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.     

Revising the definition of the crustacean seta and setal classification systems based on examinations of the mouthpart setae of seven species of decapods

The crustacean cuticle has numerous projections and some of these projections, the setae, have important mechanical as well as sensory functions. The setae display a wide diversity in their external morphology, which has led to great problems separating setae from other projections in the cuticle and problems in making a consistent classification system. Here, the cuticular projections on the mouthparts of seven species of decapods are examined by scanning and transmission electron microscopy. A new definition is given: a seta is an elongate projection with a more or less circular base and a continuous lumen; the lumen has a semicircular arrangement of sheath cells basally. From the details of the external morphology the mouthpart setae are divided into seven types: pappose, plumose, serrulate, serrate, papposerrate, simple and cuspidate setae, which are suggested to reflect mechanical functions and not evolutionary history. This classification system is compared with earlier systems.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 233–252.     

Mechanical functions of setae from the mouth apparatus of seven species of decapod crustaceans

The mouthpart setae of seven species of decapods were examined with macro-video recordings and scanning electron microscopy. The general mechanical (nonsensory) functions of the different mouthparts are described and an account of their setation is given. This offers the possibility to determine the mechanical functions of the different types of setae. Pappose setae do not participate in food handling but in general make setal barriers. Plumose setae likewise do not contact food objects but assist in current generation. Papposerrate setae are rare but they were seen to assist in pushing food particles into the mouth. Serrulate setae are very common and mainly participate in gentle food handling and grooming. Serrate setae are used for more rough food manipulation and grooming. The roughest shredding, tearing, and manipulation of prey items are handled by the cuspidate setae. Simple setae seem to be divided into two populations with very different functions. On the maxillipeds of Panulirus argus they are used for shredding, tearing, and holding the food objects, but on the basis of maxilla 2 of three other species they appear to have very little mechanical influence and only when handling small prey items. The functional scheme seems to be consistent within the Decapoda.     

Surface structures of the antennular flagella of the hermit crab Pagurus alaskensis (Benedict): A light and scanning electron microscopy study

The surface structures of the antennular flagella of Pagurus alaskensis are described in detail. Attention is directed towards the surface morphology of two types of possible sensilla: (1) exoskeletal pores (1.0–3.0 μm in diameter); (2) setae of various kinds. In addition, small (0.1–0.2 μm) pits occur in the exoskeleton which are not considered to be sensory in function. The exoskeletal pores are found at fairly specific locations on both the inner and outer flagella, particularly on the short segments of the outer flagella. Neither the inner nor the outer flagella are bilaterally symmetrical with respect to their setal armature. On the outer flagellum six groups of setae may be distinguished: lateralmesial; dorsal; ventral; accessory; aesthetasc; setae of the distal segment. On the inner flagellum setae of the mesial and lateral rows form distinctive groups. The morphology, orientation and locations of all the flagellar setae are defined and where possible the numbers of the various morphological types within the specific setal groups are given. It is noteworthy that many setal types have obvious apical pores and yet no pores could be found in the chemoreceptive aesthetasc setae. The functions of the various setae are discussed in relation to their topographical position and to existing electrophysiological and behavioral data. Some suggestions are made about future experiments to demonstrate the central connections of specific sensilla or groups of sensilla and to show their significance in the whole animal.     

Displacement sensors in the honeybee pollen basket

The mechanical importance of different groups of corbicular setae to the formation and maintenance of the pollen load was established through ablation experiments. The neurophysiological characteristics of corbicular setae were investigated and it was found that there are three types of response to mechanical displacement of individual setae. At least some of the anterior corbicular setae behave as linear displacement sensors and are capable of measuring the size of the forming pollen load during foraging.     

Morphology of the head of <Emphasis Type="Italic">Sericoderus lateralis</Emphasis> (Coleoptera,Corylophidae) with comments on the effects of miniaturization

The mouthparts and musculature of the head of Sericoderus lateralis are described. It is shown that the mouthparts of minute beetles preserve their complex structure and are not significantly affected by miniaturization. Such stability allows these features to be used in macrosystematics of groups with variable body size.     

