Graylings of Baikal Lake basin (Thymallus, Thymallidae): Diversity of forms and their taxonomic status

The morphological characters, molecular-genetic features, and patterns of the dorsal fin in different subspecies and forms of Arctic grayling Thymallus arcticus populating Lake Baikal and its tributaries, Irkutsk Reservoir, and Lake Khubsugul have been studied. Three groups are discernable. The first includes the white and black Baikal graylings T. a. baicalensis, as well as the western Siberian T. a. arcticus; the second group is represented by the Kosogol grayling T. a. nigrescens, and the third is composed of T. arcticus ssp. from the Yakchii lakes (the Verkhnyaya Angara basin) with a phenotype close to graylings populating the upper reaches of the Lena River. All of them are distinguished by some morphological characters, elements of the dorsal fin pattern, and by body coloration. The populations of black Baikal graylings are genetically uniform, and their distinctions from white Baikal graylings are insignificant, which agrees with the absence of a considerable divergence of these forms by a complex of meristic characters. It is assumed that in the black and white Baikal graylings the exchange by genetic information has either ceased quite recently, or persists, although, insignificantly. Some genetic remoteness of the west Siberian grayling from Irkutsk Reservoir and Nizhnyaya Tunguska, closely related to the Baikal grayling, is recorded. The formation of the Khubsugul subspecies is possibly a result of the contact of grayling populations during the rearrangements of the river system in the last glaciation period in the upper reaches of Yenisei and Selenga in Mongolia. The habitation in the Baikal system of the Upper Lena graylings indicates a connection between the Lena and Baikal basins in the past. The results of a multivariate analysis of meristic characters and the sequences of mitochondrial DNA confirm the conclusion made by Svetovidov (1931, 1936), concerning the absence of grounds to assign a species status to the Baikal forms.     

A complete survey of normal pores on a smooth shell ostracod (Crustacea): Landmark‐based versus outline geometric morphometrics

Pores and sensilla on ostracod shell have often been used in studies of ontogeny, taxonomy, and phylogeny of the group. However, an analysis of sexual dimorphism and variation between valves in the number and distribution of pores is lacking. Also, such studies have never been done on a widely distributed, morphologically variable, and weakly ornamented freshwater ostracod. Here, we survey pores in one such species, Physocypria kraepelini . We choose 27 homologous pores as landmarks for 2D‐geometric morphometric analysis, with the aim to assess intersexual and between valves variation in size and shape relative to the Fourier outline analysis. This species has only simple (Type A) pores with and without a lip, and each pore carries an undivided sensory seta. Our results show that the total number of pores varies (from 270 to 296), but this is not associated with a specific valve. Males carry fewer pores than females, however no sex specific pores are found. Small intrapopulation divergence of the Cyt b molecular marker (1%) indicates that morphological variability is not species related. We found that P. kraepelini exhibits directional asymmetry of size and shape, sexual size dimorphism (SSD) but lacks sexual shape dimorphism (SShD). Two geometric morphometrics methods were congruent in the estimation of SSD, SShD, and directional asymmetry of shape but differ in the statistical evaluation of directional asymmetry of size. Contrary to other animal groups, our study suggests that ostracods have more pronounced directional asymmetry of shape compared to directional asymmetry of size.     

New data on the distribution of the Upper Lena form of grayling (Thymallidae) in the basin of Lake Baikal and its taxonomic status

The habitation of the Upper Lena form of grayling in the upper reaches of the Tiya (the northwestern tributary of Lake Baikal) and Barguzin rivers, where Baikal grayling Thymallus baicalensis Dyb. also occurs, was established. The listed forms can be diagnosed from the body coloration, dorsal fin pattern, combination of meristic characters, as well as by molecular-genetic methods. The sympatric habitation of the Upper Lena and Baikal graylings in the northern tributaries of Baikal, considerable phenotypic and genetic differences, as well as the absence of individuals with intermediate characters, make it possible to consider them as independent species. The results indicate the need to revise the intraspecies structure of the Arctic grayling T. arcticus (Pall.) and the entire genus Thymallus. The localization of populations of the Upper Lena grayling in the upper reaches of Baikal tributaries testifies to its possible penetration of the Baikal basin by means of temporary river catchments that formed in the past and which are probable at present at watersheds with the Lena River. A wider distribution of this form is apparently hindered by the pressure of aboriginal species.     

