Feeding performance of Lake Victoria rock cichlids: testing predictions from morphology

Cichlid fishes that seem specialized phenotypically to exploit certain resources often act as generalists in the field. It is attempted to resolve this paradox by measuring feeding performance. There are two ways of feeding, namely suction feeding and biting, that set conflicitng demands on the anatomy. Rock-dwelling cichlids of the genus Neochromis are specialized biters, feeding largely on filamentous algae and benthic organisms. Pundamilia nyererei is one of the most specialized suction feeders among rock cichlids, its diet being zooplankton and insect larvae. A morphological analysis showed that the three Neochromis species are better adapted anatomically to biting than P. nyererei , while P. nyererei is better adapted to suction feeding. Feeding performance was tested on algal substitute and Chaoborus larvae, representing filamentous algae and zooplankton, respectively. As expected the Neochromis species fed most efficiently on algal substitute, N. rufocaudalis and N. omnicaeruleus taking per bite 2.6 times as much as P. nyererei , and N. greenwoodi 1.7 times. Unexpectedly, the species examined were all well able to collect Chaoborus. P. nyererei collected only 1.2 times as many larvae per suction act as the Neochromis species. Analysis of the stomach contents showed that P. nyererei punctured, or lacerated, the larvae with the pharyngeal jaws, while the Neochromis species swallowed them mostly intact. Thus, feeding on Chaoborus , P. nyererei may be optimizing energy gain and not intake rate.     

Functional and Genetic Integration in the Skulls of Lake Malawi Cichlids

The level of integration present among organismal traits is thought to influence evolutionary potential, and this potential should be affected by the type or types of integration displayed (e.g., functional, developmental, or genetic). Morphological integration is generally high among functionally related traits, but whether this is predominantly determined by genetic architecture, or is instead a result of biomechanical remodeling during development remains poorly understood. We examine this question in Lake Malawi cichlid fishes by combining a finite-element analysis (FEA) of bite force transmission with quantitative genetic analyses of skull morphology in order to test the hypothesis that functionally coupled traits share a common genetic basis. FEA modeling indicates that the profile of the neurocranium affects its ability to resist forces transmitted from the jaws during biting, and suggests a novel role for skull shape in fish feeding mechanics. Quantitative trait loci mapping demonstrates that the functional integration between jaw and neurocranial shape has a genetic basis, and that this association is being driven by alleles inherited from the specialized biting species. Notably, the co-inheritance of these two functionally related traits in our F₂ matches patterns of covariation within and between Lake Malawi cichlid species. Across species, jaw and neurocranial shapes covary, but the trend appears strongest among biting species. Similarly, within populations of biting species, the dimensions of the jaw and neurocranium are tightly linked, whereas this correlation disappears within populations of omnivorous and suction feeding fish. These data suggest (1) that either pleiotropy, or physical linkage maintained by selection, underlies the phenotypic integration of these two functionally related traits, and (2) that this pattern of integration may have influenced the radiation of craniofacial morphology in Lake Malawi cichlids.     

Replicated divergence in cichlid radiations mirrors a major vertebrate innovation

Decoupling of the upper jaw bones—jaw kinesis—is a distinctive feature of the ray-finned fishes, but it is not clear how the innovation is related to the extraordinary diversity of feeding behaviours and feeding ecology in this group. We address this issue in a lineage of ray-finned fishes that is well known for its ecological and functional diversity—African rift lake cichlids. We sequenced ultraconserved elements to generate a phylogenomic tree of the Lake Tanganyika and Lake Malawi cichlid radiations. We filmed a diverse array of over 50 cichlid species capturing live prey and quantified the extent of jaw kinesis in the premaxillary and maxillary bones. Our combination of phylogenomic and kinematic data reveals a strong association between biting modes of feeding and reduced jaw kinesis, suggesting that the contrasting demands of biting and suction feeding have strongly influenced cranial evolution in both cichlid radiations.     

