Analysis of the Anointing and Grooming Behavior of Several Adult Insects in Typhlocybinae (Hemiptera: Cicadellidae)

Anointing and grooming behaviors of the adult of Singapora shinshana (Matsumura) (Erythroneurini), Empoascanara (Empoascanara) mai Dworakowska (Erythroneurini), Austroasca vittata (Lethierry) (Empoascini), and Limassolla diospyri Chou et Ma (Zyginellini) were recorded and analysed. Through observation and analysis of their anointing and grooming, we found that there is a preparation stage before anointing behavior in typhlocybines. Anointing and grooming are repeated several times and the numbers and interval times differ from species to species among the Typhlocybinae. Ultramorphological examination of brochosomes and the morphology of the brochosome field, metathoracic tibiae and the structures on the head, face, mesothorax and pygofer of E. mai were done using a scanning electron microscope. Brochosomes (IBS) on the cuticle of typhlocybine adults were spherical visible above these integument. Characteristics of anointing and grooming behavior, the provide evidence that the brochosome field may function as a storage site in preparation for the next grooming session.     

Grooming as a secondary behavior in the shrimp Macrobrachium rosenbergii (Crustacea,Decapoda, Caridea)

The giant freshwater prawn, Macrobrachium rosenbergii, is a large shrimp extensively used in aquaculture whose grooming behaviors were analyzed in this study. Macrobrachium rosenbergii exhibits three unique male morphotypes that differ in their behavior, morphology and physiology: small-clawed males (SM), orange-clawed males (OC) and blue-clawed males (BC). The largest and most dominant males, BC males, are predicted to have significantly different grooming behaviors compared to females and the other two male morphotypes. These BC males may be too large and bulky to efficiently groom and may dedicate more time to mating and agonistic interactions than grooming behaviors. Observations were conducted to look at the prevalence of grooming behaviors in the absence and presence of conspecifics and to determine if any differences in grooming behavior exist among the sexes and male morphotypes. Significant differences in the grooming behaviors of all individuals (females and male morphotypes) were found. BC males tended to have the highest grooming time budget (percent of time spent grooming) while SM males had a relatively low grooming time budget. The grooming behaviors of the male morphotypes differed, indicating while these males play distinct, separate roles in the social hierarchy, they also have different grooming priorities. The conditions in which Macrobrachium rosenbergii are cultured may result in increased body fouling, which may vary, depending on the grooming efficiencies and priorities of these male morphotypes. Overall, grooming behaviors were found to be a secondary behavior which only occurred when primary behaviors such as mating, feeding or fighting were not present.     

Palatal and lingual adaptations for frugivory and nectarivory in the Wahlberg’s epauletted fruit bat (Epomophorus wahlbergi)

Wahlberg’s epauletted fruit bat (Epomophorus wahlbergi) feed on fleshy fruit and nectar of flowers and have an important role in pollination and seed dispersal. It was expected that their buccal morphological structures are adapted to this type of feeding. Consequently, buccal cavity and lingual structures of E. wahlbergi were examined by extended focus light microscopy (LM) and scanning electron microscopy (SEM). Morphology of the tongue of E. wahlbergi was similar to that of other fruit- and nectar-feeding bats. The elongated tongue of these bats possessed filiform and conical papillae as mechanical papillae and fungiform and circumvallate papillae as gustatory papillae that varied in distribution. Epomophorus wahlbergi had five palatal ridges and one post-dental palatal ridge, and relatively wide, flattened molar teeth. A hard, papillae structure at the posterior end of the upper palate of the upper plate, which has not been previously described, was observed. It appears that this structure works together with the palatal ridges and teeth, so that the bats crush fleshy fruits during feeding and extract the juices before discarding the pulp. Consequently, lingual and particularly palatal structures of E. wahlbergi show morphological adaptations for efficiently feeding on fleshy fruit and nectar.     