Sexual dimorphism in the skull geometry of newt species of <Emphasis Type="Italic">Ichthyosaura,Triturus</Emphasis> and <Emphasis Type="Italic">Lissotriton</Emphasis> (Salamandridae,Caudata, Amphibia)

In this study, we applied geometric morphometrics to explore variations in the level and pattern of sexual size dimorphism (SSD) and sexual shape dimorphism (SShD) of the ventral cranium in three different Modern Eurasian newt taxa (Ichthyosaura alpestris, Triturus species group and Lissotriton vulgaris). The ventral cranium is the part of the skull that is more directly related to foraging and feeding. Our results indicate that the level and pattern of sexual dimorphism in the ventral cranium differ among Modern Eurasian newt taxa. Regarding sexual dimorphism in skull size, Ichthyosaura alpestris and Triturus species show female-biased patterns (females are larger than males), whereas Lissotriton vulgaris appears to be non-dimorphic in skull size. In I. alpestris and Triturus species, SShD is mostly absent, whereas in L. vulgaris, SShD is more pronounced. A high level of variation between populations in both SSD and SShD indicates that local conditions may have a profound effect on the magnitude and direction of sexual dimorphism. The significant sexual differences in ventral cranium size and shape indicate possible subtle intersexual differences in ecological demands due to diet specialisation, in spite of similar general ecological settings.     

Graylings (Thymallidae) of water bodies in western Mongolia: Morphological and genetic diversity

Morphological characters, the pattern on the dorsal fin, some biological parameters, and variations of mtDNA control region were studied in graylings of the upper reaches of the Khovd River (western Mongolia). Obtained results demonstrated that grayling of this part of the river was presented by great (predatory) and small (benthophagous) forms of Mongolian grayling Thymallus brevirostris. The last form was erroneously equated to Arctic grayling Thymallus arcticus. However, genetic unity of great and small forms, homogeneity by meristic characters, and similarity in varying the color of scales and pattern on the dorsal fin attest their belonging to the same species. Water bodies of the Central Asian basin are also populated with populations of Mongolian grayling with mixed type of feeding and signs of external structure, which are typical for both forms in a different degree.     

Ontogenetic pattern,morphological sexual and side dimorphism in the saccular otolith of a scaleless killifish Aphanius furcatus (Teleostei: Aphaniidae)

The relationships between ontogenetic and sexual factors with otolith morphology are poorly studied in killifish genus Aphanius Nardo, 1827. We studied the ontogeny, morphological sexual and side dimorphism in the saccular otolith of a scaleless killifish Aphanius furcatus inhabiting a high‐salinity environment in Southern Iran. The results highlighted growth‐dependent variability in the otolith of A. furcatus, which might be useful to define its certain developmental stages. We underlined those characters that were consistent during fish development, which could be valuable to identify the species. Considering the ontogenetic pattern of otolith in A. furcatus, it was found that the otoliths of young individuals have largely been influenced by sexes and that it was reduced during the fish development. Morphological side dimorphism in the otolith of A. furcatus tends to be increased during fish development. The outcomes of this study are essential since they have provided new information on ontogenetic changes and the sexual and side dimorphism of Aphanius otolith.     

Neoboeckella loffleri (Copepoda: Calanoida), a new species and genus from Bolivian fresh waters

A new species and genus of freshwater calanoid, Neoboeckella loffleri, is described from shallow, high-altitude waters. N. kinzeli (Löffler), 1955, is also included in the new genus. Neoboeckella, like Hemiboeckella Sars from Australia, is characterized inter alia by an exceptionally high degree of sexual dimorphism in size.     

