Functional morphology of the mouthparts and associated structures of Pagurus rubricatus (Crustacea: Decapoda: Anomura) with special reference to feeding and grooming

Summary Pagurus rubricatus is predatory, detrivorous, macrophagous, and to a small degree, a suspension feeder. The crab searches for small invertebrates by digging shallow pits in the sediment. During this process it feeds on detritus obtained either directly from the sediment or scoured off gravel granules. Particles trapped by the dense setation of the 2nd and 3rd maxillipeds are brushed off and ingested.The distribution of the various types of setae on the mouthparts is mapped and structure of the mouthparts and their setae is correlated with function. Sediment collected by the pereiopods is brushed off by the endopodites of the 3rd maxillipeds and transferred to the inner mouthparts by the endopodites of the 2nd maxillipeds. The basipodites of the 1st maxillae form a filter screen through which particles of suitable size are pushed by the 2nd maxillae. Rejected particles are discarded by the exhalant stream via the currents generated by the exopodites of the maxillipeds. Specialized setae on the 2nd maxillae scour detritus from the surface of gravel granules applied to these appendages by the 2nd and 3rd maxillipeds. Interlocking setae from different appendages form a number of screens the main function of which is to retain material in the buccal region. The exopodite and endopodite of the 1st maxilliped and the endopodites of the 1st and 2nd maxillipeds form a channel which funnels the exhalant respiratory current away from the crab. The main grooming appendages are the endopodites of the 3rd maxillipeds, however, most of the other mouthparts have a self-cleaning function.     

Fine structure and functional morphology of the mouthparts of Bittacus planus and Terrobittacus implicatus (Insecta: Mecoptera: Bittacidae)

The Bittacidae are unique in Mecoptera for their adults being predaceous. However, their mouthparts have not been well documented for functional morphology to date. Here, we investigated the mouthpart morphology of the hangingflies Bittacus planus Cheng and Terrobittacus implicatus (Huang & Hua) using scanning electron microscopy. The mouthparts are of the mandibulate type and situated at the tip of an elongated rostrum. The labrum is greatly elongated, roughly twice as long as the subquadrate clypeus. The epipharynx is furnished with a row of basiconic sensilla arranged evenly as a median band extending from the apex to the base. The mandibles are slender and elongated, bearing a sharp lateral and a small mesal tooth. The maxillae are well developed, each consisting of a partially sclerotized cardo and a stipes, a hirsute galea and a lacinia, and a five-segmented maxillary palp. The sensillar pattern on the distal segment of the maxillary palp differs slightly between the two bittacid species. The labium is composed of a postmentum, a prementum, and a pair of two-segmented labial palps. The feeding mechanism of bittacids is briefly discussed in combination with the mouthpart morphology and their feeding habits.     

Functional morphology and ultrastructure of mouthparts in unfed water mite larvae Piona carnea (Koch, 1836) (Acariformes: Pionidae)

Mouthparts of unfed larvae Piona carnea (Koch, 1836) (Acariformes: Pionidae) were studied on whole-mount preparations, semi-thin sections and with TEM and SEM methods. The mouth apparatus is incorporated within the pseudotagma, gnathosoma, composed of the infracapitulum and of the chelicerae resting on the roof of the latter. The gnathosoma inclines to the long axis of the body and is inserted at its base into the idiosoma by the circumcapitular fold. The basal cheliceral segments are long and fused. An anterior projection, the proposed fused fixed digits, protrudes from the distal end of the basal cheliceral segment forward between the movable digits. The movable digits are always found in protruded position, strongly curved upward and show a groove on their inner sides. The ventral wall of the infracapitulum is made of the mentum posterior and the malapophyses anterior to the palp articulation. The malapophyses are squeezed between the large palps and envelope the distal portion of the chelicerae from the sides. The ventral portion of the fused malapophyses are provided with a characteristic ventral cuticular fork of unknown function. Each malapophysis terminates by a flexible lateral lip provided by several rigid jags looking posterad. The palps face downward and backward, and bear on the tibia the large curved palpal claws turned laterad. The palp femur bears on the ventral aspect a characteristic wide spade-like projection provided with its own muscles originating on the dorsal wall of the femur. The labrum is a thick cuticular arrow-like structure protruding forward into the preoral cavity, whereas the cervix is a thin weakly sclerotized plate. The particular labral valve projects forward from the dorsal basis of the labrum into the preoral cavity. The labrum and the cervix are provided by their own small labral and cervical muscles originating on the cervical apodemes. The pharynx is totally separated from the ventral wall of the infracapitulum and is devoid of ventral dilators. The dorsal pharyngeal dilators originate on the thick and sclerotized capitular apodeme and, posteriorly, on the paired cuticular branches, capitular apodemes, which end freely in the body cavity and are combined with the common salivary duct. The short sigmoid pieces serve for origin of the levator muscles of the chelicerae. Retraction of the gnathosoma and the chelicerae is mediated by several sets of muscles originating on the dorsal plate.     

