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The genus Carabus (Coleoptera: Carabidae: Carabinae) is a group of predacious ground beetles found throughout Holarctic temperate zones. The diversification of this group is attributed not only to allopatric speciation but also to diversification in life-history traits and morphological traits that are involved in ecological adaptation and reproductive isolation. Precise phylogenetic reconstruction of Carabus is important to understanding the factors associated with species' divergence in relation to such character divergence. Here, we analyse the phylogeny of Carabus among higher taxonomic groups and among species of the subgenus Apotomopterus , a hyper-diverse group endemic to East Asia. We used genome-wide sequence data from ultraconserved elements for phylogenetic reconstruction via maximum-likelihood and species-tree reconstruction methods. With regard to the relationships among higher taxonomic groups, the obtained phylogenetic trees resolved two structures with previously unresolved divergence. The subgenus Apotomopterus contained three lineages, i.e. the Vitalisi, Prodigus, and Sauteri groups. It showed diversification of the spinula, a sclerotized male genital structure, and variation in the female elytral notch (preapical emargination). These morphological traits may be involved in copulatory behaviour and potentially affect reproductive isolation, and their character state divergence may have facilitated species divergence of Apotomopterus. [ABSTRACT FROM AUTHOR]

Abstract Background Cichlid fishes in Lake Tanganyika exhibit remarkable diversity in their feeding habits. Among them, seven species in the genus Perissodus are known for their unique feeding habit of scale eating with specialized feeding morphology and behaviour. Although the origin of the scale-eating habit has long been questioned, its evolutionary process is still unknown. In the present study, we conducted interspecific phylogenetic analyses for all nine known species in the tribe Perissodini (seven Perissodus and two Haplotaxodon species) using amplified fragment length polymorphism (AFLP) analyses of the nuclear DNA. On the basis of the resultant phylogenetic frameworks, the evolution of their feeding habits was traced using data from analyses of stomach contents, habitat depths, and observations of oral jaw tooth morphology. Results AFLP analyses resolved the phylogenetic relationships of the Perissodini, strongly supporting monophyly for each species. The character reconstruction of feeding ecology based on the AFLP tree suggested that scale eating evolved from general carnivorous feeding to highly specialized scale eating. Furthermore, scale eating is suggested to have evolved in deepwater habitats in the lake. Oral jaw tooth shape was also estimated to have diverged in step with specialization for scale eating. Conclusion The present evolutionary analyses of feeding ecology and morphology based on the obtained phylogenetic tree demonstrate for the first time the evolutionary process leading from generalised to highly specialized scale eating, with diversification in feeding morphology and behaviour among species.

Some highly isolated oceanic islands harbour endemic ground beetles that have lost the ability to fly. Here, we investigated the origin of the possibly extinct flightless giant ground beetle Aplothorax burchelli on St Helena Island in the South Atlantic. Aplothorax burchelli was initially considered to be a member of the subtribe Calosomina (=genus Calosoma) of the subfamily Carabinae (Coleoptera: Carabidae) closely related to the genus Ctenosta (= Calosoma subgenus Ctenosta), but this proposition was questioned due to its unique external and genital morphology. We conducted a phylogenetic analysis of mitogenome sequences using historical specimens of A. burchelli and samples of representative species of Carabinae. Our analysis of 13 protein-coding gene sequences revealed that A. burchelli is definitely a member of Calosomina, most closely related to a species of Ctenosta. Further analysis using NADH dehydrogenase subunit 5 gene sequences from most groups in Calosomina showed that A. burchelli formed a monophyletic group with Ctenosta species from Africa and Madagascar. Our results suggest that the ancestor of A. burchelli , which had the ability to fly, colonized St Helena from Africa after the emergence of the island 14 Mya, and has since undergone evolutionary changes in conjunction with loss of flight. [ABSTRACT FROM AUTHOR]

