with larval feeding. The inferred evolutionary events leading to postmetamorphic feeding specialization in N. mendicus are invisible in adults; they require reconstruction from comparative developmental analysis.     

A functional morphospace for the skull of labrid fishes: patterns of diversity in a complex biomechanical system

The Labridae (including wrasses, the Odacidae and the Scaridae) is a species‐rich group of perciform fishes whose members are prominent inhabitants of warm‐temperate and tropical reefs worldwide. We analyse functionally relevant morphometrics for the feeding apparatus of 130 labrid species found on the Great Barrier Reef and use these data to explore the morphological and mechanical basis of trophic diversity found in this assemblage. Morphological measurements were made that characterize the functional and mechanical properties of the oral jaws that are used in prey capture and handling, the hyoid apparatus that is used in expanding the buccal cavity during suction feeding, and the pharyngeal jaw apparatus that is used in breaking through the defences of shelled prey, winnowing edible matter from sand and other debris, and pulverizing the algae, detritus and rock mixture eaten by scarids (parrotfishes). A Principal Components Analysis on the correlation matrix of a reduced set of ten variables revealed complete separation of scarids from wrasses on the basis of the former having a small mouth with limited jaw protrusion, high mechanical advantage in jaw closing, and a small sternohyoideus muscle and high kinematic transmission in the hyoid four‐bar linkage. Some scarids also exhibit a novel four‐bar linkage conformation in the oral jaw apparatus. Within wrasses a striking lack of strong associations was found among the mechanical elements of the feeding apparatus. These weak associations resulted in a highly diverse system in which functional properties occur in many different combinations and reflect variation in feeding ecology. Among putatively monophyletic groups of labrids, the cheilines showed the highest functional diversity and scarids were moderately diverse, in spite of their reputation for being trophically monomorphic and specialized. We hypothesize that the functional and ecological diversity of labrids is due in part to a history of decoupled evolution of major components of the feeding system (i.e. oral jaws, hyoid and pharyngeal jaw apparatus) as well as among the muscular and skeletal elements of each component. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 82 , 1–25.