Seismosensory system of the Lower Amur rayling <Emphasis Type="Italic">Thymallus tugarinae</Emphasis> (Thymallidae) from the Anyui River

Data on specific features of the seismosensory system of the Lower Amur grayling Thymallus tugarinae from the Anyui River (basin of the Lower Amur) are provided. According to the number of openings in canals of the seismosensory system, individuals from the middle course significantly differ from graylings from the lower course in greater values of interpopulation diversity and fluctuating asymmetry. According to the number of pores of canals of the seismosensory system, the Lower Amur grayling differs from the East Siberian T. arcticus pallasi, Kamchatka T. a. mertensii, and Alaska T. a. signifer graylings.     

Is sexual size dimorphism variable? Data on species of the genus <Emphasis Type="Italic">Martes</Emphasis> in the Urals

Sexual dimorphism in cranial measurements was studied in Ural populations of two species of the genus Martes, M. martes and M. zibellina. The index of sexual dimorphism averaged 8.39% in the former and 9.03% in the latter, being subject to geographic variation. For the first time, temporal variability of sexual dimorphism in dimensional characters was revealed in mammals by analyzing M. zibellina samples collected in different decades of the second half of the 20th century. The majority of corresponding microtrends had reciprocating dynamics.     

A new species of electric knifefish,genus Compsaraia (Gymnotiformes: Apteronotidae) from the Amazon River,with extreme sexual dimorphism in snout and jaw length

Abstract

A new species of the neotropical electric fish genus Compsaraia is described from the western Amazon of Peru and Brazil. Compsaraia samueli is distinguishable from all other apteronotids by sexual dimorphism in which mature males exhibit extreme elongation and slenderness of the snout and jaws. Compsaraia samueli is readily distinguishable from its only congener, C. compsa, by more caudal‐fin rays (17–18 vs. 13–16), a shorter caudal peduncle (mean length 9% vs. 34% body length to end of anal fin), a less tapering body shape in lateral profile (mean ratio of body depth at origins of anal fin and dorsal organ 93% vs. 75%), and a smaller maximum adult body size (230 vs. 305 mm). The genus Compsaraia is readily separated from other apteronotids by a pale antorbital stripe and a pale L‐shaped patch over the supra‐temporal canal. The phylogenetic position of C. samueliis estimated by inclusion in a previously published data matrixof osteological and other morphological characters. Comparisons of the cranial bones in apteronotids shows the derived morphology of C. samueli to be a composite of three developmentally and phylogenetically discrete characters: (1) positive allometric growth before sexual maturity in both sexes of the pre‐orbital region of the neurocranium, (2) positive allometric growth of the (oral) jaws, and (3) secondary sexual dimorphism of snout and jaw morphology. The genus Compsaraia represents one of at least three phylogenetically independent cases of snout elongation and one of at least four cases of jaw elongation within the Apteronotidae. Compsaraia samueli also represents one of at least four cases of secondary sexual dimorphism in snout and jaw length within the Apteronotidae. The phylogenetic distribution of snout and jaw characters within the Apteronotidae suggests the influence of both sexual and trophic functional influences on the evolution of head morphology.     