Two ancient bacterial endosymbionts have coevolved with the planthoppers (Insecta: Hemiptera: Fulgoroidea)

Background

Pseudoscorpions are chelicerates and have historically been viewed as being most closely related to solifuges, harvestmen, and scorpions. No mitochondrial genomes of pseudoscorpions have been published, but the mitochondrial genomes of some lineages of Chelicerata possess unusual features, including short rRNA genes and tRNA genes that lack sequence to encode arms of the canonical cloverleaf-shaped tRNA. Additionally, some chelicerates possess an atypical guanine-thymine nucleotide bias on the major coding strand of their mitochondrial genomes.

Results

We sequenced the mitochondrial genomes of two divergent taxa from the chelicerate order Pseudoscorpiones. We find that these genomes possess unusually short tRNA genes that do not encode cloverleaf-shaped tRNA structures. Indeed, in one genome, all 22 tRNA genes lack sequence to encode canonical cloverleaf structures. We also find that the large ribosomal RNA genes are substantially shorter than those of most arthropods. We inferred secondary structures of the LSU rRNAs from both pseudoscorpions, and find that they have lost multiple helices. Based on comparisons with the crystal structure of the bacterial ribosome, two of these helices were likely contact points with tRNA T-arms or D-arms as they pass through the ribosome during protein synthesis. The mitochondrial gene arrangements of both pseudoscorpions differ from the ancestral chelicerate gene arrangement. One genome is rearranged with respect to the location of protein-coding genes, the small rRNA gene, and at least 8 tRNA genes. The other genome contains 6 tRNA genes in novel locations. Most chelicerates with rearranged mitochondrial genes show a genome-wide reversal of the CA nucleotide bias typical for arthropods on their major coding strand, and instead possess a GT bias. Yet despite their extensive rearrangement, these pseudoscorpion mitochondrial genomes possess a CA bias on the major coding strand. Phylogenetic analyses of all 13 mitochondrial protein-coding gene sequences consistently yield trees that place pseudoscorpions as sister to acariform mites.

Conclusion

The well-supported phylogenetic placement of pseudoscorpions as sister to Acariformes differs from some previous analyses based on morphology. However, these two lineages share multiple molecular evolutionary traits, including substantial mitochondrial genome rearrangements, extensive nucleotide substitution, and loss of helices in their inferred tRNA and rRNA structures.     

Blood vascular architecture of the rat lingual papillae with special reference to their relations to the connective tissue papillae and surface structures: a light and scanning electron microscope study

Subepithelial blood vessels of the rat lingual papillae and their spatial relations to the connective tissue papillae and surface structures were demonstrated by light and scanning electron microscopy. In the rat, four types of papillae were distinguished on the dorsal surface of the tongue, i.e. the filiform, fungiform, foliate and circumvallate papillae. Vascular beds of various appearance were found in all four types of lingual papillae: a simple or twisted capillary loop in the filiform papilla; a basket- or petal-like network in the fungiform papilla; a ring-like network in the foliate papilla, and a conglomerated network surrounded by double heart-shaped capillary networks in the circumvallate papilla. These characteristic vascular beds corresponded to the shape of the connective tissue papillae and surface structures. The vascular bed beneath the gustatory epithelium in the fungiform, foliate and circumvallate papilla consisted of fine capillary networks next to the taste buds.     

Time-matched grooming in female primates? New analyses from two species

The parcelling model of reciprocity predicts that grooming partners will alternate between giving and receiving grooming within grooming bouts, and that each partner will perform approximately as much grooming as it receives within each bout (‘time matching’). Models of allogrooming based on biological markets theory predict that individuals of lower dominance rank will exchange grooming for tolerance from high-rankers, and therefore an inverse relation will be found between grooming partners' dominance rank distance and how closely they match each other's grooming contributions within each bout. We used weighted logistic regression and weighted least-squares regression to test these predictions using data from female white-faced capuchins, Cebus capucinus, and bonnet macaques, Macaca radiata. Only 5-7% of macaque grooming bouts, and 12-27% of capuchin grooming bouts, were reciprocated. However, (1) the duration of grooming by the first groomer significantly predicted whether the groomee would reciprocate at all, and (2) when bouts were reciprocated, the duration of grooming by the first groomer significantly predicted the duration of grooming by the second groomer. Grooming was most balanced among females of similar dominance ranks. Both the time-matching and rank-related effects were weak, although significant. These results indicate that although some form of time matching may be a general characteristic of grooming in female-bonded primate species, time matching accounts for relatively little of the variation in the distribution of grooming within bouts. We also draw attention to weighted regression as a technique that avoids pseudoreplication while using all available data.     