Setal morphology of grooming appendages in the spider crab,Libinia dubia

In crustaceans, grooming behaviors decrease fouling by removing debris from the exoskeleton and body structures; these grooming behaviors improve respiration, sensory reception, movement, and reproduction. Setal morphologies of the following grooming appendages in the decapod crustacean spider crab Libinia dubia are examined including the first pereiopod (cheliped), first, second, and third maxillipeds (mouthparts), and first, second, and third epipods (internal extensions of the maxillipeds). The objective of this study was to describe setal morphologies of these grooming appendages and to elucidate possible functions and efficiencies of setal structures. Spider crabs are hypothesized to have elaborate setal morphologies, mainly for cleaning specialized decorating setae as well as for cleaning inside the gill chamber, which has a higher likelihood of becoming fouled compared to other decapods such as shrimps. Fourteen setal types are documented and included several varieties of serrate and pappose setae as well as simple setae, cuspidate setae, papposerrate setae, and canoe setae. Maxillipodal epipods in the gill chamber are free of fouling, suggesting the setation on the third maxilliped protopod has an efficient functional morphology in removing debris before water enters the gill chamber. Serrate setae may function for detangling and separating structures whereas pappose setae may function for fine detailed grooming. The cheliped is the only grooming appendage that can reach decorating setae and it contains only pappose setae; thus decorating setae is not likely groomed in a manner that would greatly decrease fouling. J. Morphol. 277:1045–1061, 2016. © 2016 Wiley Periodicals, Inc.     

A new species of Caprella (Crustacea: Amphipoda) from the Mediterranean Sea

A new caprellid amphipod, Caprella tavolarensis n. sp., is described based on specimens collected from a Posidonia oceanica seagrass meadow at the Tavolara-Punta Coda Cavallo Marine Protected Area (Sardinia, Mediterranean Sea). The species is close to Caprella liparotensis, but can be clearly distinguished by smaller size, presence of a short rostrum, body elongate and dorsally smooth, absence of serrate carina on the basis of gnathopod 2 and pereopods, mouthparts scarcely setose, absence of fine setae on peduncle of antenna 1 and absence of swimming setae on antenna 2. The number of caprellid species reported from the Mediterranean Sea has increased from 23 (1993) to 41 (2010), consequently, further taxonomical studies should be addressed to properly estimate the total amphipod diversity in the Mediterranean Sea.     

Feeding modes of deep-sea lobsters (Crustacea: Decapoda: Nephropidae and Palinuridae) in Northwest Pacific waters: Functional morphology of mouthparts, feeding behaviour and gut content analysis

The setation of the mouthparts, gut contents and video recordings of live individuals of the deep-sea clawed lobsters (Nephropidae) Metanephrops formosanus, M. armatus and the spiny lobster (Palinuridae) Puerulus angulatus from northwest Pacific waters were analysed to get an insight into their feeding modes. A comparison of SEM photos shows a high degree of similarity between the morphology and setation of the mouthparts of M. formosanus and M. armatus, but that of P. angulatus was very different to Metanephrops. Serrate setae are most abundant on the feeding appendages of M. formosanus and M. armatus. The mouthparts of P. angulatus are dominated by simple and cuspidate setae. Gut contents of Metanephrops spp. contained small crustacean parts, fish and bivalves and a considerable amount of sediment (∼60% relative abundance). Guts of Puerulus contained mostly small pieces of fish and crustaceans and only a relatively minor amount of sediment (<10%). Video analysis revealed that the studied Metanephrops species are able to handle soft food items by cutting and abrading movements of the mouthparts. Puerulus would not feed on presented food items under lab conditions. The feeding appendages and their setation are clearly related to the feeding modes of the species studied. Both Metanephrops species have slender appendages with fine and sharp setae, suggesting it is a predator and/or scavenger on small crustaceans and ingest deposits to a limited extent. Puerulus angulatus has thick and shorter appendages with strong simple and cuspidate setae, possibly corresponding to a more predatory lifestyle.     

Functional morphology of the feeding apparatus of the nymph of Farrodes sp. (Ephemeroptera: Leptophlebiidae)

Farrodes nymphs are specialized periphyton/biofilm scrapers. Their maxillae are the most specialized mouthparts, but other elements, with their systems of variously modified setae, are designed to obtain and transport food particles to the pharynx with a minimum of loss. The morphology and adaptations of these mouthparts and related head areas, as well as associated musculature, are described.     