THE EVOLUTION OF SEXUAL DIMORPHISM BY SEXUAL SELECTION: THE SEPARATE EFFECTS OF INTRASEXUAL SELECTION AND INTERSEXUAL SELECTION

Libellula luctuosa, a pond dragonfly found in eastern North America, is apparently sexually dimorphic. Previous studies of the mating behavior in this species suggested that both male-male competition and female mate choice are important influences. Males compete for territories, where they attract females and where mating occurs. Female behavior influences both the copulation success and the fertilization success of males. Because of temporal and spatial separation of these episodes of sexual selection, multivariate and nonparametric statistical techniques could be used to investigate the influence of components of sexual selection on various sexually dimorphic traits. Sexual dimorphism in L. luctuosa was first quantified; then the direct effects and the form of selection were estimated. Sexually dimorphic wing size, body size, wing coloration, and body coloration are distributed either continuously or discontinuously between the sexes in L. luctuosa. These traits have apparently diverged between the sexes as a result of directional sexual selection. Body size is further influenced by stabilizing selection. Intrasexual selection (success in gaining access to a territory) and intersexual selection (success in copulation and fertilization) can influence the same or different sexually dimorphic characters. Body size is influenced by directional selection during the intrasexual phase of sexual selection and is also influenced by stabilizing selection during intersexual selection. The size of the brown wing patch is influenced by directional selection, primarily during the intersexual phase of sexual selection. There is directional selection on the white wing patch during both phases. Thus, the different proximate mechanisms of sexual selection may jointly or separately affect the evolution of sexually dimorphic characters. Further empirical and theoretical investigations into the differences in the effects of intrasexual selection and intersexual selection are needed to clarify the circumstances leading to separate consequences of these two mechanisms of sexual selection.     

Biological and morphological characteristic of the Arctic grayling Thymallus arcticus (Thymallidae) from Alpine Lakes of the basin of the upper reaches of the Angara River

Specific morphological and biological features of the populations of the Arctic grayling Thymallus arcticus from lakes Urunge Nur, Mongosha, Sadaiskoe, and Gusinoe situated in the Upper Sayan Mountains in the sources of the Oka and Kitoi rivers (the basin of the upper reaches of Angara) that have been not studied previously are considered. The results of the multivariate analysis indicated the uniformity of all samples by 12 meristic characters. Graylings of the studied lakes are similar to graylings from the Angara and Nizhnyaya Tunguska rivers and the Irkutsk Reservoir, as well as to the Baikal grayling T. a. baicalensis in body coloration and pattern of the dorsal fin. The samples studied and T. a. baicalensis considerably differ in the above characters from populations inhabiting the basin of the upper reaches of the Ob that belong to the nominative subspecies T. a. arcticus. The differences established between the Angara-Yenisei and the Upper Ob graylings give grounds to doubt the justification of including them into one taxon, T. a. arcticus. The same is evidenced by the results of molecular-genetic studies of graylings from the Palearctic (Froufe et al., 2005). The data obtained indicate the necessity of ascertaining the boundaries of the ranges of different forms of graylings in the Angara-Yenisei and Ob basins and of the revision of their taxonomic status. Populations inhabiting the Angara and Yenisei rivers, except their sources and the section of the lower reaches, should be assigned to the Baikal subspecies, T. a. baicalensis. Graylings from different lakes in the upper reaches of Oka and Kitoi differ in their linear-weight indices. The indices of growth are highest where they occupy a dominant position. Their food in the summer consists mainly of the larval and imaginal stages of amphibiotic and aerial-terrestrial insects. Only in Lake Urunge Nur do graylings of elder age groups use fish of other species for food. The extreme conditions of habitation of the grayling in Lake Gol’tsovoe are the cause of food deficiency, which determines its dwarfism.     

Phylogenetic evidence for loss of sound production and a shift in sexual recognition signals in Hamadryas butterflies (Nymphalidae: Biblidinae)

The neotropical butterfly genus Hamadryas Hübner comprises 20 species that exhibit an intriguing variation in their natural history traits. Although revised in 1983, no phylogenetic hypothesis was presented: the first phylogenetic hypothesis is estimated here based on 93 characters and including species from the three other genera in the tribe Ageroniini. The phylogeny is used to test the monophyly of the genus, establish the sister group of Hamadryas and identify its apomorphies. The tree allows the inference of patterns of character change in sound production and sexual dimorphism. Implied weights show that Hamadryas is monophyletic and corroborate Ectima Doubleday as a sister genus. Previously suggested subgenera for Hamadryas were non‐monophyletic, with the exception of the laodamia clade, supported by the presence of a complete sterigma. Sound production is inferred to be a derived condition in Hamadryas that has been lost in the laodamia clade. This, plus the presence of androconial organs and sexual dimorphism in the laodamia clade, suggests a shift in sexual recognition signalling. Furthermore, the phylogeny indicates that the colour pattern of males in the laodamia clade is novel, supporting a Darwinian origin of sexual dimorphism.     