Motor Control Across Trophic Strategies: Muscle Activity of Biting and Suction Feeding Fishes

Many fishes use a powerful bite of the oral jaws to captureor tear their prey. This behavior has received less study fromfunctional morphologists and physiologists than suction feeding,and presents an opportunity to examine motor control of fishfeeding across alternative prey-capture strategies. We usedelectromyography to compare muscle activity patterns of thefeeding bite in five teleost fishes representing at least threelineages in which biting has been independently acquired: twoparrotfish (Cetoscarus bicolor and Scarus iseri), a wrasse (Cheilinuschlorourus), and two serrasalmines, a pacu (Piaractus brachypomus)and a piranha (Pygocentrus nattereri). Multivariate analysisindicated that muscle activity patterns differed significantlyamong species, although a four-way ANOVA designed to test fordifferences within a phylogenetic hierarchy revealed that thebiting motor pattern was largely similar for both narrow andbroad phylogenetic comparisons. A comparison of the motor patternsof biting and suction feeding species revealed that biters hadsignificantly shorter durations of the epaxialis and sternohyoideusand significantly longer relative onset times of the epaxialis,adductor mandibulae, and sternohyoideus. Character mapping oftiming variables suggested that short relative onset times areprimitive for suction feeders and that this characteristic isgenerally retained in more advanced species. Despite these differences,all species overlap extensively in multivariate EMG space. Ourresults demonstrate that change in the feeding motor patternhas accompanied morphological and behavioral change in transitionsfrom suction to biting, which suggests that the neuromotor systemhas not acted as a constraint on the evolution of the feedingsystem in fishes.     

The influence of oral anatomy on prey selection during the ontogeny of two percoid fishes,Lagodon rhomboides and Centropomus undecimalis

Synopsis Ontogenetic increases in mouth size and changes in dentition of percoid fishes may affect the size and species of prey selected, thus influencing the fundamental trophic niche. To examine the influence of oral anatomy on prey selectivity by pinfish, Lagodon rhomboides, and snook, Centropomus undecimalis, two co-occurring percoid fishes with contrasting mouth morphologies, the mouth size, dentition, stomach contents, and available prey during ontogeny were quantified. Based on the presence of prey fragments in stomach contents and direct behavioral observation, prey were categorized by the feeding mode used during capture (suction/ramfeeding or biting). Centropomus has a larger size-specific gape than Lagodon during all ontogenetic stages. Although both feeding modes were used by Lagodon during ontogeny, the amount of prey captured using suction/ram-feeding declined and the amount of prey captured by biting increased with standard length. This change in feeding mode was associated with a change in incisor shape and width: Lagodon < 39 mm SL possessed narrow, pointed incisors and strongly selected amphipods, which are captured using suction/ram-feeding; Lagodon> 40 mm SL possessed wide, flat-topped incisors and significantly increased their selectivity for polychaetes, which are captured by biting. Centropomus used ram-feeding to capture prey at all ontogenetic stages. Size-selective feeding by Centropomus was apparent but could not be due to gape-limitation alone, because average prey body depth was only 45% of gape and was not proportional to absolute mouth size increase during ontogeny. Dietary diversity was greatest during the transition from suction/ram-feeding to biting in Lagodon. Lagodon had a higher dietary diversity at all ontogenetic stages than Centropomus, due in part to Lagodon's use of multiple feeding modes.     

The feeding habits of a group of herbivorous rock-dwelling cichlid fishes (Cichlidae: Perciformes) from Lake Malawi,Africa

Synopsis The feeding; habits of a group of tropical herbivorous rock-dwelling cichlid fishes from Lake Malawi, Africa, are investigated using stomach content analyses. The various species fed selectively on the periphyton of the rocky shores. Blue-green alga of the genus Calothrix was the most common item ingested by the group. Diatoms (Chrysophyta) also were abundant food items. Discriminant analysis showed that dietary items were good variables to identify species. Interspecific dietary differences showed a continuum from those species feeding primarily on Calothrix to those feeding primarily on diatoms. Algal resources exhibit distinct patterns of spatial variation. Diet was correlated with foraging behavior and trophic morphology. Interspecific differences in diet could possibly facilitate ecological coexistence among various species. Such coexistence would contribute to the maintenance of the high diversity fish faunas characteristic of the Great Rift Lakes of Africa.     

Evolutionary patterns in tropical marine reef fish feeding

The majority of tropical reef fishes are acanthopterygians. Most of them are percomorphs and thus are likely monophyletic. In accordance to modern systematics, the primitive types among the latter are large-mouthed suction feeders. Species from advanced families often have biting oral jaws with a reduced number and complexly shaped teeth. Mouth sizes decrease from the primitive towards the advanced reef fishes when ranked according to increasing family numbers (Nelson 1984). To create a functional resource axis, Randall's (1967) and Hobson's (1974) data on tropical reef fish feeding were re-interpreted by ranking food items from mobile to sessile prey. The primitive paracantho pterygian and acanthopterygian reef fishes are large-mouthed, suction-feeding predators on mobile prey. Most of the advanced, small-mouthed species are browsers and grazers, but often feed on mobile prey too. Obligatory specialists (monophagous and unable to switch) seem to be relatively rare among modern reef fishes. The trends stated above indicate a wealth of parallel developments in many advanced families of reef fishes towards small, often biting oral jaws. This parallelism may be the result of comparable regimes of selection pressures in reefs and of the need for newly evolved species to establish themselves within the already existing guilds.     

