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.     

The setae of Austropotamobius pallipes (Crustacea: Astacidae)

The author presents here the results of an introductory study into the setal armature of the British species of crayfish, Austropotamobius pallipes (Lereboullet, 1858). Using an ordinary optical microscope, it is demonstrated that a considerable variety of setae exist on the body and appendages of A. pallipes. These setae are described, and their distribution indicated. The setae fall into two main groups:
(1) Setae with relatively thick walls and inconspicuous ampulla, and having no basal septum.
(2) Those setae with relatively thin walls, a well-developed ampulla and basal septum. Further subdivisions of these two groups are made on the bases of outgrowths and outlines of the setal wall. Finally, the significance of the distribution and the variation, is discussed in relation to crayfish behaviour and function, linking the observed morphological facts with the results of some current electro-physiological experiments performed on related decapods.     

Morphology and distribution of setae on the antennules of the Caribbean spiny lobster Panulirus argus reveal new types of bimodal chemo-mechanosensilla

This study describes the morphology and distribution of setae on the lateral and medial flagella of the antennules of the spiny lobster Panulirus argus in an effort to identify antennular chemoreceptors in addition to the well-studied aesthetasc chemosensilla. Setae were examined using light and electron microscopy, and their distribution on flagellar annuli was analyzed. We identified ten setal types based on external morphology: hooded, plumose, short setuled, long simple, medium simple, short simple, aesthetasc, guard, companion, and asymmetric setae, with the last four types being unique to the "tuft" located on the distal half of the lateral flagellum. The three setal types whose ultrastructure was examined--hooded, long simple, and medium simple setae--had characteristics of bimodal (chemo-mechanoreceptive) sensilla. The antennules have four distinct annular types based on their setal complement, as shown by cluster analysis. This basic distribution of non-tuft setal types is similar for both lateral and medial flagella. Annuli in the tuft region have tuft setal types superimposed on a basic organization of non-tuft setal types. These results show that the antennules possess a diverse set of setae, that these setae have a highly ordered arrangement on the antennules, that at least four (and probably many more) of these setal types are chemosensilla, and suggest that most antennular chemosensilla are bimodally sensitive.     

Ultrastructure and development of forked and capillary setae in the polychaetes Orbinia bioreti and Orbinia latreillii (Annelida: Orbiniidae)

Abstract. Recent investigations into chaetogenesis of certain types of annelid setae provide important results for unravelling the phylogenetic relationships within several taxa of poly-chaetous annelids. This paper presents data on ultrastructure and development of 2 types of orbiniid setae. The analysis of the crenulate capillaries in Orbinia latreillii reveals a formation process which clearly differs from the development of Equisetum -like setae of lingulid bra-chiopods. For the investigation of forked setae, which up to now have been neglected in the discussion on the phylogenetic significance of annelid setae, notopodial setal sacs of O. latreillii and O. bioreti were studied by light- and electron microscopy. In the setal sacs, stages of forked setae are restricted to a dorsocaudal pouch, which represents the site of setal formation. The 2 diverging, stout tines of the fork bear spines on their inner margins, each of which is preformed by a single microvillus. After retraction of the microvilli, a characteristic pattern of the setal canals inside of the spines remains. The present study belongs to a series of comparative studies into chaetogenesis of forked setae. These special setae are also found in other orbiniid taxa as well as some paraonids, scalibregmatids, and nephtyids. Ultrastructural investigations into the development of these forked setae might suggest homology.     

Setal morphology of the grooming appendages of Macrobrachium rosenbergii (Crustacea: Decapoda: Caridea: Palaemonidae) and review of decapod setal classification

