Re‐establishment of biting mouthparts in desert‐living dung beetles (Scarabaeidae: Scarabaeinae) feeding on plant litter—Old structures reacquired or new ones evolved?

Evolution of mouthparts in adult dung beetles (Scarabaeidae: Scarabaeinae) for eating moist, fresh dung was linked with a loss of the ability to chew. However, the desert‐living genus Pachysoma, probably evolved from a wet‐dung feeding, Scarabaeus‐like ancestor, has switched to a diet of dry fecal pellets (of rodents or small ruminants) and plant litter that requires re‐establishment of chewing. Indeed, gut contents of a litter‐feeding Pachysoma species indicate efficient food comminution. Based on scanning electron microscopy, cutting and grinding mouthpart structures in six Pachysoma species, of two lineages and with different food preferences, are described and compared with homologous structures in wet‐dung feeding Scarabaeus species. In Pachysoma, cutting and breaking of large food items is probably performed by a clypeal scraper, a prominent epipharyngeal tooth and large maxillary galeal hooks. Further comminution is achieved by a large, grinding area evolved on the mandibular molae. Interspecific differences and the probable function and evolution of these structures are discussed. Particularly, the unique tools for cutting/breaking are completely novel structures and not results of some reacquisition of normal biting mouthparts. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Comparative morphology of the tentorium and hypopharyngeal–premental sclerites in sporophagous and non‐sporophagous adult Aleocharinae (Coleoptera: Staphylinidae)

We elucidate the configuration of the tentorium and the sclerites of the hypopharynx–prementum complex in selected spore‐ (pollen‐) and non‐spore‐feeding Aleocharinae (Staphylinidae) by presenting the first comparative 3D reconstructions of these structures for 19 staphylinoid beetle species (six outgroups, 13 Aleocharinae). General organization of the tentorium follows the groundplan previously proposed for adult Staphylinidae, although some taxa have reduced or lost the dorsal (all Aleocharinae studied, Agathidium mandibulare [Leiodidae]) or anterior (Omalium rivulare [Omaliinae], Anotylus sculpturatus [Oxytelinae]) tentorial arms. All species investigated have premental and hypopharyngeal sclerites that are partly homologizable across taxa. We clarified that Musculus praementopalpalis externus originates from the premental sclerite, resolving its unclear origin reported in our previous publications. Eight of 13 investigated Aleocharinae species are spore/pollen feeders, six obligatorily. Three of these six (Eumicrota, Gyrophaena fasciata, G. gentilis) have grinding pseudomolae and a fully developed hypopharyngeal suspensorium with posterior bridge and anterior elongations; the remaining three (Oxypoda, Pagla, Polylobus) lack pseudomolae and suspensorial bridge, but have the suspensorium elongated anteriorly. The dorsolateral side of the hypopharyngeal sclerite interacts with the pseudomola. Obligate sporophagy/pollinivory apparently arose at least three times in Aleocharinae, not always involving the pseudomola–hypopharynx grinding mechanism.     

Utility of geometric morphometrics for inferring feeding habit from mouthpart morphology in insects: tests with larval Carabidae (Insecta: Coleoptera)

Feeding habits are important life‐history traits in animals; however, methods for their determination are not well established in many species. The larvae of the beetle family Carabidae are an example. The present study tested the utility of geometric morphometrics of mouthpart morphology to infer the feeding habits of carabid larvae. Using Pterostichus thunbergi as a model system, larval feeding habits were inferred using geometric morphometrics of mouthparts and the results were compared with those obtained from rearing experiments. The rearing experiments indicated that P. thunbergi larvae are carnivores that require snails as an essential part of the diet. Through geometric morphometrics, associations between mouthpart morphology and larval feeding habits were confirmed for species in which these two traits are known. A discriminant analysis using these associations classified P. thunbergi larvae as snail/slug feeders, which is a result compatible with the rearing experiments. Geometric morphometrics also revealed that morphological integration and ontogenetic shape change might play roles in the diversification of mouthpart morphology. Overall, these results demonstrate the utility of the geometric morphometrics of mouthparts to infer feeding habit and to clarify the mechanisms of mouthpart morphological diversification in the study group, and the results also serve as a basis for future studies of other insect groups.     