Feeding biomechanics of five demersal Antarctic fishes

There is scant information available on the ecomorphology of Antarctic fishes, and especially on their feeding capabilities. We measured interspecific variation in mechanical advantage (MA), force-producing capability, and suction index for the jaws of the five dominant taxa of high-Antarctic fishes: the nototheniid Trematomus bernacchii; the zoarcids Pachycara brachycephalum, Lycodichthys dearborni, and Ophthalmolycus amberensis; and the liparid Paraliparis devriesi. Analysis of variance indicated significant differences in jaw metrics, and ordinations of morphological traits identified three loosely defined groups reflecting their family-level taxonomy. Principal component analyses showed distinct segregation between the nototheniid and the liparid, indicating that they are at the extremes of the feeding performance continuum. The zoarcids fell in the middle, suggesting that they utilize a combination of feeding modes to capture prey. The liparid had the lowest MA and bite force, but a large epaxialis implied a ram-suction-feeding mode. The large adductor mandibulae in the zoarcids P. brachycephalum and L. dearborni suggest that they are capable of grasping mobile prey and manipulating sedentary, hard-shelled macroinvertebrates. The zoarcids had a smaller epaxialis than the liparid and may not be as efficient as suction-feeders. Values for mechanical advantage ratios and suction indices in Antarctic fishes were within the range known for non-Antarctic fishes. The five Antarctic species do not possess dentition specialized for durophagous feeding; however, the high mechanical advantage ratio in the nototheniid and, to a lesser extent, in the zoarcids, suggests that durophagy may be possible.     

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.     

Genetic and morphological population differentiation in the rock-dwelling and specialized shrimp-feeding cichlid fish species <Emphasis Type="Italic">Altolamprologus compressiceps</Emphasis> from Lake Tanganyika,East Africa

With about 250 endemic species, Lake Tanganyika contains an extraordinarily diverse cichlid fish fauna, and thus represents an ideal model system for the study of pathways and processes of speciation. The Lamprologini form the most species-rich tribe in Lake Tanganyika comprising about 100 species in seven genera, most of which are endemic to the lake. They are territorial substrate-breeders and represent a monophyletic tribe. By combined analysis of population genetics and geometric morphometric markers, we assessed gene flow among three populations of the highly specialized shrimp-feeding rock-dweller Altolamprologus compressiceps, separated by geographic distance and ecological barriers. Five highly polymorphic microsatellite markers were analyzed in conjunction with 17 landmarks in order to compare genetic differences to body shape differences among populations. Both genetic and morphological analyses revealed significant differentiation among the three studied populations. A significant, but overall relatively low degree of genetic differentiation supports a very recent divergence. Phenotypic differentiation was primarily found in the head region of A. compressiceps. In agreement with findings in other cichlid species, similar adaptations to specialized feeding mechanisms can consequently lead to marginal shape changes in the trophic apparatus.     

Architectonic constraints on the hyoid's optimal starting position for suction feeding of fish

During suction feeding teleost fish have to start mouth opening prior to other expansion movements of the head such as operculo-suspensorium abduction. The distribution of the input force over the various expansion movements is determined by the position of the hyoid in the expansion apparatus. Based on a three-dimensional (3-D) kinematic model of this apparatus it can be calculated at which positions of the hyoid operculo-suspensorial abduction is precluded. For 73 cichlid species from various African lakes and covering a wide array of feeding types and adult sizes, it is demonstrated that these optimal positions for the onset of suction feeding can be attained or closely approached by species whose regular diet only requires suction feeding but not by species whose regular diet is dominated by items requiring forceful biting. It is argued that the suboptimality of the biters is due to an architectonic constraint, viz. an increase in head width necessary to accommodate their enlarged m. adductor mandibulae. Although it is theoretically feasible to optimize the model's parameters for every head width, the biters apparently have not achieved such an adaptive change. As these parameters also feature in the execution of other functions, it is likely that conflicting demands on their optimal value overrule their optimization for the starting position of the hyoid of biters. The results hold for cichlids of independently evolved species flocks and therefore concern general rules for biter-sucker transformations in cichlids. © 1996 Wiley-Liss, Inc.     