Although parental care is known to occur in a wide range of teleost fishes, postnatal provisioning of nutrition has been documented rarely. Here, we describe a novel example of bi-parental care in a teleost, i.e. mucus-provisioning behaviour in the scale-eating cichlid Perissodus microlepis endemic to Lake Tanganyika. Field observations revealed that young guarded by their parents frequently glanced towards the body surface of both parents. Furthermore, analyses of stomach contents of the young found the presence of ingested mucus, confirming that the young feed on the mucus secretions of their parents. The frequency of glancing behaviour increased with size of the young up to ~13 mm in standard length, but then declined with further growth. Additionally, the frequency of glancing of young towards their parents was higher when the frequency of foraging on plankton was lower. Underwater cage experiments revealed a higher rate of growth in the young kept in direct contact with their parents than in those not allowed direct contact. We conclude that glancing behaviour in young P. microlepis is a form of direct parental nourishment that confers growth benefits to the young when food abundance is low. [ABSTRACT FROM AUTHOR]

Unravelling the evolution of complex social organization in animals is an important aim, not least because it helps to understand the evolutionary roots of human sociality. Recent advances in comparative methods allow to approach this question in a phylogenetic context. The validity of such comparative approaches depends strongly on the quality of information regarding the behaviour, sociality, and reproduction of animals in natural systems, and on the quality of the phylogenetic reconstruction. Applying a novel comparative approach, a recent study of Dey et al. (, Nature Ecology & Evolution, 1, 137) concluded that evolutionary transitions to cooperative breeding in cichlid fishes were not associated with the social mating pattern. Here we argue that this result was adversely affected by equivocal classifications of mating patterns, and inadequate phylogenetic data. In order to illustrate the impact of the mating system misclassifications, we scored mating patterns as reported in the original literature and re‐analysed the dataset based on Dey et al.'s tree topology. The result suggests that the mating system does in fact significantly explain the evolutionary transition to cooperative breeding in lamprologine cichlids, but we submit that a reliable conclusion cannot be reached before improving the behavioural information and the underlying phylogenetic reconstruction. The problems identified in this case study are not unique and we urge caution in the interpretation of results from comparative phylogenetic studies in general. We do agree with Dey et al. () though that the lamprologine cichlids of Lake Tanganyika may constitute a fundamental test case for the theory of social evolution, but better information on their behaviour and phylogenetic relationships is needed to allow meaningful analyses. [ABSTRACT FROM AUTHOR]

Several studies in the last 20 years have revealed that morphological asymmetry in fish can be characterized as 'antisymmetry'. Antisymmetry is a lateral dimorphism in which each population consists of individuals with well-developed left sides (lefties) and well-developed right sides (righties). This dimorphism influences predator-prey interactions. In some piscivorous fishes, it has been found that predators can catch more prey of the opposite morphological type to themselves (cross-predation) than of the same morphological type (parallel-predation). Our previous work clarified that the predominance of cross-predation is caused by lateralized behaviors of predators and prey that correspond to their morphological antisymmetry. Moreover, based on the results of our behavioral observations, we hypothesized that parallel-predation can predominate when predators encounter the potential prey frontally. To test this hypothesis, in the present study we investigated the relationship between lateral morphological types of anglerfish ( Lophiomus setigerus) and those of the prey fishes found in their stomachs. Anglerfish attract potential prey using their first dorsal fin (illicium) as a lure, and their frontal encounters with potential prey fishes were photographed in situ and observed in an aquarium. The results of a stomach contents analysis indicated that parallel-predation predominated in five benthopelagic prey fish species (perches and eels). By contrast, five benthic prey fishes (gobies and weevers) exhibited the predominance of cross-predation. These results not only demonstrate the predominance of parallel-predation in a natural fish community, but also suggest that the relationship between morphological types of predator and prey species can be reversed depending on the lifestyle of prey. [ABSTRACT FROM AUTHOR]

The scale-eating cichlid Perissodus microlepis with asymmetric mouth is an attractive model of behavioral laterality: each adult tears off scales from prey fishes’ left or right flanks according to the direction in which its mouth is skewed. To investigate the development of behavioral laterality and mouth asymmetry, we analyzed stomach contents and lower jaw-bone asymmetry of various-sized P. microlepis (22≤SL<115mm) sampled in Lake Tanganyika. The shapes of the pored scales found in each specimen’s stomach indicated its attack side preference. Early-juvenile specimens (SL<45mm) feeding mainly on zooplankton exhibited slight but significant mouth asymmetry. As the fish acquired scale-eating (45mm≤SL), attack side preference was gradually strengthened, as was mouth asymmetry. Among size-matched individuals, those with more skewed mouths ate more scales. These findings show that behavioral laterality in scale-eating P. microlepis is established in association with development of mouth asymmetry which precedes the behavioral acquisition, and that this synergistic interaction between physical and behavioral literalities may contribute to efficient scale-eating. [ABSTRACT FROM AUTHOR]