Acanthodiaptomus denticornis another omnivorous calanoid copepod: description of its mouth appendages and feeding experiments on animal prey

The mouthpart morphology of the freshwater calanoid copepod Acanthodiaptomus denticornis was examined with optical microscopy. The mouthparts have sharp teeth and stout appendages with clawlike setae, typical of omnivorous calanoid mouthpart morphology. Observation of the buccal aperture with Scanning Electron Microscopy shows a large opening permitting feeding on prey as large as Keratella cochlearis. These observations agree with our feeding experiments which show that A. denticornis feed on K. cochlearis.     

Cleansing mechanisms of caprellid amphipods (Crustacea) from North America

There are three cleaning mechanisms utilized by intact caprellids separable by the body region being cleansed: antennae, mouthparts, and the remainder of the body. Antennal cleansing is a function of the first gnathopods and maxill'ipeds, mouthpart cleaning utilizes all of the mouthparts, and body cleansing relies solely on the first gnathopods. Behaviorally, filter feeders groom and feed approximately as much as predators, but filter feeders cleanse their entire body whereas predators primarily cleanse only both pair of antenna. The possibility that filter feeding was evolutionarily derived from cleaning activities is proposed.     

The mouth apparatus of Lithodes maja (Crustacea: Decapoda) – form,function and biological role

Being able to utilize many different food resources is probably an important aspect of the success of decapod crustaceans which fill a wide range of various ecological niches worldwide. The phenomenon is facilitated by the complex mouth apparatus found in this group, whose representatives possess six pairs of mouthparts– mandibles, maxillae 1, and maxillae 2 and three maxillipeds, the first three pairs of thoracic appendages which are also specialized to food manipulation. These six pairs are able to perform a number of movements for transporting, aligning, crushing and cutting. Studies into the functional morphology of mouthparts have already been carried out in some decapod species. This study focuses on Lithodes maja, a species of the hitherto understudied king crabs (Lithodidae), chosen on the grounds of their remarkable evolutionary history as ‘derived hermit crabs’. Individuals were filmed while being presented with different kinds of food. To obtain structural information on the individual mouthpart elements as naturally arranged in relation to one another, the shape of the mouthparts was 3D‐reconstructed from micro‐CT scans. These data were complemented by scanning electron microscopy, to analyse the surface structures in detail. There is evidence that the various elements of the mouthparts of L. maja can be sorted into six functional groups: (i) transporting mouthpart elements, (ii) aligning/sorting mouthpart elements, (iii) clutching/holding mouthpart elements, (iv) tearing/cutting/crushing mouthpart elements, (v) current‐generating mouthpart elements and (vi) grooming mouthpart elements. According to our 3D reconstruction, there only seem to be minor differences in morphology and relative position between the mouthparts of L. maja and those of the closely related species, Pagurus bernhardus.     

Mouthparts of flower-visiting insects

This review deals with the morphology and function of adult insect mouthparts modified to feed on nectar, pollen or petals. Specialization to nectar-feeding is evident in formation of proboscides of various lengths and designs. Proboscides of many Hymenoptera and Diptera function according to adhesion mechanisms that load nectar onto extensible apical mouthpart regions before fluid is conveyed along the food canal to the mouth by capillarity and suction. Predominantly suctorial proboscides evolved once in Lepidoptera, probably twice in Coleoptera, variously in some Hymenoptera and several times with similar design in Diptera. Many of them are particularly long and have sealed food tubes, specialized apical regions, new proboscis resting positions and modified feeding movements. Mouthparts of obligate pollen-feeding insects can be characterized by modified mandibles, specialized bristles for pollen manipulation and elaborate feeding movements. Often saliva is crucial for pollen retention and ingestion. In Coleoptera, intact pollen is gathered by sweeping movements of mouthparts; in Diptera, it is suspended in saliva prior to suction. Pollen is crushed by asymmetrical mandibles in aglossatan Lepidoptera and one group of basal Hymenoptera. Pollen-piercing mouthparts occur in Thysanoptera and one group of Diptera. Some butterflies and few Diptera extract nutrients from pollen by mixing it externally with saliva on their mouthparts. No mouthpart specializations to petal-feeding are reported in flower-visiting insects.     