Setae are vital in grooming activities and aiding in the removal of epibionts and sedimentary fouling from the body surfaces of decapod crustaceans. Thus, the setal structures and their arrangement on the grooming appendages and sensory structures of the commercially important shrimp, Macrobrachium rosenbergii, were examined using scanning electron microscopy. Macrobrachium rosenbergii is extensively grown in aquaculture and exhibits unique male morphological forms, termed morphotypes. The three male morphotypes are termed blue‐clawed males, orange‐clawed males, and small‐clawed or undifferentiated males and all three differ in their dominance, behavior, body morphology, and reproductive success. Seven setal types, two of which have never been described in the literature, are identified on the grooming appendages (third maxillipeds, first, second, and fifth pereopods) and antennae: simple, serrate, serrulate, spiniform, pappose, crinoid, and spinulate. The latter two setae are newly identified. Certain setal types, such as serrate and serrulate setae were located and associated with specific grooming appendages such as the first pereopods. The types of setae on the grooming appendages varied among females and male morphotypes and the novel setal types (crinoid and spinulate) were found only on two of the male morphotypes. A literature review of terminology related to the structure of setae and setal types in decapod crustaceans is offered as the usage of various terms is ambiguous and conflicting in the literature. The intention of this review is to provide future authors with a comprehensive collection of terms and images that can be used to describe various aspects of setal morphology in decapods. J. Morphol. 275:634–649, 2014. © 2014 Wiley Periodicals, Inc.     

Morphology and distribution of antennular setae of scyllarid lobsters (Scyllarides aequinoctialis, S. latus, and S. nodifer) with comments on their possible function

Abstract. Lateral flagella of the antennules of scyllarid lobsters were examined for setal morphology and distribution via scanning electron microscopy. Setal distribution patterns were mapped directly for 3 regions of the antennule ( base, tuft , and tip ) and analyzed for differences: (1) between left and right antennules, (2) between males and females within a species, and (3) among species by comparing counts of setae per annulus in the ventral tuft region only. Six types of antennular setae were identified based on their external morphology: aesthetases, simple, modified simple, asymmetric, hemi-plumose, and toothbrush setae. These different types were organized in a clear pattern over the ventral and dorsal surfaces of the lateral flagella of the antennule. Aesthetase, asymmetric, modified simple, and hemi-plumose setae were found only on annuli in the tuft region between the distal and proximal ends of the flagellum. Simple setae were found on all annuli of all regions of the antennule, and toothbrush setae were mainly concentrated on all annuli of the base region and on proximal annuli of the tuft region . All species of scyllarids examined had the same general pattern of setal distribution and no differences were found between left and right, or male and female antennules. Similar setae located on the lateral antennules of species from the families Nephrophidae and Palinuridae (clawed and spiny lobsters) have been previously described as chemo- and/or mechanoreceptive for use in distance chemoreception (i.e., detection and orientation to olfactory stimuli). Based on work on clawed and spiny lobsters, we predict that the aesthetases on slipper lobsters have a chemoreceptive function and that simple and toothbrush setae may have a bimodal chemo- and mechanoreceptive function.     

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.     

The structure and growth of the statocyst in the Australian crayfish Cherax destructor

The morphology of the statocyst of the Australian crayfish Cherax destructor was examined using scanning electron microscopy. It resembles in general structure, size, and position the statocysts of crayfish described previously, and the size and distribution of the fields of setae on the floor of the capsule are similar but not the same. Over the size range examined, the relationship between the carapace length, the length of the basal antennular segment, the diameter of the statocyst capsule, and the total number of setae are all linear. The number and position of setae on the floor of the statocyst capsule were mapped for animals in two size classes (small, ca. 20 mm; large, ca. 50 mm) to test for changes in their arrangement during growth. The change in the ratio of setal number to statocyst size between the two size classes was about three times greater for the anterior setal field than for the other fields. We propose that differential development of the setal fields may be related to changes in the force-monitoring requirements of the animals as they increase in size, but this remains to be experimentally tested.     

Ultrastructural Similarities Between Setae of Brachiopods and Polychaetes1

Larvae of the brachiopod Terebratalia transversa Sowerby have 2 bundles of setae on each side of their mantle lobe. Each seta grows out of a follicle of epidermal cells. The cisternae and associated vesicles of the Golgi bodies in lateral follicular cells are filled with electron dense material. The basal follicular cell, or chaetoblast, has apical microvilli projecting into the basal ends of the longitudinal channels of the seta. The channel partitions are composed of fine fibers oriented parallel to the setal axis. The close structural and developmental similarities between polychaete and brachiopod setae are discussed.     

