Functional morphology of comminuting feeding structures of Trichodactylus borellianus (Brachyura,Decapoda, Trichodactylidae), an omnivorous freshwater crab

Crustaceans exhibit great diversity of feeding structures with morphological traits that are useful to infer the general trophic habits of species. In this study, we analyzed the functional morphology of comminuting feeding structures (mandibles, chelipeds, gastric mill) of the freshwater crab Trichodactylus borellianus directly related with the food fragmentation. The heterochely and mechanical advantage (MA) of the chelae were also studied. In both analyses, we considered the relationship between morphology and the natural diet. We expected to find a consistent relation between feeding habits and morphological traits. In general, we found simple structures armed with uniform setal systems and feeding appendages without pronounced teeth or spines. Mandibles have primarily cutting functions, helping with the food anchoring and fragmentation with mandibular palps armed with pappose setae. Chelipeds were covered with spines and simple setae. Adult males exhibited right-handedness with high MA of the major chelae. The ingested, relatively large pieces of food are finally chewed by a gastric mill equipped with sharp cusps characteristic of decapods with low ingestion of crude fiber material. The morphology of the feeding apparatus revealed that it is well adapted to an omnivorous diet, being able to cope with dietary changes.     

Development and metamorphosis of the digestive system of larval lobsters,Homarus americanus (Decapoda: Nephropidae)

This study provides a detailed account of the development of the digestive system of larval lobsters (Homarus americanus H. Milne Edwards, 1837) and the morphological changes that occur at metamorphosis. The most dramatic of these changes involves the gastric mill of the cardiac stomach. First-and second-stage lobsters lack the medial and lateral teeth characteristic of the grinding stomach of adult lobsters. Clearly recognizable, heavily cuticularized teeth first appear in the third stage, and accessory lateral teeth do not appear until the fourth stage. In place of the teeth of the gastric mill, first- and second-stage stomachs have a series of pads and ridges which are the apparent rudiments of the teeth. The development of the gastric mill during the larval stages enables lobsters to deal successfully with the more substantial food they encounter in the benthic environment, and corresponds to the drastic change of habitat and diet which occurs at metamorphosis. Confusion about the extent of the midgut and hindgut in larval lobsters has been clarified. The results of this study have shown that the larvae have a long midgut, which lacks a cuticle, and a short hindgut with a cuticular lining, just as in adult lobsters. The junction between midgut and hindgut lies in the sixth abdominal segment in all of the first four stages, as well as in the adult.     

Silicification of the medial tooth in the blue crab Callinectes sapidus

Observations of cuticular structures mineralized with silica within the Crustacea have been limited to the opal teeth of copepods, mandibles of amphipods, and recently the teeth of the gastric mill in the blue crab Callinectes sapidus. Copepod teeth are deposited during premolt, with sequential elaboration of organic materials followed by secretion of silica into the tooth mold. The timing of mineralization is in stark contrast to that of the general integument of crustaceans in which calcification is completely restricted to the postmolt period. To determine the timing of molt‐related deposition and silicification of the teeth of the gastric mill, the medial tooth of the blue crab C. sapidus was examined histologically and ultrastructurally across the molt cycle. Histological data revealed deposition of the organic matrix of the epicuticle and exocuticle during premolt. No evidence of postmolt changes in the thickness of the epicuticle and exocuticle, or any deposition of endocuticle, was observed. Scanning electron microscopy revealed degradation of the outer surface of the old tooth during premolt. During premolt, epithelial structures resembling papilla appeared to secrete a fibrous web that coalesces to become the matrix of the new tooth. Semi‐quantitative elemental analyses indicated simultaneous deposition of silica and organic matrix, and demonstrated a homogeneous distribution of silicon throughout the epicuticle of the tooth at all stages. However, there is evidence of deposition (presumably silicification) during postmolt as spaces between the papillae become filled in. Thus, the pattern and timing of deposition and silicification of the tooth are different from both teeth of copepods and the general exoskeleton of decapods, and may facilitate rapid resumption of feeding and consumption of the exuvia in early postmolt. J. Morphol. 277:1648–1660, 2016. © 2016 Wiley Periodicals, Inc.     