Exploring sexual differences in external morphology and limb muscles of Hylarana guentheri (Anura: Ranidae) during non-breeding season

Studying sexual dimorphism facilitates better understanding of the general intersexual divergence of the same species and gives insights into the impact of selective forces on each sex. The sexual dimorphism in anuran external morphology or limb muscles has been well studied in reproductive context, but less so outside the breeding season. Herein, the intersexual differences in 12 external morphological characteristics, 11 forelimb and 12 hindlimb muscles of Hylarana guentheri during non-breeding season were investigated. We found that the males possessed significantly greater head width and tympanum diameter, while the females had significantly larger body size and head length; there was no sexual differences in the examined limb muscles, except for the mass of flexor carpi radialis. The larger body size means greater fertility for the females, and the longer head may be related to the allometry and reduce resource competition. For the males, the wider head is likely correlated with prey size and male–male competition, and also, the larger tympanum and heavier flexor carpi radialis probably aid the productive success. This study provides the comprehensive morphological accounts about the sexual differences of H. guentheri during non-breeding season, which will contribute to clarify the sex-specific resource allocation and reproductive strategies of anurans.     

Sexual Selection and Dynamics of Jaw Muscle in Tupinambis Lizards

Sexual dimorphism patterns provide an opportunity to increase our understanding of trait evolution. Because selective forces may vary throughout the reproductive period, measuring dimorphism seasonally may be an interesting approach. An increased male head size may be important in intersexual and intrasexual interactions. In Tupinambis lizards, a big head is attributed in part to a large adductor muscle mass. Competition for mating can differ in species with different sex ratio and different degrees of sexual size dimorphism. We examined sexual differences in mass of the pterygoideus muscle, its temporal variation throughout the reproductive period and the relationship between muscle and reproductive condition in Tupinambis merianae and T. rufescens. We characterized sexual size dimorphism and sex ratio in both species. Mature males had larger jaw muscles than mature females in both species, mainly during the reproductive season. The dimorphism in jaw muscle was due to an increase in muscle mass in sexually active males. Seasonal increases in muscle mass and variation between immature and mature individuals suggest that the jaw muscle might be a secondary sexual character. We propose that the pterygoideus muscle may act as a signal of reproductive condition of males because it is associated with testis size and sperm presence. The patterns of sexual dimorphism in jaw muscle in both species were similar; however, the comparison shows how sexual characters remain dimorphic in different competition contexts and in species with different degrees of body size dimorphism. Our results suggest that jaw muscle as sexual character could be influenced by inter- and intrasexual selective pressures.     

Comparative analysis of oocyte growth in two forms of Baikal grayling <Emphasis Type="Italic">Thymallus baicalensis</Emphasis> (Thymallidae)

Size characteristics of oocytes of the elder generation of the end of previtollogenesis, as well as of the beginning and middle of the phase of cytoplasm vacuolization in two forms of Baikal grayling Thymallus baicalensis were studied. In each form, the dependence of oocyte parameter on the age and size of females is traced. During the studied phases of development—at the termination of the period of previtellogenesis and at the proper beginning of the period of vitellogenesis—oocytes of the white and black graylings have similar sizes; cytoplasm vacuolization in white grayling proceeds less actively. It was established that differences in the diameter of mature ovicells in the black and white Baikal graylings result from dissimilar rate of accumulation of trophic substances in the oocytes of the given forms of this species and are determined by differences in the rate of growth of oocytes during the formation in them of yolk inclusions.     