The setal morphology of the larval phases of the Argentinean red shrimp Pleoticus muelleri Bate, 1888 (Decapoda: Solenoceridae)

Summary

The setal morphology is described for the larval stages of the solenocerid shrimp Pleoticus muelleri based upon scanning electron microscope studies. Sixty-five different types of setae are recognized and some currently used terms for describing setae are reviewed.     

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.     

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.     

A scanning electron microscope study of tarsal adhesive setae in the Coleoptera

Scanning electron micrographs of the tarsal adhesive setae of 84 species of beetle are described. These show a vast range of setal structure and distribution.     

Structure of dactyl sensilla in the kelp crab,Pugettia producta

In the kelp crab, Pugettia producta, flat plate setae cover all but the ventral surfaces of the walking leg dactyls. Dendrites enter the setal shaft located inside the plate superstructure, and extend to a region of the setal tip that contains a system of minute pores resembling the pore systems found in chemosensory sensilla of insects. Presumably, much of the chemosensitivity of the dactyls in the kelp crab is mediated by the plate setae. In the interior of the dactyl, supporting cells and the neurons innervating plate setae, other types of setae, and other presumptive sensilla form scolopidia. Large scolopidia, containing as many as 12 dendrites, appear to innervate some of the plate setae and also large ventral rodlike setae that might be chemosensory. Two of the dendrites of large scolopidia usually have more densely packed microtubules, longer ciliary axonemes, slightly larger rootlets, and dark A fibers with arms, characteristics indicative of mechanosensory function. Some dactyl setae, therefore, could be both mechanosensory and chemosensory. Small scolopidia containing two or three dendrites that exhibit mechanosensory characteristics appear to innervate small, rodlike setae, which presumably are strictly mechanosensory. The two types of structures located on the epicuticular cap, elliptical structures resembling campaniform sensilla and small cones in pits resembling CAP organs, appear to be dually innervated and presumably are mechanosensory, although other functions are possible. The internal positions of the scolopidia, together with the support afforded by an extracellular dendritic sheath, by the scolopale, and by desmosomelike and septate junctions, may serve to protect internal portions of setal dendrites, some of which appear to remain functional in nonmolting adults that have abraded setae.     

Prey finding behavior and mechanosensilla of larval Toxorhynchites brevipalpis Theobald (Diptera : Culicidae)

Behavioral experiments demonstrated that starved 3rd-instar Toxorhynchites brevipalpis (Diptera : Culicidae) will attack a glass probe in response to vibrations alone. Frequencies in the range of 80–200 Hz elicited 85% of the attacks. The observation that most attacks occurred while the larva was drifting forward towards the probe, substantiates that the predatory behavior of these organisms is basically opportunistic.The main setae on the thorax and abdomen arise from setal support plates and are of 4 general morphological types: dendritic, smooth, spinulate spiniform, and unbranched aciculate setae. The numbers and lengths of the setae on 3rd- and 4th-instar larvae are given. Each seta is innervated by a bipolar neuron with a tubular body in the dendrite, a characteristic of mechanosensilla. By virtue of their abundance and location, it seems probable that one or more types of these setae sense the water vibrations capable of eliciting an attack response.The tubular bodies of the 4 types of setae are somewhat unusual in that they lack electron-dense material, except near sites of attachment of the microtubules to the dendritic membrane, yet possess a large number of complexly arranged microtubules. Among the setal types, variations were observed in the prominence of microtubular attachment sites, proportion of tubules associated with the sites, and orientation of microtubules. In spinulate spiniform setae, the arrangement of microtubules varies within the same tubular body; peripherally most of the microtubules are in one direction with little convergence and have only one distinct attachment site, whereas medially they converge considerably resulting in 2 groups, each associated with its own well-defined site.The attachment sites, which presumably through linkage with the dendritic membrane provide a means of initiating depolarization, are associated with the distal ends of almost all of the microtubules. This suggests that in these tubular bodies depolarization may be initiated through mechanical force acting along the length of the microtubules, that is, stretching and/or compression.     