Foregut morphology and ontogeny of the spider crab Maja brachydactyla (Brachyura,Majoidea, Majidae)

We describe the morphology of the foregut of the spider crab Maja brachydactyla Balss, 1922, from first larval stage to adult, with detailed stage‐specific documentation using light and scanning electron microscopy. A total of 40 ossicles have been identified in the foregut of adults of M. brachydactyla using Alizarin‐Red staining. The morphological pattern of the ossicles and gastric mill is very similar to other Majoidea species with only a few variations. The foregut of the zoeae stages appeared as a small and simple cavity, with a cardio‐pyloric valve that separates the stomach into cardiac and pyloric regions. The pyloric filter is present from the first zoea, in contrast to the brachyuran species which have an extended larval development. Calcified structures have been identified in the cardio‐pyloric valve and pyloric region of the zoeal stages. The most significant changes in foregut morphology take place after the metamorphosis from ZII to megalopa, including the occurrence of the gastric mill. In the megalopa stage, the foregut ossicles are recognizable by their organization and general morphology, but are different from the adult phase in shape and number. Moreover, the gastric teeth show important differences: the cusps of the lateral teeth are sharp (no molariform); the dorsal tooth have a small, dentate cusp (not a well‐developed quadrangular cusp); and the accessory teeth are composed of one sharp peak (instead of four sharp peaks). The gastric mill ontogeny from megalopa to adult reveals intermediate morphologies during the earlier juvenile stages. The relationship between gastric mill structures with food preferences and their contribution to the brachyuran phylogeny are briefly discussed. J. Morphol. 276:1109–1122, 2015. © 2015 Wiley Periodicals, Inc.     

The gastric mill of euphausiid crustaceans: a comparison of eleven species

The morphology of the gastric mill of 11 species in the family Euphausiidae is described. Two distinct arrangements can be recognized with respect to presence or absence of lateral teeth. One is the well-developed gastric mill of Thsanoessa gregaria, T. inermis, T. macrura and Pseudeuphausia sinica; the other is the gastric mill without lateral teeth of Nematoscelis atlantica, N. difficilis, N. megalops, Nematobrachion flexipes, N. sexspinosum, Stylocheiron abbreviatum, and S. carinatum. This supports the close relationships of the four species of both genera, Thysanoessa and Pseudeuphausia on the one hand and seven species belonging to Nematobrachion, Nematoscelis, and Stylocheiron on the other hand. The hypothesis that species in the same genus have similar gastric mill morphology, which was once rejected in the genus Thysanopoda, is accepted in all five genera examined here. Functional morphology of euphausiid foreguts is discussed.     

Comparative digestive biology between the ponyfishes from the Pranburi River estuary,Thailand

We investigated the digestive biology of two prevalent leiognathid species in Pranburi River estuary, Thailand: the decorated ponyfish (Nuchequula gerreoides) and the splendid polyfish (Eubleekeria splendens). A total of 632 samples collected from February to April and September to November 2017 were analysed using morphological and histological approaches. The overall structures were similar between the species: a short mucous-cell-rich oesophagus region, a well-developed gastric gland uniformly present across the stomach's mucosal layer, and three finger-like pyloric caeca between the stomach and intestine. However, there were marked differences in the mouth, gill raker, and intestinal coefficient (IC). N. gerreoides had a relatively longer mouth, smoother gill rakers, and an IC of 1.08 ± 0.01, similar to those of other carnivorous fish. In contrast, the gill raker of E. splendens had more villiform teeth that can filter-feed better, and their IC was 2.16 ± 0.02 (i.e., longer intestine). Although digestive structures were generally similar between the ponyfishes, these differences suggest that N. gerreoides is relatively carnivorous with stronger suction, whereas E. splendens may be an omnivorous or herbivorous filter-feeder.     