Functional morphology of the feeding apparatus and evolution of proboscis length in metalmark butterflies (Lepidoptera: Riodinidae)

An assessment of the anatomical costs of extremely long proboscid mouthparts can contribute to the understanding of the evolution of form and function in the context of insect feeding behaviour. An integrative analysis of expenses relating to an exceptionally long proboscis in butterflies includes all organs involved in fluid feeding, such as the proboscis plus its musculature, sensilla, and food canal, as well as organs for proboscis movements and the suction pump for fluid uptake. In the present study, we report a morphometric comparison of derived long‐tongued (proboscis approximately twice as long as the body) and short‐tongued Riodinidae (proboscis half as long as the body), which reveals the non‐linear scaling relationships of an extremely long proboscis. We found no elongation of the tip region, low numbers of proboscis sensilla, short sensilla styloconica, and no increase of galeal musculature in relation to galeal volume, but a larger food canal, as well as larger head musculature in relation to the head capsule. The results indicate the relatively low extra expense on the proboscis musculature and sensilla equipment but significant anatomical costs, such as reinforced haemolymph and suction pump musculature, as well as thick cuticular proboscis walls, which are functionally related to feeding performance in species possessing an extremely long proboscis. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 291–304.     

Structural mouthpart interaction evolved already in the earliest lineages of insects

In butterflies, bees, flies and true bugs specific mouthparts are in close contact or even fused to enable piercing, sucking or sponging of particular food sources. The common phenomenon behind these mouthpart types is a complex composed of several consecutive mouthparts which structurally interact during food uptake. The single mouthparts are thus only functional in conjunction with other adjacent mouthparts, which is fundamentally different to biting–chewing. It is, however, unclear when structural mouthpart interaction (SMI) evolved since this principle obviously occurred multiple times independently in several extant and extinct winged insect groups. Here, we report a new type of SMI in two of the earliest wingless hexapod lineages—Diplura and Collembola. We found that the mandible and maxilla interact with each other via an articulatory stud at the dorsal side of the maxillary stipes, and they are furthermore supported by structures of the hypopharynx and head capsule. These interactions are crucial stabilizing elements during food uptake. The presence of SMI in these ancestrally wingless insects, and its absence in those crustacean groups probably ancestral to insects, indicates that SMI is a groundplan apomorphy of insects. Our results thus contradict the currently established view of insect mouthpart evolution that biting–chewing mouthparts without any form of SMI are the ancestral configuration. Furthermore, SMIs occur in the earliest insects in a high anatomical variety. SMIs in stemgroup representatives of insects may have triggered efficient exploitation and fast adaptation to new terrestrial food sources much earlier than previously supposed.     

Profound head modifications in Claviger testaceus (Pselaphinae,Staphylinidae, Coleoptera) facilitate integration into communities of ants