Seasonal aspects of the life cycle of solifuges (Arachnida, Solifugae) as compared with pseudoscorpions (Arachnida, Pseudoscorpiones)

Solifuges (order Solifugae) and pseudoscorpions (order Pseudoscorpiones) united into the superorder Haplocnemata (Shultz, 2007) are nevertheless characterized by essential differences both in morphological and biological characters. Analysis of available information on the biology and life cycles of these arachnids revealed a clear difference between the daily rhythms of activity: their presence in solifuges and their absence in pseudoscorpions. However, this concerning the seasonal adaptations in the two orders is not simple since they demonstrate not only differences but also a lot of similarities. All the studied solifuges are characterized by the seasonally timed stenochronous (heterodynamic) type of development which is characteristic of species with uni-, bi-, and semi-voltine development (i.e., to life cycles completed within a year, half a year, and several years), as well as to species combining different forms of voltinism. This type of development is not only prevalent in solifuges (as in pseudoscorpions and other arachnids) but appears to be the only one, since no cases of eurychronous (homodynamic) development have been found in solifuges; whereas pseudoscorpions and other arachnids develop both steno- and eurychronously. The initial ontogenetic stages remain in underground shelters (brood burrows in solifuges and brood chambers in pseudoscorpions). The first nymphal stages (I instar nymphs in solifuges, protonymphs in pseudoscorpions) are embryonized; active life outside the brood burrows starts with II instar nymphs in solifuges and with deutonymphs in pseudoscorpions.     

Cuticular structures of the priapulid Halicryptus spinulosus: A scanning electron microscopical study

Summary The surface anatomy and the structures lining the pharynx of Halicryptus spinulosus were examined by scanning electron microscopy (SEM). The structures were compared and contrasted with those reported for other priapulids, particularly those features previously studied with SEM. Buccal papillae and pharyngeal teeth of two types were described. Surface structures observed with SEM were: scalids, abdominal setae, anal papillae, posterior warts and ring papillae. The latter three structures are unique among described priapulids. The anal papillae are composed of several rounded, perhaps pedunculate, structures; the posterior warts are composed of mitriform structures in close association with columnar structures. Both are located in separate depressions in the posterior integument. The ring papillae occur on the annuli close to the posterior end. Halicryptus spinulosus was previously thought to lack these structures.     

Do exaggerated chelicerae function as weapons or genitalia in a long‐jawed spider? Functional allometric analysis yields an answer

From the elongated neck of the giraffe to the elaborate train of the peacock, extreme traits can result from natural or sexual selection (or both). The extreme chelicerae of the long‐jawed spiders (Tetragnatha) present a puzzle: do these exaggerated chelicerae function as weapons or genitalia? Bristowe first proposed that Tetragnatha chelicerae function as a holdfast because these spiders embrace chelicerae during mating. This hypothesis has remained untested until now. Here, we use functional allometry to examine how extreme chelicerae develop and perform in the long‐jawed spider Tetragnatha elongata. Similar to other Tetragnatha species, chelicerae were longer in adult males than in adult females. Overall, we confirm Bristowe's hypothesis: elongation only occurred in the adult stage. However, we propose that chelicerae function as more than a holdfast in T. elongata. Male chelicerae exhibited positive allometry, which suggests scaling as weapons rather than genitalia. However, fieldwork revealed that the operational sex ratio is female‐biased and both adult male–male competition and sexual cannibalism were rarely observed. Consequently, we propose that the positive allometry of male chelicerae may result from sexual selection to mechanically mesh with larger and more fecund females. Evidence for mechanical mesh includes multiple traits ranging from apophyses and grooves to guide teeth on the basal portion of the chelicerae. In contrast, we propose that chelicerae of females are analogous to the female peacock's tail: shortened by natural selection limiting the exaggeration of sexually selected traits. Indeed, females had increased foraging efficiency compared to males and exhibited negative cheliceral allometry. We discuss the implications for the evolution of elongated chelicerae in Tetragnatha.     