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.     

External morphology of Lightiella monniotae (Crustacea, Cephalocarida) in the light of Cambrian 'Orsten' crustaceans

The species-poor meiofaunal Cephalocarida have played an important role in discussions of the phylogeny and evolution of Crustacea since their discovery in 1955. One reason may be that the morphology of cephalocarids includes some aspects of putatively ancient appearance, such as the simple roof-shaped head shield, the anterior three head appendages resembling those of a nauplius larva, or the trunk-limb-like second maxilla. Cephalocarida have even been suggested to represent the sister taxon to all other Eucrustacea. Presence of possibly plesiomorphic characters, however, does not necessarily point to a basal position in the system. Growing evidence demonstrates that the modification of the fourth post-antennular cephalic appendage, the ‘maxilla’, into a “mouth part” may have occurred independently in the different eucrustacean lineages, so a trunk-limb-like maxilla is an ancient feature that does not hold only for cephalocarids. Retention of its plesiomorphic shape and function in the Cephalocarida remains, however, noteworthy. Cephalocarids are still little studied and incompletely known, especially their external morphology. By examining several adults and one young specimen of Lightiella monniotae Cals and Delamare Deboutteville, 1970 from New Caledonia, we aimed to a) document as many details as possible, and b) compare these data with other species of Cephalocarida. We also aimed to reconstruct aspects of the ground pattern of Cephalocarida, which is a pre-requisite for any comparisons in a broader perspective of crustacean phylogeny. Among the new findings or conclusions are: (1) Lightiella is in need of a revision since several assumed differences between the species are questionable or subject to intra-specific variability; (2) the cuticle of the trunk-limb basipod is sub-divided into a number of smaller sclerotized areas as in various exceptionally 3D preserved fossil crustaceans from Cambrian ‘Orsten’ faunal assemblages; (3) a small transitional portion on the post-maxillulary limbs in the area where the endopod and basipod connect is discussed as either a reduced, proximal endopod segment or as an evolutionary new joint of the basipod to enhance its flexibility; (4) the so-called pseud-epipod is interpreted as an outer branch of the exopod; (5) compared to ‘Orsten’ crustaceans many characters of the Cephalocarida are more modified than previously assumed, including the morphology of the trunk-limb basipod, and the unique, ring-shaped appearance of the abdominal segments. Also the development is not as plesiomorphic as sometimes assumed, at least not compared to that of the strictly anamorphic series of the ‘Orsten’ eucrustacean Rehbachiella kinnekullensis. The application of SEM techniques has again proved to be especially appropriate because of the small size of these animals, and because it permits direct comparisons with other similarly small crustaceans and the ‘Orsten’ crustaceans and their larvae.     

Chemosensory neurons in the mouthparts of the spiny lobsters Panulirus argus and Panulirus interruptus (Crustacea: Decapoda)

We studied electrophysiological properties of single chemosensory neurons in the mouthparts of the spiny lobsters Panulirus argus and Panulirus interruptus to complement our growing understanding of the behavioral roles of mouthparts of decapod crustaceans. Food mixtures and 13 single compounds were used to characterize the response specificity, sensitivity, and time course of individual neurons in the endopods of maxilliped 2 and 3. Additional chemoreceptors were found in the mandibular palp and basis of maxilliped 1 but they were not characterized. Neurons were broadly tuned, with the five most potent single compounds being ammonium, adenosine-5′-monophosphate, taurine, glutamate, and aspartate. Cluster analysis indicated that the neurons constitute a heterogeneous population that could be placed into seven groups linked according to their most excitatory compound. These neurons in the mouthparts had concentration-dependent responses, with thresholds between 10⁻⁷ and 10⁻⁴ M and without saturation even at 10⁻³ or 10⁻² M. They also quickly adapted when exposed to their best compounds at 10⁻⁴ and 10⁻³ M. A comparison of the response properties of these neurons in the mouthparts with those of chemosensory neurons in other crustacean appendages shows that neurons in the mouthparts have relatively broad tuning biased toward detecting and resolving high concentrations. Based on these comparisons, we suggest a functional distinction among the chemosensors on the different appendages: long distance detection by the antennae, precise location and collection by the pereiopods, and detailed assessment of quality by the mouthparts.     