Functional morphology of mouthparts and digestive system during larval development of the cleaner shrimp Lysmata amboinensis (de Man, 1888)

Mouthpart and alimentary canal development was examined in Lysmata amboinensis larvae using scanning electron microscopy and histology. The gross morphological features of external mouthparts and internal digestive tract structures of larvae at different developmental stages indicate that ingestive and digestive capabilities are well developed from early on. With increasing age of the larvae the mouthpart appendages increased in size, the hepatopancreas in tubular density and the midgut in length. The density of setae and robustness of teeth and spines of individual structures increased. The most pronounced changes from early to late stage larvae involved formation of pores on the paragnaths and labrum, transformation of the mandibular spine‐like teeth to molar cusps, development of the filter press in the proventriculus and of infoldings in the previously straight hindgut. The results suggest that early stage L. amboinensis larvae may benefit from soft, perhaps gelatinous prey, whereas later stages are better equipped to handle larger, muscular or more fibrous foods. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

The setae of Eurydice pulchra (Crustacea: Isopoda)

The setal armature of Eurydice pulchra Leach is described using an ordinary optical microscope. On the basis of size, setae are divided into macrotrichs and microtrichs. Macrotrichs are further classified according to structural similarities into four groups. Only two types of microtrichs have been recognized. The distribution of the different setae on both adults and juveniles is described and where possible an attempt is made to relate morphology and distribution to function.  

Summary:

The setal armature of E. pulchra has been divided into macrotrichs, varying in length from 0·025 to 0·4 mm, and microtrichs varying in length from 2 to 10 μm. The macrotrichs have been classified into groups based on structural similarities and 17 distinct types have been recognized. Each differ in length, degree of chitinization and modification of the shaft wall. The distribution of the macrotrichs has been described and particular types have been shown to occur in distinct positions on the body. Many of the macrotrichs are peculiar to one position on the body.
Two types of microtrichs have been described, a single type approximately 10 μm in length and a smaller type 2 μm in length. The latter occurs in short crescentic rows principally on the pleopods and oostegites. All setal types occurring in the adult have been shown to be present in the newly emerged juvenile although the number developed is very much less. Similar setal types are present in males and females. An attempt has been made to relate the morphology and distribution of both macrotrichs and microtrichs to possible functions.     

Structure and sensilla of the antennae and mouthparts of Loxocephala perpunctata Jacobi (Hemiptera: Fulgomorpha: Eurybrachidae)

The ultramorphology of the antennae and mouthparts of the adult Loxocephala perpunctata Jacobi was studied through a scanning electron microscope. Seven types of sensilla were found on antennomeres, including a Böhm bristle on the scape, sensillum trichoideum and plaque organ on the pedicel, two subtypes of sensilla chaetica and two subtypes of sensilla campaniformia on these two antennomeres; and Bourgoin's organ with sensory pegs and sensilla basiconicum on the basal bulb of the flagellum. The mouthparts of L. perpunctata are of the typical piercing-sucking type, similar to mouthparts found in other hemipteran insects. In general, six types of sensilla (i.e., four subtypes of sensilla chaetica, sensillum basiconicum, subapical labial sensillum, uniporous peg-like sensillum, multiporous peg-like sensillum and two subtypes of bristle-like sensilla) were detected on different locations of the labium, with the last three, and numerous cuticular processes, present on the labial tip. The potential functions of these sensilla are discussed.     

Mechanosensory neurons with bend- and osmo-sensitivity in mouthpart setae from the spiny lobster Panulirus argus

The mouthparts of the spiny lobster Panulirus argus hold primarily two types of setae--simple setae and cuspidate setae. Mechanosensory neurons from these setae were examined by electrophysiological recordings. The population of simple setae contained two types of mechanosensory neurons: displacement-sensitive neurons, which responded to deflection at the setal base; and bend-sensitive neurons, which responded to bending of the setal shaft. Displacement-sensitive neurons, in general, responded phasically and only during actual displacement. Typically, their response changed with alteration of the direction, amplitude, and velocity/acceleration of the mechanical stimulus. Bend-sensitive neurons, in general, responded phaso-tonically and carried information on the direction and region of bending. This is the first experimental demonstration of bend sensitivity for arthropod setae. Cuspidate setae contain highly sensitive mechanosensory neurons; however, due to the rigid nature of these setae, whether they were bend sensitive or displacement sensitive could not be determined, and they were thus called "tactile neurons." Bend-sensitive neurons, but not displacement-sensitive neurons or tactile neurons, showed graded responses to changes in osmolarity. The osmosensitivity of these neurons could mediate behavioral responses to changes in the osmolarity of seawater or food.     