Clavigeritae is a group of obligate myrmecophiles of the rove beetle subfamily Pselaphinae (Staphylinidae). Some are blind and wingless, and all are believed to depend on ant hosts through feeding by trophallaxis. Phylogenetic hypotheses suggest that their ancestors, as are most pselaphines today, were free-living predators. Morphological alterations required to transform such beetles into extreme myrmecophiles were poorly understood. By studying the cephalic morphology of Claviger testaceus, we demonstrate that profound changes in all mouthpart components took place during this process, with a highly unusual connection of the maxillae to the hypopharynx, and formation of a uniquely transformed labium with a vestigial prementum. The primary sensory function of the modified maxillary and labial palps is reduced, and the ventral mouthparts transformed into a licking/‘sponging’ device. Many muscles have been reduced, in relation to the coleopteran groundplan or other staphylinoids. The head capsule contains voluminous glands whose appeasement secretions are crucial for the beetle survival in ant colonies. The brain, in turn, has been shifted into the neck region. The prepharyngeal dilator is composed of an entire series of bundles. However, the pharynx does not show any peculiar adaptations to taking up liquid food. We demonstrate that far-reaching cephalic modifications characterize C. testaceus, and that the development of appeasement glands and adaptation of the mouthparts to trophallaxis determine the head architecture of this extreme myrmecophile.     

Mouthparts of larvae of Opifex fuscus and Aedes australis (Diptera: Gulicidae); a scanning electron microscope study

Abstract

Larvae of the New Zealand culicid species Opifex fuscus and Aedes australis have previously been reported to show dimorphism in the structure of their labral brushes, some larvae having pectinate bristles and others only simple hairs. In the scanning electron microscope all larvae showed some degree of pectination of hairs in these brushes. There is also a gradation in the pectination. Some bristles are only sparsely pectinate; because the dimensions of their teeth are close to the limit of resolution by the compound microscope, the pectination had previously gone undetected. The mouthparts of both species are intermediate in character between those typical of filter-feeding larvae and those typical of browsing larvae. The SEM appearance of maxillary sensoria and bristles on the ventral surfaces of the mandibular brushes is described; the latter bristles comb food particles out of the labral brushes and towards the mouth. Features of the mouthparts are illustrated with scanning electron micrographs.     

Mouthparts of the bark beetle (Ips acuminatus) as a possible carrier of pathogenic microorganisms

The bark beetle Ips acuminatus Gyllenhal (Coleoptera: Curculionidae: Scolytinae) has been recently reported as one of the most serious secondary pests of pine trees. Since the adult beetles chew the sapwood to create tunnels, they have developed effective drilling mouthparts enough to make galleries directly into the heartwood of the tree. The mouthparts of this bark beetle exhibit typical morphology of mycophagous coleopteran beetles and have those characteristics of chewing mouthparts. Both maxillary and labial palpi have the functions of directing the food to the mouth and holding it while the mandibles chew the food. Although this bark beetle did not have prothoracic mycangial cavities, yeast-like spores were concentrated at the invaginated surface of mouthparts where cuticular hairs are densely packed. In particular, the cuticular surface around the base segments of these palpi has sufficient spaces to accommodate microorganisms during the series of drilling or feeding processes. Therefore, this paper reports detailed observation of the cuticular structure of the mouthpart using the field emission scanning electron microscopy (FESEM) for the purpose of demonstrating its possible implication to act as external carriers of pathogenic microorganisms.     

Morphological and trophic specialization in a subterranean amphipod assemblage

相似文献   

The Structure of the Mouthparts of Some Galypterate Dipteran Larvae in Relation to Their Feeding Habits

The anatomy of the head and the structure of the mouthparts is described for calypterate larvae of differing feeding habits. The way in which the skeletal parts of the cephalopharynx and the musculature are adapted to the type of food is discussed and particular reference is made to the action of the mouthparts in the carnivorous larva of Limnophora riparia Fallen.     