Operation tests were applied to fuel cell with perfluorosulfonic acid (PFSA) polymer for membranes and electrodes. Degradation of PFSA polymer in catalyst layers alter the tests was investigated using 19F NMR. As the decomposition products, six kinds of the side-chain fragments were acquired from the water and the methanol extracts, and the each weight of the fragment was acquired. The side-chain fragments indicate that side-chain scission occurs at two positions in the side-chain. Moreover, reduction of relative equivalent weight for all PFSA polymer (EW*all) and amounts of PFSA polymer in catalyst layer after the tests suggests that main-chain scission occurs. Hydrogen peroxide vapor (HPV) exposure tests were applied to PFSA polymer membranes. PFSA polymer degradation after the tests was investigated using ion chromatography and total organic carbon analysis for trap water, and using 19FNMR for water/methanol extracts of membranes. Decomposition products other than fluoride ion accounted for a significant portion of degradation weight loss, and their generation rates were not proportional to fluoride ion release rate (FRR). The differences between the results of FRR and other decomposition product analysis strongly suggest the presence of two or more decomposition mechanisms in PFSA degradation. [ABSTRACT FROM AUTHOR]

The inheritance patterns of asymmetry in mouth opening in zebrafish were investigated using crossing experiments. Zebrafish exhibit asymmetric laterality in mouth opening, with each individual having either a leftward (righty) or rightward (lefty) bias. All righty incrosses produced only righty F1, whereas all lefty incrosses resulted in an F1 L:R ratio of 2:1. All test crosses between lefty and righty individuals resulted in an F1 L:R=1:1. These results were consistent with the hereditary pattern for Japanese medaka, three Tanganyikan cichlids, and a Japanese riverine goby. The pattern suggests a one-locus two-allele Mendelian model of inheritance, with the lefty allele being dominant over righty and the dominant homozygote being lethal. To determine the reason for the absence of lefty homozygotes, the survival rates of the offspring were examined according to developmental stage. Survival did not differ among combinations of parent laterality. Thus the mechanism underlying the lethality of the dominant homozygote remains unclear. This study showed that the mouth-opening laterality of zebrafish is genetically determined and that the direction follows a Mendelian inheritance pattern that is shared among cypriniform zebrafish, beloniform medaka, perciform cichlids, and a goby, suggesting a common genetic background in mouth-opening laterality among these species. [ABSTRACT FROM AUTHOR]

Vertebrates show left-right biases in turning direction, limb usage, predator-escape response and use of sensory organs. In particular, some fishes are known to have lateral biases in predatory behaviors corresponding to their morphological antisymmetry. To reveal the effects of these laterally biased behaviors on predator-prey interaction, we conducted behavioral tests of predatory events between largemouth bass, Micropterus salmoides, and freshwater gobies, Rhinogobius sp., both of which have individuals with a well-developed left side and individuals with a well-developed right side. The left-developed bass tended to approach the goby clockwise from behind, whereas right-developed individuals tended to approach counterclockwise. Congruently, left-developed gobies began their escape maneuvers at a longer distance from bass when they were approached clockwise than when they were approached counterclockwise, whereas right-developed gobies showed the reverse tendency. The longer the distance between bass and gobies at the start of goby escape, the more the subsequent bass strike or dash was delayed. Under these conditions, predation should be more successful when a left (right)-developed bass meets a right (left)-developed goby, and less successful when a left (right)-developed bass meets a left (right)-developed goby. This prediction was consistent with the difference in predation success in our test and in field data from Lake Biwa, Japan. We conclude that lateral biases in the behavioral direction of each morphological type will generate bias in predation success between different combinations of predator and prey types, leading to the maintenance of antisymmetric dimorphism through negative frequency-dependent natural selection. [ABSTRACT FROM AUTHOR]