Functional morphology of the feeding apparatus of ten species of Pacific butterflyfishes (Perciformes,Chaetodontidae): an ecomorphological approach

Synopsis The functional morphology of the feeding apparatus and the feeding ecology of an assemblage of ten species of butterflyfishes was investigated using a comparative ecomorphological approach. Behavioral observations in situ and in vivo, morphological measurements of fresh-killed specimens, scanning electron microscopy, and kinematic modeling were utilized. The fishes show varying degrees of morphological and behavioral specialization and generalization. The more specialized species group according to how they feed, rather than what they feed on. The feeding guild concept is therefore not very instructive in understanding the functional significance of the feeding apparatus. Many of the morphologically specialized butterflyfishes demonstrate evolutionary convergence in feeding morphology. Whereas the more morphologically specialized species do generally have more ecologically specialized diets, these data do not particularly support the ecomorphological hypothesis in that similar diets do not correspond to similar morphologies.     

The functional morphology of the feeding appendages and gut of Hippolyte varians (Crustacea: Natantia)

The musculature of the feeding appendages are described and compared with those few early accounts of other Decapoda, particularly those of Pandalus danae and Astacus fluviatilis . While the musculature is very similar to that of the latter, the relative importance of the various muscles may differ markedly and a number of previously undescribed muscles have been found, while others present in Pandulus and Astacus are not present in Hippolyte . The alimentary canal is also described, together with its associated musculature. The function of the various appendages and the parts of the gut are described from observations of living specimens.     

Comparative morphology of the feeding basket of five species of Euphausia (Crustacea,Euphausiacea) in the western North Pacific,with some ecological considerations

The feeding basket morphology and stomach content analyses of five Euphausia species (E. recurva, E. nana, E. pacifica, E. mutica and E. similis) were compared to elucidate their feeding ecology. The filter areas of feeding basket of E. pacifica and E. nana were proportionally larger than those of others in the size classes 13-20 mm and <13 mm, respectively, suggesting a high filtering efficiency in these species at each size class. Based on the secondary setal distance, it is suggested that there are three types of feeding basket in five Euphausia species. One is the fine mesh (<5 μm) of E. pacifica and E. nana; a second is the medium (10-20 μm) of E. recurva and E. mutica; and the third is the coarse (20-30 μm) of E. similis. The ability to feed on particles <5 μm would give both species, E. pacifica and E. nana, a great advantage over other species when number of flagellates is high. Stomach content analyses indicated a more omnivorous feeding mode in E. mutica, E. recurva and E. similis and a more herbivorous in E. pacifica and E. nana. This is in agreement with morphological studies of feeding baskets. Increments in the primary and secondary setal distances with increasing size of four Euphausia species, except E. pacifica, were evident, reflecting interspecific differences in food particle sizes utilized. In E. pacifica, however, the morphological similarity should produce intraspecific competition in diet. This revised version was published online in September 2006 with corrections to the Cover Date.     

Observations on the feeding mechanism,structure of the gut,and digestive physiology of the european lobster Homarus gammarus (L.) (Decapoda: Nephropidae)

Investigations have been made on the feeding mechanism, structure of the gut, and digestive physiology of the European lobster Homarus gammarus (L.).Ciné-photography has shown that the mandibles do not possess a masticatory function, merely serving to grip food morsels during the tearing process effected by the pulling action of the third maxillipeds. The remaining maxillipeds, together with the maxillae, then direct food fragments to the mouth for ingestion.Ingestion is facilitated by mucoid secretions discharged from the oesophageal tegumental glands; the glands do not appear to produce any enzymes which directly contribute to the digestive processes.The hepatopancreas is the principal organ concerned with digestion. It possesses a complex tubular organization in which sequential cellular differentiation culminates in the discharge of enzymes from the B-cells for extracellular digestion in the cardiac stomach. The enzymes are synthesized within vacuoles contained in the B-cell precursors (F-cells) and are secreted in three bursts of activity at 0–15 min, 1–2 h, and 3.5–5 h after a meal. The initial secretory phase is holocrine. Extracellular digestion involves esterases, arylamidases, and lipases; endopeptidases have not been positively identified by histochemical means despite the fact that Homarus is a carnivore. There is an intracellular digestive phase, not previously described in decapod crustaceans, at the 7–9 h post-ingestive stage in the hepatopancreatic R-cells which is effected by arylamidases and lipases.Various phosphatase enzymes have been identified in the hepatopancreatic cells. Acid and alkaline phosphatases are apparently concerned with several stages in the digestive cycle, including enzyme synthesis and secretion, and the absorption of digestive products. Adenosine triphosphatase activity is primarily associated with granules located in the distal R-cell cytoplasm; the possible significance of these granules in the elimination of metabolic wastes is discussed. Acid phosphatases and esterases are present in the midgut epithelium. The possibility of a passive uptake of material from the midgut lumen is considered.Faecal material in the hindgut is bound by mucoid secretions derived from the tegumental glands of this alimentary region; the mucus may also assist in defaecation.A complete digestive cycle in Homarus occupies ≈ 12 h.Food reserves in the gut consist principally of fat deposits in the R-cells, but minute amounts of glycogen can also be detected.No evidence of calcium, copper or ferric iron deposition in any part of the alimentary tract was found.     