Development and metamorphosis of the feeding apparatus of the stone crab,Menippe mercenaria (brachyura,xanthidae)

Analysis of the feeding apparatus of the stone crab, Menippe mercenaria (Brachyura, Xanthidae), has demonstrated that substantial internal and external morphological alterations occur at metamorphosis and suggests that the mastication of food shifts from the mandibles to the gastric mill at that time. These changes correspond to the changes in environment and diet that take place at metamorphosis, when the previously planktotrophic larvae begin benthic life. A detailed account of the structure and development of the mandibles is presented. The mandibles of all zoeal stages are similar: The incisor process has a series of teeth and denticles and the prominent molar process appears to be well adapted for grinding food. Megalopal mandibles are transitional but have the form that is typical of all subsequent stages: The expanded incisor process is rounded and toothless and the molar process is less prominent and has lost its grinding denticles. The cardiac stomach of the zoeal stages has no gastric mill; the medial and lateral teeth of the mill first appear in the megalopa. A very simple procedure is described for preparing larval mandibles for scanning electron microscopy using the molted exoskeletons from larval rearing experiments.     

Anatomy and histology of the gastrointestinal tract of the neo-tropical opossum (Didelphis marsupialis insularis,Allen 1902)

The morphology of the gastrointestinal tract (GI) is a strong indicator of the dietary habits of a species. The goal of this study was to describe the gross and microanatomy of the digestive tract of the neo-tropical opossum (Didelphis marsupialis insularis) and relate them to the animals’ food habits. GI tracts from 12 adult animals were used for this study. Results found the small intestine made up 65.9% of the GI tract, and the stomach was simple with a prominent caecum. Histologically, the oesophageal mucosa was non-keratinized and glands were found throughout the oesophagus. The large intestine showed a great number of goblet cells, the jejunum possessed well-developed villi, and Peyer's patches were absent in the ileum. The absence of keratinization of the epithelial lining of the oesophagus and stomach and a high lymphocytic infiltration throughout the small and large intestine reflected a more carnivorous diet, whereas the presence of a well-developed caecum in the large intestine indicated the ability to digest plant matter. Overall, the morphology of GI tract of D. m. insularis displayed both carnivorous and herbivorous features, allowing us to conclude that it is an omnivorous animal.     

Ecomorphological correlates of craniodental variation in bears and paleobiological implications for extinct taxa: an approach based on geometric morphometrics

Relative warp analyses of landmarks describing cranial and mandibular shape are used for investigating patterns of morphological variation among extant bears (Mammalia, Carnivora, Ursidae) indicative of diet and feeding behavior. These patterns are used for deriving inferences about the autecology of two extinct species previously assumed to have had different dietary preferences, the North American giant, short-faced bear Arctodus simus and the Eurasian cave bear Ursus spelaeus . Results reveal a set of shared craniodental traits among the herbivorous bears, including short and vaulted skulls with well-developed zygomatic arches, lateralized orbits and small canines, concave jaws with a highly positioned condyle, large moment arms for the temporalis and masseter muscles, and long cheek teeth. In contrast, those bears that consume animal resources have long skulls with small zygomatic arches, frontalized orbits and well-developed canines, and long jaws with a deep mandibular symphysis, low muscle leverages, a condyle situated at the level of the tooth row and reduced cheek teeth. The craniodental morphology of omnivorous bears is intermediate between those of faunivores and herbivores. This is also the case of the short-faced bear and the cave bear, which suggests that previous reconstructions of the feeding ecology of these extinct species (highly carnivorous for A. simus and herbivorous for U. spelaeus ) should be revised.     