The cusp of evolution and development: a model of cichlid tooth shape diversity

Tooth shape is a hallmark of repeated evolutionary radiations among cichlid fishes from East Africa. Cusp shape and number vary both within populations and among closely related species with different feeding behaviors and ecologies. Here, we use histology and scanning electron microscopy to chart the developmental trajectory of tooth shape differences in fishes from Lake Malawi. We demonstrate that species with bi- or tricuspid adult (replacement) teeth initially possess a first-generation unicuspid dentition. Notably, the timing of turnover from first-generation to replacement teeth differs among species and is correlated with feeding ecology. Next, we use field data for cichlid species with adult unicuspid, bicuspid, and tricuspid teeth to demonstrate a strong and positive relationship between the number of teeth in a row and tooth shape. We discuss cichlid tooth ontogeny in the context of morphogenetic models designed to explain the developmental basis of tooth shape variation in mammals. We suggest that the dramatic differences in cichlid dentitions can be explained by variation in the expression of common activators and inhibitors acting at multiple stages of odontogenesis.     

Feeding biomechanics and theoretical calculations of bite force in bull sharks (Carcharhinus leucas) during ontogeny

Evaluations of bite force, either measured directly or calculated theoretically, have been used to investigate the maximum feeding performance of a wide variety of vertebrates. However, bite force studies of fishes have focused primarily on small species due to the intractable nature of large apex predators. More massive muscles can generate higher forces and many of these fishes attain immense sizes; it is unclear how much of their biting performance is driven purely by dramatic ontogenetic increases in body size versus size-specific selection for enhanced feeding performance. In this study, we investigated biting performance and feeding biomechanics of immature and mature individuals from an ontogenetic series of an apex predator, the bull shark, Carcharhinus leucas (73–285 cm total length). Theoretical bite force ranged from 36 to 2128 N at the most anterior bite point, and 170 to 5914 N at the most posterior bite point over the ontogenetic series. Scaling patterns differed among the two age groups investigated; immature bull shark bite force scaled with positive allometry, whereas adult bite force scaled isometrically. When the bite force of C. leucas was compared to those of 12 other cartilaginous fishes, bull sharks presented the highest mass-specific bite force, greater than that of the white shark or the great hammerhead shark. A phylogenetic independent contrast analysis of anatomical and dietary variables as determinants of bite force in these 13 species indicated that the evolution of large adult bite forces in cartilaginous fishes is linked predominantly to the evolution of large body size. Multiple regressions based on mass-specific standardized contrasts suggest that the evolution of high bite forces in Chondrichthyes is further correlated with hypertrophication of the jaw adductors, increased leverage for anterior biting, and widening of the head. Lastly, we discuss the ecological significance of positive allometry in bite force as a possible “performance gain” early in the life history of C. leucas.     

Proportions of the jaw mechanism of cichlid fishes changes and their meaning

The jaw mechanism of cichlid fishes is an intricate apparatus with complex force transmission from muscles to environment. The proportions of this apparatus change considerably during growth mainly due to scale effects. In adult fishes, the proportions differ, corresponding with the type of preferred food. In such a complex mechanism, it is very hard to gain insight into the functional meaning of the differences in proportions, unless a biomechanical model is constructed, describing kinematics and force equilibria of the apparatus.Such a model has been constructed and by means of perturbation analysis anatomical points could be selected of which the positions are very important for the function of biting. These hot spots need only small displacements in order to obtain an increase in biting force. By means of a newly developed mathematical technique, it was possible to move the anatomical points in very small steps towards more favourable positions for the function of biting.In this way, an improved version of an existing not very powerful biting insect feeding species, Haplochromis elegans, was calculated. This calculated version resembles a more powerful biting species Haplochromis nigricans very closely. Also, the functional meaning of the proportional changes during growth of the insect feeding Haplochromis elegans could be evaluated. In both cases compromises could be indicated where functional demands are in conflict with each other on the level of the anatomical design of the jaw mechanism.     