Effects of jaw adductor hypertrophy on buccal expansions during feeding of air breathing catfishes (Teleostei,Clariidae)

Some species of Clariidae (air breathing catfishes) have extremely well developed (hypertrophied) jaw closing muscles that increase the maximal biting force of these species. As these enlarged jaw muscles tightly cover the suspensoria, which are firmly connected to the neurocranium, we expect diminished lateral expansions during suction for species with hypertrophied jaw muscles. In turn, this could imply a reduced suction performance for these species. Compared to Clarias gariepinus, which has relatively small jaw closers, Clariallabes longicauda shows a clear hypertrophy of the jaw adductors. A kinematic analysis of prey capture in these two species is presented here. As predicted, Clariallabes longicauda shows less lateral expansion (average abduction of the hyoids of 19.0°) than Clarias gariepinus (abduction of 31.1°). However, our data indicate that the decrease in lateral expansion capacity in the species with excessive adductor development is compensated for by a larger and faster ventral expansion of the buccal cavity by depression of the hyoid.     

Feeding behaviour, digestive physiology and lipid content in macropterous females of Pyrrhocoris apterus (L.) (Heteroptera: Pyrrhocoridae)

Abstract. Macropterous females of Pyrrhocoris apterus (L.) reared under short-day conditions (LD 12:12 h) were analysed for temporal patterns of feeding and drinking behaviour, activities of digestive enzymes in the gut, and lipid and glycogen content in the haemolymph and fat body. Peaks of drinking activity were recorded at days 3, 7 and 10 during the first 14 days after imaginal ecdysis. Feeding activity peaked on the third day, ceasing completely after the fourth day of adult life. Esterase, protease, amylase and aminopeptidase activities exhibited the highest overall activity in the first days after imaginal emergence; then enzyme activities decreased. In the fat body, the content of lipids was highest on day 5, then a decrease of about 40% was observed at day 14; the amount of glycogen was highest on day 1 at 11 μg of glucose equivalents/mg of fat body, then decreased to 2 μg at day 14 after the imaginal moult. In the haemolymph, the lipid content rose until day 8 when it reached almost 0.3 μmol/μl; at day 14 the value was slightly lower. The association of fasting with reproductive arrest in macropterous females of P. apterus, accompanied by a decrease in digestive enzyme activities and a mobilization of lipid reserves from the fat body, was demonstrated.     

Developmental changes in the structure and function of mouthparts of phyllosoma larvae of the packhorse lobster, Jasus verreauxi (Decapoda: Palinuridae)

Mouthpart morphology and feeding behaviour of Jasus verreauxi phyllosomas (instars 1-13) were examined using scanning electron microscopy and video analysis, respectively, to better understand ingestion and processing mechanisms and to identify developmental changes in feeding biology. The density, robustness and complexity of mouthpart setation increased with development, oral field increased and there were a greater number of spinose projections on maxillae 1 in mid and late instar phyllosomas. The second and third maxillipeds were able to sweep a larger area due to their increased length, which effectively increased the size of the oral field. Changes in feeding behaviour were consistent with these morphological differences between instars. In late instars, the shredding and tearing efficiency of maxillae 1 increased, larger pieces of prey were pushed between the mandibles, and the mandibles were able to effectively grind food due to a slight rotation away from the transverse plane. Both morphological and behavioural observations suggests that the absolute size range of prey increases with phyllosoma body size and the prey processing (i.e. ability to capture, manipulate and grind prey) becomes more efficient with development. We suggest early instar phyllosoma are most suited to a diet comprising softer prey items, whereas later-instar phyllosoma are better equipped to deal with larger, fleshier prey.     