Dental Histology of a Characoid Fish from the Plio-Pleistocene of Acre, Brazil

The histology of premaxillary teeth of Colossoma sp. from the Solimöes Formation of Northern Brazil is described, in comparison with Recent material. Analysis by light and scanning electron microscopy shows that they share with those of the serrasalminids, histological (though not anatomical) features of carnivorous fishes. The enameloid in both groups displays surface-parallel images of calcified fibres along the periphery of the teeth while internally, these images present a random distribution. Such similarities suggest that both the serrasalminids and myleinids may have evolved from carnivorous ancestors. The serrasalminids remain carnivorous, while the herbivorous habit of the myleinids may have been acquired secondarily. This change of diet is probably related to changes in the shape of the teeth which are used for crushing fruits, seeds and leaves. The teeth in adult Colossoma are unicuspid, though not conical. The molar or incisive form of these teeth may have evolved independently from the conical-shaped primitive dentition of the characoids, without passing through any stage of fusion of dental papillae, which is observed in the ontogenetic development of other characoids.     

Feeding Habits of Percnon gibbesi (H. Milne Edwards, 1853) in the Sicily Strait

The subtropical grapsid crab Percnon gibbesi (H. Milne Edwards, 1853) is one of the most recent alien decapods found in the Mediterranean, where it was discovered at Linosa (Pelagie Islands, Sicily Strait) in summer 1999. At present, the invasion of this species has been recorded in several other insular localities. We studied the feeding habits of the crab in an attempt to understand its success in the Mediterranean. The morphological characteristics of the chela, the feeding adaptations of the gastric mill and the analysis of stomach contents indicate that P. gibbesi is a strictly herbivorous species, a characteristic not shared with any other large-sized infralittoral Mediterranean crab.     

Marginal tooth-bearing bones in the lower jaw of the recent australian lungfish,Neoceratodus forsteri (Osteichthyes,Dipnoi)

Developmental studies of the Recent Australian lungfish, Neoceratodus forsteri, show that this species has two sets of functional tooth-bearing bones in the lower jaw of young hatchlings. These coincide with an early stage in the life history when the fish is strictly carnivorous. In N. forsteri, a paired tooth-bearing dentary and an unpaired symphyseal bone and tooth develop slightly later than the permanent vomerine, prearticular, and pterygopalatine tooth plates, which appear at stage 44 of development, and erupt with the permanent dentition between stages 46 and 48, when the hatchling first starts to feed on small aquatic invertebrates. At these stages of development, all of the teeth are long, sharp, and conical and help to retain prey items in the mouth. Disappearance of the transient dentition coincides with complete eruption of the permanent tooth plates and precedes the change to an omnivorous diet. Existence of a transient marginal dentition in this species of lungfish suggests that the presence of an apparently similar marginal dentition in adults of many species of Palaeozoic dipnoans should be considered in phylogenetic analyses of genera within the group, and when analysing the relationships of dipnoans with other primitive animals. © 1995 Wiley-Liss, Inc.     

Costs of carnivory: tooth fracture in Pleistocene and Recent carnivorans

Large, carnivorous mammals often break their teeth, probably as a result of tooth to bone contact that occurs when carcasses are consumed more fully, a behaviour likely to occur under conditions of food stress. Recent studies of Pleistocene predators revealed high numbers of teeth broken in life, suggesting that carcass utilization and, consequently, food competition was more intense in the past than at present. However, the putative association between diet and tooth fracture frequency was based on a small sample of large, highly carnivorous species. In the present study, a greater diversity of extant carnivorans is sampled, including insectivorous, omnivorous, and carnivorous forms, ranging in size from weasels to tigers. Species that habitually consume hard foods (bones, shells) had the highest fracture frequencies, followed by carnivores, and then insectivorous and/or omnivorous species. Predator and prey sizes were not associated with tooth fracture frequency, but more aggressive species did break their teeth more often. Comparison of the modern sample with five Pleistocene species confirms the previous finding of higher tooth breakage in the past, although some extant species have fracture frequencies that approach those of extinct species. Thus, the Pleistocene predator guild appears to have been characterized by relatively high levels of competition that are rarely observed today.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 68–81.     

