Specialized morphology for a generalist diet: evidence for Liem's Paradox in a cichlid fish

The stable isotope ratio and seasonal changes in diet of Alluaud's haplo Astatoreochromis alluaudi, a cichlid fish with massive pharyngeal jaws well known for its ability to process hard‐bodied prey, are described. The diet of A. alluaudi was quantified in Lake Saka, Uganda, over a period of 30 months. Variation in physico‐chemical variables (mean monthly rainfall, water temperature, turbidity and dissolved oxygen), as well as potential competitor density and food abundance, was measured throughout the second half of the study (14 months). Stomach contents and isotope analysis revealed a diet comprised mainly of fishes and insects, with a low contribution of molluscs (0–33%) in any given month. No correlation was detected between diet and either macroinvertebrate abundance or competitor abundance. The running average rainfall was positively related to the percentage of fish consumed per month. Although A. alluaudi exhibits an apparent molluscivorous trophic morphology in Lake Saka, molluscs did not appear to compose a major portion of its diet. Gradients of rainfall seemed to be the most important environmental predictor of diet choice in Lake Saka. These results are discussed with reference to Liem's Paradox that apparently morphologically specialized fishes often function as generalist feeders in the wild.     

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.     

Convergence in a mechanically complex phenotype: detecting structural adaptations for crushing in cichlid fish

Morphological convergence provides strong evidence that evolution is adaptive. However, putatively convergent morphology is often examined in two dimensions with no explicit model of function. In this study, we investigated structural and mechanical similarities of the lower pharyngeal jaw (LPJ) in cichlid fish that have evolved the ability to crush hard-shelled molluscs. Using a novel phylogeny, we demonstrated molluscivory has been gained and/or been lost numerous times in Heroine cichlids. Within this comparative framework, we produced three-dimensional computed tomography (CT) scans for LPJs of both morphotypes in the trophically polymorphic Herichthys minckleyi and six evolutionarily independent pairs of closely related species. Like H. minckleyi , these species exhibit divergence between a molluscivore and a nonmolluscivore. Using the CT scans, we generated finite element models of papilliform H. minckleyi LPJs to determine where stress would concentrate in a jaw not modified to crush molluscs. Then, we examined whether stress in the papilliform LPJ predicted structural modifications in molariform H. minckleyi and other molluscivorous species. Despite potential constraints, stresses imposed during prey processing explain 40% of LPJ variation in mollusc crushing species. The structural and mechanical analyses also suggest divergence found in polymorphic species could provide the substrate for trophic differences found in reproductively isolated cichlids.     

A unique feeding strategy of the extinct marine mammal Kolponomos: convergence on sabretooths and sea otters

Mammalian molluscivores feed mainly by shell-crushing or suction-feeding. The extinct marine arctoid, Kolponomos, has been interpreted as an otter-like shell-crusher based on similar dentitions. However, neither the masticatory biomechanics of the shell-crushing adaptation nor the way Kolponomos may have captured hard-shelled prey have been tested. Based on mandibular symphyseal morphology shared by Kolponomos and sabre-toothed carnivores, we hypothesize a sabretooth-like mechanism for Kolponomos prey-capture, whereby the mandible functioned as an anchor. Torque generated from jaw closure and head flexion was used to dislodge prey by prying, with prey then crushed using cheek teeth. We test this hypothesized feeding sequence using phylogenetically informed biomechanical simulations and shape analyses, and find a strongly supported, shared high mandibular stiffness in simulated prey-capture bites and mandibular shape in Kolponomos and the sabre-toothed cat Smilodon. These two distantly related taxa converged on using mandibles to anchor cranial torqueing forces when prying substrate-bound prey in the former and sabre-driving forces during prey-killing in the latter. Simulated prey-crushing bites indicate that Kolponomos and sea otters exhibit alternative structural stiffness-bite efficiency combinations in mandibular biomechanical adaptation for shell-crushing. This unique feeding system of Kolponomos exemplifies a mosaic of form-function convergence relative to other Carnivora.     

Similar ecology,different morphology: Three new species of oral-mollusc shellers from Lake Edward

Lake Edward, East Africa, harbours a largely understudied assemblage of haplochromine cichlids that displays a range of adaptions to various specialised trophic niches. In this system, we discovered specimens of Haplochromis with morphologies similar to those of oral-mollusc shellers from Lake Victoria. These morphologies are characterised by short oral jaws with stout teeth that are used either to crush molluscs or to grab the soft bodies of snails and wrench them out of their shells. A morphometric study on 47 specimens from Lake Edward revealed the presence of three new species with an oral-shelling morphology: Haplochromis concilians sp. nov., H. erutus sp. nov. and H. planus sp. nov. All three species are formally described. Stomach-content observations confirmed an opportunistic oral-shelling ecology for H. concilians sp. nov. and H. erutus sp. nov. Within H. planus sp. nov., only large specimens displayed a specialised oral-shelling morphology, but their stomachs were nearly empty, while small specimens consumed mainly Ostracoda and Hydrachnidia. Remarkably, the three species differed considerably in morphology from each other, but they each resembled oral-sheller species from Lake Victoria.     

Hammer-toothed 'marsupial skinks' from the Australian Cenozoic

Extinct species of Malleodectes gen. nov. from Middle to Late Miocene deposits of the Riversleigh World Heritage Area, northwestern Queensland, Australia are enigmatic, highly specialized, probably snail-eating marsupials. Dentally, they closely resemble a bizarre group of living heterodont, wet forest scincid lizards from Australia (Cyclodomorphus) that may well have outcompeted them as snail-eaters when the closed forests of central Australia began to decline. Although there are scincids known from the same Miocene deposits at Riversleigh, these are relatively plesiomorphic, generalized feeders. This appears to be the most striking example known of dental convergence and possible competition between a mammal and a lizard, which in the long run worked out better for the lizards.     

Evolution of the relative width of the head and alveolar surfaces in map turtles (Testudines: Emydidae: Graptemys)

Replicate adaptive radiations are dichotomous morphological patterns related to resource use and repeated in a series of isolated habitats. Biologists have long noted a dichotomy in relative head width and alveolar width among species of map turtles ( Graptemys ), particularly with regard to adult females, which are much larger dian adult males in all 12 species. I measured plastron length (PL), head width (HW), and alveolar width (AW) of nearly 2300 specimens representing the 12 recognized species, and used allometric regressions to investigate this genus as an example of both sexual dimorphism in trophic morphology and replicate adaptive radiation. Sexual dimorphism was noted in HW after correction for PL, with females having wider heads than males of similar body sizes, but AW as corrected for HW was not dimorphic, thus absolute differences in AW are the simple result of differences in HW. Sexual differences in HW probably relate to the dimorphic niches of Graptemys , as females of three species have been reported to occupy deeper water further from shore than conspecific males. Based on predicted HW at maximum PL in adult females, species were categorized as mega-, meso-, or microcephalic. Differences in relative head width are related to differences in degree of molluscivory, with little molluscivory in microcephalic females, moderate to high molluscivory in mesocephalic females, and high molluscivory in megacephalic females. Distributions of Graptemys species in sympatry and allopatry reveal patterns consistent with structuring of distributions via competitive interactions. Superimposing data on relative HW on a phylogeny of the genus suggests there may have been one or two episodes of character displacement, widi subsequent character assortment, in Graptemys evolution. This conclusion is similar to the current model for the evolution of body size during the radation of Anolis lizards in the northern Lesser Antilles.