Rapid radiation and cryptic speciation in squat lobsters of the genus Munida (Crustacea, Decapoda) and related genera in the South West Pacific: molecular and morphological evidence

Squat lobsters (genus Munida and related genera) are among the most diverse taxa of western Pacific crustaceans, though several features of their biology and phylogenetic relationships are unknown. This paper reports an extensive phylogenetic analysis based on mitochondrial DNA sequences (cytochrome c oxidase subunit I and 16S rRNA) and the morphology of 72 species of 12 genera of western Pacific squat lobsters. Our phylogenetic reconstruction using molecular data supports the recent taxonomic splitting of the genus Munida into several genera. Excluding one species (M. callista), the monophyly of the genus Munida was supported by Bayesian analysis of the molecular data. Three moderately diverse genera (Onconida, Paramunida, and Raymunida) also appeared monophyletic, both according to morphological and molecular data, always with high support. However, other genera (Crosnierita and Agononida) seem to be para- or polyphyletic. Three new cryptic species were identified in the course of this study. It would appear that the evolution of this group was marked by rapid speciation and stasis, or certain constraints, in its morphological evolution.     

Larval stages of the deep-sea lobster Polycheles typhlops (Decapoda,Polychelida) identified by DNA analysis: morphology,systematic, distribution and ecology

A total of 25 specimens of Eryoneicus larvae were collected near the Balearic Archipelago (Western Mediterranean Sea) in 2009 and 2010. Detailed morphological examination indicated that the smallest individual corresponded with the first zoea (ZI) stage of Polycheles typhlops hatched from a berried female by Guerao and Abelló (J Nat Hist 30(8):1179–1184, 1996). Only two species of deep-sea polychelid lobster, namely P. typhlops and Stereomastis sculpta, are known to occur in the Mediterranean. Genetic distance comparisons and phylogenetic analysis of the mitochondrial 16S rDNA and Cox I genes of this early larva together with adults from several Polycheles and Stereomastis species allowed us to assign it to P. typhlops. This is the first wild-caught larval stage of a polychelid lobster being identified using molecular techniques. The remaining specimens were attributed to zoeal stages II–III and decapodid stage based on morphological comparison. The arrangement of spines along the anterior part of the middorsal line (R, 1, 1, 1, 2, C₁), characteristic of the former species E. puritanii, discriminates these larvae from other Eryoneicus found in the Mediterranean. The clear presence of epipods on the third maxilliped and pereiopods of the decapodid stage gives further support to the identification of E. puritanii as the larval stages of P. typhlops. Additionally, information on the ecology of these larvae, their abundances during different seasons, as well as their bathymetric distribution is reported.     

The ontogeny of feeding behavior and cranial morphology in the leopard shark Triakis semifasciata (Girard 1854): A longitudinal perspective

While much is known about the ecological and functional morphology of feeding in lower vertebrates, studies of elasmobranch feeding over ontogeny are broadly lacking. In this study, the ontogeny of feeding behavior and morphology were investigated in neonatal and young-of-the-year leopard sharks Triakis semifasciata using morphometric measurements of growth and high-speed videography in a longitudinal study. Five food types were used during filming sessions to facilitate differentiation of modulation and variation over ontogeny. Functional aspects of muscle and buccal volume scaling were investigated through dissection. Growth over ontogeny was shown to influence numerous kinematic variables, while intra- and inter-individual variability was the dominant factor affecting feeding behavior. Modulation of feeding behavior based on food size and elusiveness was present for timing variables and predator motion during the strike, but not for food motion or the relative extent of buccal expansion. Allometric growth occurred in all aspects of external cranial morphology measured, resulting in a shallower head profile, anterior displacement of the mouth, and relatively larger jaw musculature over ontogeny. While the degree to which morphology constrains or enhances behavior cannot be directly quantified, variability in behavior greatly exceeds variability in morphology over early ontogeny. Maintenance of a behaviorally and morphologically versatile feeding apparatus throughout ontogeny is proposed to enhance the exploitation of resources and facilitate a diverse diet in T. semifasciata under variable environmental conditions.     

Functional morphology of the head of Hawaiian and Mid-Pacific butterflyfishes (Perciformes,Chaetodontidae)

Synopsis The functional morphology of the jaws of six species of butterflyfishes was investigated and related to their feeding behaviors. Utilizing observations and measurements of fresh-killed specimens as well as scanning electron microscopy of their dentitions, interspecific differences in the size, shape, orientation of the mouth, dentition, degree and function of jaw protrusion were related to their different feeding behaviors. The jaws of the six species showed a variety of adaptations for feeding on or over the reef as well as a repertoire of modulated jaw movements hitherto unnoticed in many studies. This radiation in morphologies was believed to be due to relatively simple changes of a few structural elements, changes that could occur relatively rapidly in the evolutionary time scale. The benefit of naturalistic observations in such functional morphological studies was emphasized.