Light- and Scanning-Electron-Microscopic Analyses of the Plica glossoepiglottica lateralis of the neonate and adultPan troglodytes troglodytes — Blumenbach (1799)

A unique type of papillae is evident in the plicae glossoepiglotticae laterales of the tongue ofPan troglodytes troglodytes. Areas of occurrence show flat lobes of varying length with elongated miniature processes on their free tips. The function of these papillae is presumably the convection of liquid food into the pharynx (along with the lateral folds). Furthermore, it is assumed that the papillae are sensory organs which relay taste, temperature, pain, and pressure, similar to the papillae filiformes of the tongue of other mammalia (Kunze 1969). Free nerve fibres and nerve endings were found in both the epithelium and the connective-tissue. Nerve structures resembling the “Organs of Meissner” were also discovered in the subepithelial connective-tissue. Goblet cells are found in the surface layers in the epithelium of the papillae of the newborn. These are absent in the papillae of the adultPan troglodytes troglodytes. The secretion of the goblet cells functions as a mechanical and chemical protectant for the papillae.     

Are the morphometrics of chelicerae correlated with diet in mesostigmatid mites (Acari)?

Gamasid mites are prominent predators of soil communities. Some species are said to be specialized predators feeding on nematodes or arthropods. A conspicuous difference between species is the structure of the chelicerae. We test the hypothesis that measurable attributes of the chelicerae are related to diet. We measured ten morphometric parameters of the chelicerae from 52 species within 29 genera. For 28 species and 26 genera we were able to extract food preferences from the literature and own observations. We found some weak indications that arthropod predators have proportionally larger chelicerae compared to similar-sized species feeding on nematodes or other worm-like diets. The latter are characterized by a larger number of teeth along the digitus fixus and digitus mobilis.     

Morphology of sensory papillae on the feeding proboscis of cone snails (Mollusca,Gastropoda)

Cone snails, predatory marine gastropods, have developed a specialized prey capture method in which a long, distensible proboscis is used to identify prey and inject venom via a hydraulically propelled hollow radular tooth. Using brightfield, epifluorescence, confocal, and transmission and scanning electron microscopy, we describe the morphology of ciliated sensory structures concentrated on the tip of the proboscis. The number and morphology of these sensory papillae are linked to the type of preferred prey: cone snails feeding on worms and mollusks have short, cone or finger‐shaped papillae, whereas fish‐hunting cone snails have long tubular papillae in addition to short conical papillae. Sensory papillae are well positioned to provide information necessary to locate, identify, and dispatch prey. Proboscis tips and their sensory papillae regenerated within 10 d following experimental ablation, and snails with regenerated proboscis tips were able to locate and envenomate prey. The remarkable intrageneric variation found in the morphology of these sensory structures is probably linked to the specialized prey that cone snail species have evolved to hunt.     

Oviposition behaviour of the soil mite Veigaia cerva (Acari: Veigaiidae)

We observed the oviposition behaviour of the soil mite Veigaia cerva (Kramer) (Acari: Veigaiidae) using continuous video-monitoring. Five phases could be recognized. Phase I involved inspection of the substrate. In phase II the female rhythmically moved her gnathosoma and first pair of legs. After an inactive phase III, the soma was raised (IV), and the egg was laid (V). In the actual egg laying three sub-phases could be distinguished: internal egg movement, placing the egg in front of the gnathosoma, and depositing the egg using the chelicerae. The palps and first pair of legs were used to position the egg between the chelicerae. The whole process took on average 333 ± 22 s.     