Trophic strategies and functional morphology of feeding appendages, with emphasis on setae, of Upogebia omissa and Pomatogebia operculata (Decapoda: Thalassinidea: Upogebiidae)

Although species of Upogebiidae historically have been considered filter feeders, recent studies show that many species of this group also deposit feed. In this study, the degrees of trophic specialization of two species of this family, Upogebia omissa and Pomatogebia operculata , were analysed. Feeding mechanisms, stomach contents and morphology of the feeding appendages, with emphasis on setae, were examined. U. omissa , found in sandy substrate, is a generalistic feeder while P. operculata inhabits burrows inside corals, being more specialized in filter feeding. Only 21% of the 57 setal types described were common to both species. Setal types were clustered in three main categories: plumed, serrate and plumodenticulate. No simple setae were found. P. operculata has lower setal diversity, with higher ratio of plumodenticulate to serrate setal types than U. omissa. The 1st and 2nd pereiopods have an important role in collecting food. The mouthparts have two main functions: to brush and retain particles so that food can be transported from the pereiopods to the mouth. Generally in these appendages, the dactyli and basal endites are responsible for brushing particles and the meri and coxal endites for particle retention. The diversity of setal types and complexity of their distribution on the appendages may be related to the necessity to select and triturate particles prior to ingestion, reflecting the differences in trophic strategies utilized by U. omissa and P. operculata. Setal characters appear to be indicators of the relative importance of a specific feeding mode for species of this group.     

Morphological diversity of setae on the grooming legs in Anomala (Decapoda: Reptantia) revealed by scanning electron microscopy

The fifth pereiopods (P5) in Anomala are specialized appendages used mainly for grooming. We studied the articulated cuticular outgrowths, setae, on the distal segments of the P5 in 40 species from 18 Anomala families using light and scanning electron microscopy. Five general classes of setae can be found on the P5: serrate, serrulate and simple setae which all appear bristle-like, and tooth-like and scale-like cuspidate setae. We classified the bristle-like setae according to criteria of shape and the arrangement of distinct outgrowths – denticles and setules – on the shaft of the seta. In this way we were able to distinguish eleven mainly serrate and serrulate types of seta. Some setal types imply homology due to their distinctness and could thus help to solve problematic phylogenetic questions. One setal type, for example, is only present in pagurid hermit crabs and king crabs, which corroborates the theory that these two morphologically very dissimilar groups are, in fact, closely related.     

Fine structure and sensory apparatus of the mouthparts of the pear psyllid,Cacopsylla chinensis (Yang et Li) (Hemiptera: Psyllidae)

The pear psyllid, Cacopsylla chinensis (Yang et Li) (Hemiptera: Psyllidae), is one of the most significant economic pests of pear in China, causing direct damage through feeding by the highly specialized piercing–sucking mouthparts. The ultrastructural morphology and sensory apparatus of the mouthparts of the adult were examined using scanning and transmission electron microscopy. The piercing–sucking mouthparts of C. chinensis are composed of a three-segmented labium with a deep groove in the anterior side, a stylet fascicle consisting of two mandibular and two maxillary stylets, and a pyramid-shaped labrum. Proximal to the labium, the stylet fascicle forms a large loop within a membranous crumena. Mandibles, with more than ten teeth on the external convex region, can be seen on the distal extremity. Smooth maxillary stylets are interlocked to form a larger food canal and a smaller salivary canal. One dendritic canal housing 2 dendrites is also found in each mandible. Two types of sensilla trichodea, four types of sensilla basiconica, single as well as groups of sensilla campaniformia, and oval flattened sensilla occur in different locations on the labium, whereas a kind of sensilla basiconica is at the junction of the labrum and anteclypeus. Sensilla trichodea and sensilla campaniformia, always present with denticles, are present on the middle labial segment. Three types of sensilla basiconica, two types of sensilla trichodea and two oval flattened sensilla are located on the distal labial segment. The mouthpart morphology and abundance of sensilla located on the labium in C. chinensis are illustrated, along with a brief discussion of their taxonomic and putative functional significance.