Food utilisation and digestive ability of aquatic and semi-terrestrial crayfishes, <Emphasis Type="Italic">Cherax destructor</Emphasis> and <Emphasis Type="Italic">Engaeus sericatus</Emphasis> (Astacidae,Parastacidae)

Both Engaeus sericatus and Cherax destructor are omnivorous crayfishes consuming a variety of food items. Materials identified in the faeces of both E. sericatus and C. destructor consisted of mainly plant material with minor amounts of arthropod animals, algae and fungi. The morphology of the gastric mill of C. destructor suggests that it is mainly involved in crushing of food material while the gastric mill of E. sericatus appears to be better suited to cutting of food material. Given this, the gastric mill of E. sericatus may be better able to cut the cellulose and hemicellulose fibres associated with fibrous plant material. In contrast, the gastric mill of C. destructor appears to be more efficient in grinding soft materials such as animal protein and algae. Both species accumulated high amounts of lipids in their midgut glands (about 60% of the dry mass) which were dominated by triacylglycerols (81–82% of total lipids). The dominating fatty acids were 16:0, 16:1(n-7), 18:1(n-9), 18:2(n-6), and 18:3(n-3). The two latter fatty acids can only be synthesised by plants, and are thus indicative of the consumption of terrestrial plants by the crayfishes. The similarity analysis of the fatty acid patterns showed three distinct clusters of plants and each of the crayfish species. The complement of digestive enzymes, proteinases, total cellulase, endo-β-1,4-glucanase, β-glucosidase, laminarinase and xylanase within midgut gland suggests that both C. destructor and E. sericatus are capable of hydrolysing a variety of substrates associated with an omnivorous diet. Higher activities of total cellulase, endo-β-1,4-glucanase and β-glucosidase indicate that E. sericatus is better able to hydrolyse cellulose within plant material than C. destructor. In contrast to E. sericatus, higher total protease and N-acetyl-β-d-glucosaminidase activity in the midgut gland of C. destructor suggests that this species is better able to digest animal materials in the form of arthropods. Differences in total cellulase and gastric mill morphology suggest that E. sericatus is more efficient at digesting plant material than C. destructor. However, the contents of faecal pellets and the fatty acid compositions seem to indicate that both species opportunistically feed on the most abundant and easily accessible food items.     

Characterization of the ultrastructure of the gastrointestinal tract mucosa, stomach contents and liver enzyme activity of the pinfish during development

Carnivorous juveniles (<16 mm L_S) of pinfish Lagodon rhomboides apparently lacked gastric glands in the stomach while larger fish, intermediates (30-33 mm L_S) and herbivorous adults (>80 mm L_S) had numerous gastric glands. Two cell types were identified in the gastric glands of larger fish: mucous secreting cells and secretory cells which had ultrastructure features typical of digestive enzymes and acid secretion. Lipid absorption occurred throughout the caeca and intestine in all sizes of fish. Microvilli found on the rectal epithelial cells of the intermediate and adult pinfish occurred on stalks and were possibly associated with water reabsorption. Liver enzyme activities changed in small fish (26–39 mm L_S) compared to the adult fish. Alanine amino transferase (ALT) and fructose diphosphotase (FDP) activities declined while pyruvate kinase (PK) activities increased significantly. These changes were consistent with a change in diet from a carnivorous (high protein) diet in juveniles to an omnivorous (lower protein) diet determined by stomach content analyses.     