All the better to see you with: eyes and claws reveal the evolution of divergent ecological roles in giant pterygotid eurypterids

Pterygotid eurypterids have traditionally been interpreted as active, high-level, visual predators; however, recent studies of the visual system and cheliceral morphology of the pterygotid Acutiramus contradict this interpretation. Here, we report similar analyses of the pterygotids Erettopterus, Jaekelopterus and Pterygotus, and the pterygotid sister taxon Slimonia. Representative species of all these genera have more acute vision than A. cummingsi. The visual systems of Jaekelopterus rhenaniae and Pterygotus anglicus are comparable to that of modern predatory arthropods. All species of Jaekelopterus and Pterygotus have robust crushing chelicerae, morphologically distinct from the weaker slicing chelicerae of Acutiramus. Vision in Erettopterus osiliensis and Slimonia acuminata is more acute than in Acutiramus cummingsi, but not to the same degree as in modern active predators, and the morphology of the chelicerae in these genera suggests a grasping function. The pterygotids evolved with a shift in ecology from generalized feeder to specialized predator. Pterygotid eurypterids share a characteristic morphology but, although some were top predators, their ecology differs radically between genera.     

Seasonal aspects of the life cycles of pseudoscorpions (Arachnida, Pseudoscorpiones)

Analysis of available information on the seasonal features of life cycles in pseudoscorpions (Arachnida, Pseudoscorpiones) shows that in the temperate climate of Europe with distinct seasonality, the development of these peculiar arachnids can be either eurychronous (= homodynamic), with a poorly pronounced seasonal arrangement, or stenochronous (= heterodynamic), with a distinct seasonal arrangement. This is similar to some other arachnids, particularly spiders and harvestmen. Eurychronous pseudoscorpions are characterized by the approximately year-round development (quite often with winter activity in the whole or part of the population), an overlapping of consecutive generations, the presence of alternative development with or without the dormant state (at all the postembryonic life cycle stages), lack of brood chamber production by females, and the start of free-living life at the protonymphal stage. Stenochronous pseudoscorpions, on the contrary, possess clearly arranged (usually univoltine) development with overwintering deutonymphs and tritonymphs (more rarely adults), a clear separation of adjacent generations, the production of brood chambers where the regressive protonymphs develop until their molt into deutonymphs, and the start of free-living life at the deutonymphal stage. These two types of life cycles are exemplified by two pseudoscorpions from South England, namely Neobisium muscorum with eurychronous development, and Chthonius orthodactylus with stenochronous, univoltine development (Goddard, 1976). There is no correlation between the taxonomic position and the seasonal characters of life cycles in representatives of Neobisiidae, Chthoniidae, and Chernetidae. There is, instead, a close connection between the eco-physiological state of these arachnids and the type of their gas exchange (continuous, diffusive and non-cyclic in the active state, as opposed to discontinuous, with cyclic emission of CO₂ and uptake of O₂ in the dormant state). The latter information may be useful for distinguishing different kinds of dormancy (diapause and quiescence) in these arachnids.     

Surface ultrastructure of the tegument of Clonorchis sinensis newly excysted juveniles and adult worms.

The tegumental structures of newly excysted juveniles and adult worms of Clonorchis sinensis were studied using scanning and transmission electron microscopy. After excystation the juvenile's tegumental surface is characterized by knoblike protuberances and is armed almost entirely with numerous rows of small spines encircling the body. These spines are double- or triple-pointed on the anterior portion of the body and become single-pointed posteriorly. Four types of presumed sensory structures were observed as follow: A) ciliated knoblike papillae and B) nonciliated platelike papillae, both of which are arranged in rougly a bilaterally symmetrical pattern dorsally, ventrally, and laterally; C) rounded swellings of nonciliated papillae on the lips of the ventral and oral suckers, which are characterized in the transmission electron microscope by a rounded dense body in the apical bulb; and D) a sensory receptor with a bulbous projection having the appearance of a modified cilium, which was not found with SEM likely owing to its being enclosed by an extension of the tegument. In full-grown adult worms, the tegumental surface is knobbed or lobulated in various forms without surface spines. The tegumental structures in the adults appear to be clearly differentiated from those in the juveniles. Upraised, buttonlike papillae, each topped by a short cilium, which are similar to the Type A papillae in the juveniles, are distributed thickly around the oral and ventral suckers, and are rather randomly scattered over the remainder of the body. Some nonciliated swollen papillae were found on the lip of the ventral sucker.     