The modality of the dominance signal in snapping shrimp (Alpheus heterochaelis) and the corresponding setal types on the antennules

Previous behavioural experiments showed that snapping shrimp lacking lateral antennular filaments, i.e. without chemosensory aesthetascs, lose the ability to distinguish between conspecifics that are inexperienced in fighting and former winners. A chemosensory dominance signal was assumed to be present, although other receptors unique to the lateral filaments may have been responsible for the behavioural changes. In the present study, the antennules of snapping shrimp were examined for differences between the lateral and medial antennule filaments to identify the modality of the dominance signal. We found six different types of setae and two types of pores. A new probably bimodal setal type is described, the broad long simple seta. Only the chemosensory aesthetascs and their associated hydrosensory companion setae are unique to the lateral filament. Thus we conclude that the dominance signal is chemical, because a hydrodynamic signal would be also received by the simple setae distributed on both filaments.     

The modality of the dominance signal in snapping shrimp (Alpheus heterochaelis) and the corresponding setal types on the antennules

Previous behavioural experiments showed that snapping shrimp lacking lateral antennular filaments, i.e. without chemosensory aesthetascs, lose the ability to distinguish between conspecifics that are inexperienced in fighting and former winners. A chemosensory dominance signal was assumed to be present, although other receptors unique to the lateral filaments may have been responsible for the behavioural changes. In the present study, the antennules of snapping shrimp were examined for differences between the lateral and medial antennule filaments to identify the modality of the dominance signal. We found six different types of setae and two types of pores. A new probably bimodal setal type is described, the broad long simple seta. Only the chemosensory aesthetascs and their associated hydrosensory companion setae are unique to the lateral filament. Thus we conclude that the dominance signal is chemical, because a hydrodynamic signal would be also received by the simple setae distributed on both filaments.     

A new species of Anuretes Heller, 1865 (Copepoda: Caligidae) from the yellowbanded sweetlips Plectorhinchus lineatus (Haemulidae) off New Caledonia

A new species of caligid copepod, Anuretes justinei n. sp., is described from off New Caledonia. It is parasitic on the gill filaments of a haemulid fish, the yellowbanded sweetlips Plectorhinchus lineatus (Linnaeus). The new species is distinguished from its congeners by the combination of the following character states: (1) the fourth pedigerous somite is covered dorsally by the expanded free posterior margin of the cephalothorax; (2) a maxillary whip is present; (3) the relatively small genital complex is less than half the length of the cephalothorax; (4) leg 3 is armed with nine setae on the terminal exopodal segment and six setae on the terminal endopodal segment; and (5) leg 4 is long and slender with a setal armature of I, III twisted spines. The new species is an addition to the possibly monophyletic group of seven species that is characterised by the possession of a maxillary whip, all of which are found on haemulid hosts. The host-specificity of Anuretes is relatively high, its species being largely parasitic on reef-associated fishes, such as the families Haemulidae (eight species), Ephippidae (four species), Acanthuridae (four species) and Pomacanthidae (one species).     

The idiosomal chaetotaxy of astigmatid mites

Within the Astigmata, setal homologies with the general chaetotaxic systems developed for acariform mites by F. Grandjean have never been convincingly established. This study deals with all body regions, exclusive of legs and gnathosoma, mostly utilizing as models astigmatid species from three different families. Six hypotheses which attempt to explain the ontogeny of segments, setae and cupules in the caudal bend region are compared using three-dimensional views obtained by scanning electron microscopy. The hypotheses are evaluated by in-group comparisons testing their consistency with presumed segmental boundaries, and by out-group comparisons with oribatid mites. The strongest, most parsimonious hypothesis suggests that larval astigmatid mites possess segment F, but not its setae, and the two pairs posterior to segment F are h _l and h ₂, whilst the five setae added in the protonymph are h ₃, f ₂ and ps _1–3. Six pairs of structures on the deutonymphal anal plate are identified as modified setae, and evidence supporting their homologies with setae of other instars is discussed for the first time. A comparison of the principal chaetotaxic systems used for the dorsal and anal regions of the Astigmata is presented. Application of Grandjean's setal signatures to these regions, as well as to the coxisternal and genital regions, is discussed and illustrated by examples.