High prevalence of non-synonymous substitutions in mtDNA of cichlid fishes from Lake Victoria

When a population size is reduced, genetic drift may fix slightly deleterious mutations, and an increase in nonsynonymous substitution is expected. It has been suggested that past aridity has seriously affected and decreased the populations of cichlid fishes in Lake Victoria, while geographical studies have shown that the water levels in Lake Tanganyika and Lake Malawi have remained fairly constant. The comparably stable environments in the latter two lakes might have kept the populations of cichlid fishes large enough to remove slightly deleterious mutations. The difference in the stability of cichlid fish population sizes between Lake Victoria and the Lakes Tanganyika and Malawi is expected to have caused differences in the nonsynonymous/synonymous ratio, ω (= d_N/d_S), of the evolutionary rate. Here, we estimated ω and compared it between the cichlids of the three lakes for 13 mitochondrial protein-coding genes using maximum likelihood methods. We found that the lineages of the cichlids in Lake Victoria had a significantly higher ω for several mitochondrial loci. Moreover, positive selection was indicated for several codons in the mtDNA of the Lake Victoria cichlid lineage. Our results indicate that both adaptive and slightly deleterious molecular evolution has taken place in the Lake Victoria cichlids' mtDNA genes, whose nonsynonymous sites are generally conserved.     

Dealing with food and eggs in mouthbrooding cichlids: structural and functional trade-offs in fitness related traits

Background

As in any vertebrate, heads of fishes are densely packed with functions. These functions often impose conflicting mechanical demands resulting in trade-offs in the species-specific phenotype. When phenotypical traits are linked to gender-specific parental behavior, we expect sexual differences in these trade-offs. This study aims to use mouthbrooding cichlids as an example to test hypotheses on evolutionary trade-offs between intricately linked traits that affect different aspects of fitness. We focused on the oral apparatus, which is not only equipped with features used to feed and breathe, but is also used for the incubation of eggs. We used this approach to study mouthbrooding as part of an integrated functional system with diverging performance requirements and to explore gender-specific selective environments within a species.

Methodology/Principal Findings

Because cichlids are morphologically very diverse, we hypothesize that the implications of the added constraint of mouthbrooding will primarily depend on the dominant mode of feeding of the studied species. To test this, we compared the trade-off for two maternal mouthbrooding cichlid species: a “suction feeder” (Haplochromis piceatus) and a “biter” (H. fischeri). The comparison of morphology and performance of both species revealed clear interspecific and intersex differences. Our observation that females have larger heads was interpreted as a possible consequence of the fact that in both the studied species mouthbrooding is done by females only. As hypothesized, the observed sexual dimorphism in head shape is inferred as being suboptimal for some aspects of the feeding performance in each of the studied species. Our comparison also demonstrated that the suction feeding species had smaller egg clutches and more elongated eggs.

Conclusions/Significance

Our findings support the hypothesis that there is a trade-off between mouthbrooding and feeding performance in the two studied haplochromine cichlids, stressing the importance of including species-specific information at the gender level when addressing interspecific functional/morphological differences.     

Interspecific differences in prey items in relation to morphological characteristics among four lutjanid species (Lutjanus decussatus, L. fulviflamma, L. fulvus and L. gibbus)

Clarifying interspecific differences in prey items in relation to morphological characteristics is a fundamental aspect to understand the mechanism enabling the diversity of feeding ecology of fishes. The aim of the present study was to clarify the relationship between prey items and body shape variation, teeth and mandible characteristics for four lutjanid species: Lutjanus decussatus, L. fulviflamma, L. fulvus and L. gibbus. Stomach contents analysis revealed that the main prey items of L. decussatus were fishes, L. fulviflamma were crabs and fishes, and L. fulvus and L. gibbus was crabs. Body shape analysis revealed that L. decussatus and L. fulviflamma had a shallower body depth whereas L. fulvus and L. gibbus had a deeper body depth. The two species with a shallower body had long teeth whereas the other two species with a deeper body depth had shorter teeth. The jaw-lever mechanics were compared and L. decussatus and L. fulviflamma have a faster mouth opening–closing mechanism. Canonical correspondence analysis revealed that fishes was the major prey item for the species having a shallower body depth, higher teeth length and lower ratios of in-lever to out-lever of mandibles, whereas crabs was the major prey item for the species having a deeper body depth, lower teeth length and higher ratios of in-lever to out-lever of mandibles. It is suggested that the interspecific differences in main prey items among the four species are directly related to behavioral differences based on body shape, teeth characteristics and jaw-lever mechanics.     