The Absence of Grooming for Rank-Related Benefits in Female Assamese Macaques (Macaca assamensis)

Seyfarth’s model of social grooming proposes that by grooming females higher ranking than themselves, females can gain access to important rank-related benefits, such as agonistic support. This, in turn, produces a distinctive pattern of grooming in which females direct their grooming up the female dominance hierarchy and compete for access to the highest ranking individuals. We aimed to test to what extent the grooming behavior of female Assamese macaques (Macaca assamensis) fits the assumptions and predictions of Seyfarth’s model. During two 1-yr sampling periods (October 2007–September 2008, May 2010–April 2011) we collected >2100 focal hours of data from a single wild group in their natural habitat at Phu Khieo Wildlife Sanctuary, Thailand. Subjects included all adult female group members (N?=?12 in 2007/8; N?=?15 in 2010/11). We collected detailed data on grooming interactions, approaches, and departures as well as all aggressive and submissive behaviors between all subjects. We found no evidence that grooming was exchanged for rank-related benefits. In line with this we found no evidence that the grooming of female Assamese macaques fits the pattern predicted by Seyfarth’s model. These results are surprising given that such deviations from Seyfarth’s model are relatively rare among macaques. We propose that our findings are best explained as a lack of a need for rank-related benefits by females in this group.     

Fine structure and distribution of epidermal projections associated with taste buds on the oral papillae in some loricariid catfishes (Siluroidei: Loricariidae)

Two morphologically distinct structures occur on the surfaces of the oral papillae in several loricariid catfish species; namely, (1) typical vertebrate taste buds composed of receptor and sustentacular cells and (2) brushlike projections, termed epidermal brushes, that represent specialized epidermal cells containing keratin. Both of these structures were studied with the combined use of light microscopy and scanning and transmission electron microscopy. The general body surface, fins, and rostral cutaneous processes of some loricariid catfishes are covered with taste or terminal buds but lack the epidermal brushes. It is suggested that the epidermal brushes found on the oral papillae serve as protective devices for the taste buds and as abrasive surfaces for substrate scraping during feeding. The taste buds on the oral papillae may detect any gustatory stimuli from the resulting substrate disturbance. Comparative studies reveal many differences in the number and spatial arrangement of these two structures on the oral papillae among the several species of the Loricariidae examined. These differences may represent functional adaptations to the various modes of life in the Loricariidae.     

The influence of kinship and dominance hierarchy on grooming partner choice in free-ranging <Emphasis Type="Italic">Macaca mulatta brevicaudus</Emphasis>

In group-living animals, individuals do not interact uniformly with their conspecifics. Among primates, such heterogeneity in partner choice can be discerned from affiliative grooming patterns. While the preference for selecting close kin as grooming partners is ubiquitous across the primate order, the selection of higher-ranking non-kin individuals as grooming partners is less common. We studied a group of provisioned rhesus macaques (Macaca mulatta brevicaudus) on Hainan Island, China, to examine rank-related benefits of grooming exchanges and the influence of kin relationships. We tested four hypotheses based on Seyfarth’s model: (1) there will be kin preference in grooming relationships; (2) grooming between non-kin individuals will be directed up the dominance rank; (3) grooming between non-kin individuals will reduce aggression from higher-ranking ones; and (4) non-kin individuals will spend more time grooming with adjacent ranked ones. We found that grooming relationships between kin individuals were stronger than those between non-kin individuals. For non-kin relationships, lower-ranking individuals received less aggression from higher-ranking ones through grooming; a benefit they could not derive through grooming exchanges with individuals related by kinship. Individuals spent more time grooming adjacent higher-ranking non-kin individuals and higher-ranking individuals also received more grooming from non-kin individuals. Our results supported Seyfarth’s model for predicting partner choice between non-kin individuals. For relationships between kin individuals, we found results that were not consistent with prediction for the exchanges of aggression and grooming, indicating the importance to control for the influence of kinship in future studies.