Comparative morphology of the head of selected sporophagous and non‐sporophagous aleocharinae (Coleoptera: Staphylinidae): Musculature and hypopharynx‐prementum complex

To investigate whether specialization to spore‐ (or pollen‐) feeding in advanced Aleocharinae is mirrored by their head anatomy, we compiled and compared synchrotron X‐ray micro‐tomography datasets for 11 Aleocharinae in conjunction with previous data for two aleocharine and six outgroup species (two nonstaphylinids, four staphylinids). We describe the presence/absence of head muscles and investigate the variability of points of origin by character mapping analyses. Monophyly of Aleocharinae is supported by the absence of M. 48 (M. tentoriobuccalis anterior), and by changes in the origins of Mm. 1, 2, 17, 18, 28, 29, 30. Within Aleocharinae the origins of the labial muscles (Mm. 28–30) have shifted posteriorly to the gula, which might enhance the movement posterad of the hypopharynx and partly compensate for the loss of M. 48. We also analyzed the general organization of the hypopharynx‐prementum complex and the fine structure of the mandibles through SEM studies. In the absence of grinding mandibular molae like those of most mycophagous Coleoptera, seven aleocharine species studied have evolved “pseudomolae” at the ventral side of the mandibles that replace true molae as secondary grinding surfaces. In these species, the hypopharynx is elevated and displaced anteriorly, bearing a bowl‐like depression on its surface that functions as a mortar where spores are ground between the hypopharynx and the mandibles. Two of these species are not yet known to feed on spores or pollen. Another species (Oxypoda alternans) is thought to feed on fungus material but bears no pseudomolae on its mandibles. J. Morphol. 271:910–931, 2010. © 2010 Wiley‐Liss, Inc.     

Conservation and diversification of gene function during mouthpart development in Onthophagus beetles

The evolutionary success of insects is in part attributable to the tremendous diversification of their mouthparts, which permitted insects to radiate into novel food niches. The developmental genetic basis of mouthpart development has been well studied in at least two insect taxa possessing derived mouthparts, the hemipteran Oncopeltus fasciatus and Drosophila. However, much less is known about the regulation of mouthpart differentiation of the presumed ancestral mandibulate type. Here we aim to extend current insights into the patterning of mandibulate mouthparts through a functional genetic analysis of three leg gap genes, homothorax (hth), dachshund (dac), and Distal-less (Dll), in the dung beetle Onthophagus taurus, a species whose mouthpart arrangement has in part retained, as well as diverged form, the ancestral mandibulate mouthpart type. We specifically include in this study a first functional genetic analysis of the adult labrum, an enigmatic mouthpart whose appendicular origin has been the subject of a long-standing debate. Our results support a functional role of all three patterning genes in the development of the labium, maxilla, as well as the labrum. In contrast, mandible development appeared to rely only on the patterning functions of hth and dac, but not Dll. Here, our results raise the possibility that evolutionary changes in the dac-patterning may have played an important role in the evolutionary transition from a short, triangular mandible adapted for chewing to the elongated, flat, and blade-like mandible of modern filter-feeding scarabaeine beetles. In general, our results contribute to a growing body of studies that suggest that basic patterning genes can contribute to morphological evolution of adult features while maintaining traditional patterning responsibilities at earlier developmental stages or in other body regions.     

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.     

Drinking with a very long proboscis: Functional morphology of orchid bee mouthparts (Euglossini,Apidae, Hymenoptera)

Neotropical orchid bees (Euglossini) possess the longest proboscides among bees. In this study, we compared the feeding behavior and functional morphology of mouthparts in two similarly large-sized species of Euglossa that differ greatly in proboscis length. Feeding observations and experiments conducted under semi-natural conditions were combined with micro-morphological examination using LM, SEM and micro CT techniques. The morphometric comparison showed that only the components of the mouthparts that form the food tube differ in length, while the proximal components, which are responsible for proboscis movements, are similar in size. This study represents the first documentation of lapping behaviour in Euglossini. We demonstrate that Euglossa bees use a lapping-sucking mode of feeding to take up small amounts of fluid, and a purely suctorial technique for larger fluid quantities. The mouthpart movements are largely similar to that in other long-tongued bees, except that the postmentum in Euglossa can be extended, greatly enhancing the protraction of the glossa. This results in a maximal functional length that is about 50% longer than the length of the food canal composing parts of the proboscis. The nectar uptake and the sensory equipment of the proboscis are discussed in context to flower probing.