Morphologie des Gebisses und Nahrungspraferenzen von dreizehn adriatischen Blenniiden (Teleostei, Perciformes)

The morphology and arrangement of the teeth in 13 sympatric species of the genus Blennius were investigated by SEM. All the species were collected in the upper littoral of the Northern Adriatic near Rovinj (Yugoslavia) and represent the total species inventory of the genus Blennius in this area. After analysing the content and length of the intestine, four groups with different feeding habits could be distinguished to which the morphological adaptations of the teeth could be correlated. There is a non-selective omnivorous group of Aufwuchs-grazer which take the inhabiting vagilfauna {Blennius gattorugine, B. incognitas, B. tentacularis ), a herbivorous group ( B. sanguinolentus, B. sphynx, B. canevae ), a carnivorous group ( B. trigloides, B. dalmatinus, B. rouxi ), and a linking group between omnivorous grazers and carnivores (B. adriaticus, B. pavo, B. nigriceps). B. galerita shows a preponderance of comblike teeth and feeds exclusively scraping microscopic algae from the rocky substrate. The morphology and position of the caniniforms vary greatly within the investigated species. The dentition on the vomer is lacking in five species and seems of no consequence in the uptake of food.     

THE DIET OF OSTRICH DINOSAURS (THEROPODA: ORNITHOMIMOSAURIA)

Abstract:  The diets of ornithomimosaurian dinosaurs (Theropoda: Ornithomimosauria) have proved to be contentious owing to a dearth of unambiguous evidence in support of carnivory, omnivory or herbivory. Re-assessment of anatomical, taphonomical and palaeoecological evidence, and estimates of daily minimal energy budgets for two derived ornithomimosaurian genera, indicate that suspension-feeding and carnivory were unlikely. The combined presence of a keratinized rhamphotheca and gastric mill is strongly indicative of a herbivorous habitus for these dinosaurs. Herbivorous and omnivorous forms are rare among the non-avian Theropoda, but are more common than previously suspected. Rejection of carnivorous habits for derived ornithomimosaurs redresses apparent discrepancies in the relative abundances of the herbivore and carnivore guilds of several Late Cretaceous faunas.     

Tooth and consequences: Heterodonty and dental replacement in piranhas and pacus (Serrasalmidae)

Tooth replacement in piranhas is unusual: all teeth on one side of the head are lost as a unit, then replaced simultaneously. We used histology and microCT to examine tooth‐replacement modes across carnivorous piranhas and their herbivorous pacu cousins (Serrasalmidae) and then mapped replacement patterns onto a molecular phylogeny. Pacu teeth develop and are replaced in a manner like piranhas. For serrasalmids, unilateral tooth replacement is not an “all or nothing” phenomenon; we demonstrate that both sides of the jaws have developing tooth rows within them, albeit with one side more mineralized than the other. All serrasalmids (except one) share unilateral tooth replacement, so this is not an adaptation for carnivory. All serrasalmids have interlocking teeth; piranhas interdigitate lateral tooth cusps with adjacent teeth, forming a singular saw‐like blade, whereas lateral cusps in pacus clasp together. For serrasalmids to have an interlocking dentition, their teeth need to develop and erupt at the same time. We propose that interlocking mechanisms prevent tooth loss and ensure continued functionality of the feeding apparatus. Serrasalmid dentitions are ubiquitously heterodont, having incisiform and molariform dentitions reminiscent of mammals. Finally, we propose that simultaneous tooth replacement be considered as a synapomorphy for the family.     

Foregut morphology and ontogeny of the mud crab Dyspanopeus sayi (Smith, 1869) (Decapoda,Brachyura, Panopeidae)

The morphology of the foregut of the Say's mud crab Dyspanopeus sayi was described in adults and larvae. The ossicle system was illustrated based on a staining method with Alizarin-Red. The gastric teeth and cardio-pyloric valve were dissected and examined using optical and scanning electron microscopy. In the adults, the morphology of ossicles and gastric teeth of D. sayi is very similar to the related species Rhithropanopeus harrisii. The foregut of first zoea (ZI) presented a functional cardio-pyloric valve while the filter press was lacking. The filter press was observed in the pyloric chamber from ZII. The most significant changes in morphology take place after metamorphosis from ZIV to megalopa, including the occurrence of the gastric mill. The organization and morphology of many megalopal foregut ossicles are recognizable in the adult phase, although the morphology of the gastric teeth differs from the morphology of adults. A correlation of gastric mill structures with food preferences and their contribution to the phylogeny are briefly discussed.