Males are expected to adjust testes investment according to the varying level of sperm competition that they experience. Spatial and temporal variation in population density likely influences sperm competition. In herbivorous aquatic organisms, densities often decrease along a vertical depth gradient, because their food is photosynthetic and thus becomes less abundant in deeper regions where less light penetrates. This decrease should be dramatic on a steep slope, which allows testing of the association between density and testes investment at the within-population level. We tested the effect in the socially monogamous herbivorous cichlid fish Variabilichromis moorii living on a steep slope in Lake Tanganyika. We examined competitor density and food abundance as ecological factors, territory defense behaviors and phenotypic traits (testes investment and somatic investment), and compared them between shallow (4-6 m depth) and deep habitats (10-13 m depth) separated by several dozen meters. We found that food availability drastically decreased with increasing depth and that V. moorii was much more abundant in shallower habitats. Males in shallower habitats were in better physical condition (based on fat and liver mass) despite experiencing greater costs in terms of territory defense. Testes investment differed in areas with different competitor density and food abundance along a vertical depth gradient, but competitor density was the most explanatory factor of the difference. This suggests that this herbivorous fish would change testes investment in response to population density. [ABSTRACT FROM AUTHOR]

Behavioral lateralization has been documented in many vertebrates. The scale-eating cichlid fish Perissodus microlepis is well known for exhibiting lateral dimorphism in its mouth morphology and lateralized behavior in robbing scales from prey fish. A previous field study indicated that this mouth asymmetry closely correlates with the side on which prey is attacked, but details of this species' predation behavior have not been previously analyzed because of the rapidity of the movements. Here, we studied scale-eating behavior in cichlids in a tank through high-speed video monitoring and quantitative assessment of behavioral laterality and kinematics. The fish observed showed a clear bias toward striking on one side, which closely correlated with their asymmetric mouth morphologies. Furthermore, the maximum angular velocity and amplitude of body flexion were significantly larger during attacks on the preferred side compared to those on the nonpreferred side, permitting increased predation success. In contrast, no such lateral difference in movement elements was observed in acoustically evoked flexion during the escape response, which is similar to flexion during scale eating and suggests that they share a common motor control pathway. Thus the neuronal circuits controlling body flexion during scale eating may be functionally lateralized upstream of this common motor pathway. [ABSTRACT FROM AUTHOR]

Background: Asymmetry in animal bodies and behavior has evolved several times, but our knowledge of their linkage is limited. Tanganyikan scale-eating cichlids have well-known antisymmetry in their bodies and behavior; individuals open their mouths leftward (righty) or rightward (lefty), and righties always attack the right flank of the prey, whereas lefties attack the left. This study analyzed the morphological asymmetry in a scale-eating characiform, Exodon paradoxus, and its behavioral handedness. Methodology/Principal Findings: Each eight E. paradoxus was observed for 1-h with a prey goldfish in an aquarium to detect the behavioral handedness. Following the experiment, the lateral differences in the mandibles and head-inclination of these eight and ten additional specimens were analyzed. Both measurements on the morphology showed a bimodal distribution, and the laterality identified by these two methods was always consistent within a given individual, indicating that the characin has morphological antisymmetry. Furthermore, this laterality significantly corresponded to behavioral handedness; that is, lefties more often rasped scales from the right flank of the prey and vice versa. However, the correlation between laterality and handedness is the opposite of that in the cichlids. This is due to differences in the feeding apparatus and technique. The characin has cuspids pointing forward on the external side of the premaxilla, and it thrusts its dominant body side outward from its body axis on the flank of the prey to tear off scales. By contrast, the cichlids draw their dominant body side inward toward the axis or rotate it to scrape or wrench off scales with the teeth lined in the opened mouth. Conclusions/Significance: This study demonstrated that the antisymmetry in external morphology and the corresponding behavioral handedness have evolved in two lineages of scale-eating fishes independently, and these fishes adopt different utilization of their body asymmetry to tear off scales. [ABSTRACT FROM AUTHOR]