Feeding Kinematics,Suction, and Hydraulic Jetting Performance of Harbor Seals (Phoca vitulina)

The feeding kinematics, suction and hydraulic jetting capabilities of captive harbor seals (Phoca vitulina) were characterized during controlled feeding trials. Feeding trials were conducted using a feeding apparatus that allowed a choice between biting and suction, but also presented food that could be ingested only by suction. Subambient pressure exerted during suction feeding behaviors was directly measured using pressure transducers. The mean feeding cycle duration for suction-feeding events was significantly shorter (0.15±0.09 s; P<0.01) than biting feeding events (0.18±0.08 s). Subjects feeding in-water used both a suction and a biting feeding mode. Suction was the favored feeding mode (84% of all feeding events) compared to biting, but biting comprised 16% of feeding events. In addition, seals occasionally alternated suction with hydraulic jetting, or used hydraulic jetting independently, to remove fish from the apparatus. Suction and biting feeding modes were kinematically distinct regardless of feeding location (in-water vs. on-land). Suction was characterized by a significantly smaller gape (1.3±0.23 cm; P<0.001) and gape angle (12.9±2.02°), pursing of the rostral lips to form a circular aperture, and pursing of the lateral lips to occlude lateral gape. Biting was characterized by a large gape (3.63±0.21 cm) and gape angle (28.8±1.80°; P<0.001) and lip curling to expose teeth. The maximum subambient pressure recorded was 48.8 kPa. In addition, harbor seals were able to jet water at food items using suprambient pressure, also known as hydraulic jetting. The maximum hydraulic jetting force recorded was 53.9 kPa. Suction and hydraulic jetting where employed 90.5% and 9.5%, respectively, during underwater feeding events. Harbor seals displayed a wide repertoire of behaviorally flexible feeding strategies to ingest fish from the feeding apparatus. Such flexibility of feeding strategies and biomechanics likely forms the basis of their opportunistic, generalized feeding ecology and concomitant breadth of diet.     

Prey capture and processing behaviors vary with prey size and shape in Australian and subantarctic fur seals

When hunting at sea, pinnipeds should adapt their foraging behaviors to suit the prey they are targeting. We performed captive feeding trials with two species of otariid seal, Australian fur seals (Arctocephalus pusillus doriferus) and subantarctic fur seals (Arctocephalus tropicalis). This allowed us to record detailed observations of how their foraging behaviors vary when presented with prey items that cover the full range of body shapes and sizes encountered in the wild. Small prey were captured using suction alone, while larger prey items were caught in the teeth using raptorial biting. Small fish and long skinny prey items could then be swallowed whole or processed by shaking, while all prey items with body depths greater than 7.5 cm were processed by shaking at the water's surface. This matched opportunistic observations of feeding in wild Australian fur seals. Use of “shake feeding” as the main prey processing tactic also matches predictions that this method would be one of the only tactics available to aquatic tetrapods that are unable to secure prey using their forelimbs.     

PHENOTYPIC PLASTICITY AND HETEROCHRONY IN CICHLASOMA MANAGUENSE (PISCES,CICHLIDAE) AND THEIR IMPLICATIONS FOR SPECIATION IN CICHLID FISHES

Cichlid fishes in African rift lakes have undergone rapid speciation, resulting in “species flocks” with more than 300 endemic species in some of the lakes. Most researchers assume that there is little phenotypic variation in cichlid fishes. I report here extensive phenotypic plasticity in a Neotropical cichlid species. I examined the influence of diet on trophic morphology during ontogeny in Cichlasoma managuense. Two groups of full siblings were fed two different diets for eight months after the onset of feeding; thereafter both groups were fed a common diet. Phenotypes that differed significantly at 8.5 months converged almost completely at 16.5 months. If feeding on two different diets is continued after 8.5 months, the phenotypes remain distinct. Differences in diet and possibly in feeding mode are believed to have caused these phenotypic changes. Phenotypic plasticity is described in terms of a qualitative model of heterochrony in which phenotypic change in morphology is explained as retardation of the normal developmental rate. If phenotypic expression of morphology is equally plastic in African cichlid species as it may be in the American cichlids, as exemplified by C. managuense, then taxonomic, ecological, and evolutionary analyses of “species flocks” may be in need of revision. However, Old World cichlids may be less phenotypically plastic than New World cichlids, and this may contribute to the observed differences in speciation rate and degree of endemism.