Phylogenetic analyses of the family Tetraonchidae (Platyhelminthes: Monogenea)

A phylogenetic reconstruction of the monogenean family Tetraonchidae was carried out by methods of parsimony-based cladistics. The analysis included 20 species of tetraonchids and two out-groups (Sundanonchus tomanorum and Dactylogyrus amphibothrium) and was based on 34 morphofunctional characters. Software PAUP 4.0 and Winclada were used for the phylogenetic reconstructions. Obtained results allow proposing a preliminary phylogenetic hypothesis of the family Tetraonchidae along with the discussion of host-parasite association. According to the current taxonomic view, the family Tetraonchidae included two genera. Cladistic analysis showed a monophyly of the family and the genus Tetraonchus Diesing, 1858. Two representative of the former genus, Tetraoncus monenteron and T. borealis, parasitize the pikes (Esocoformes: Esocidae) and the grayling (Salmonidae: Thymallinae) respectively. The genus Salmonchus Spassky et Roytman, 1958 has a complicated structure and its intrageneric relationships were not completely resolved; in general, the analysis allows to recognise several species groups: Salmonchus oncorhynchi--the parasite of the Oncorhynchus masou smolt living during the first year of life in fresh water; four species (S. variabilis, S. gussevi, S. grumosus, S. alaskensis) inhabiting specifically the whitefishes (Salmonidae: Coregoninae); all reminder of Salmonchus species occurring on the salmons (Salmonidae: Salmoninae). The bootstrap test gives a support only for the following clades: family Tetraonchidae (75%), genus Tetraonchus (88%); a group of Salmonchus species associated with the whitefishes (93%) and grouping of four species (S. huhonis, S. pseudolenoki, S. skrjabini and S. lenoki) from the lenoks (Brachymystax) and taimens (Hucho) (61%).     

Diet and sexual dimorphism in the very catholic lizard genus,Leiocephalus of the Bahamas

Summary Over 5,000 prey items from specimens of Bahamian Leiocephalus lizards were measured and identified taxonomically. The diet in general consists mainly of arthropods, but much plant matter is also eaten, including flowers and buds as well as fruit. Lizards comprise about 2% of the diet by volume. Individuals inhabiting relatively small islands are more likely to have eaten plant matter than those from relatively large islands. Within the most widespread species (carinatus), sexual dimorphism in size is greater, the smaller the number of sympatric species in its structural habitat. Prey-size differences between differently sized Leiocephalus are greater, the greater the dimorphism. However, even the most dimorphic sexes take rather similar prey sizes. For all Bahamian species combined, the inverse correlation of sexual dimorphism with sympatric species is not as strong as an inverse correlation with latitude. We suggest that sexual selection on female size to increase the clutch size that can be carried may have affected sexual dimorphism in the genus.     

Phylogenetic analysis of Micrathena and Chaetacis spiders (Araneae: Araneidae) reveals multiple origins of extreme sexual size dimorphism and long abdominal spines

The phylogenetic relationships amongst the New World spiny orb‐weaving spiders Micrathena and Chaetacis were assessed through parsimony and Bayesian analyses of morphological characters. A total of 146 characters was scored for ten outgroup taxa and 37 Micrathena and four Chaetacis species. The results indicate that Chaetacis nests within Micrathena and we propose Chaetacis as a junior synonym of Micrathena. Twelve subgeneric species groups of Micrathena are recognized and diagnosed. Species with extremely long spines evolved at least eight times in the genus and we suggest that this may be related to antipredator defences. Micrathena is primitively sexually monomorphic and extreme sexual size dimorphism has arisen at least six times in the genus. Most of these events are because of enlargement of the female in relation to the ancestral size, although in two cases sexual dimorphism was attained through male reduction, adding more data to the ‘giant females’ vs. ‘dwarf males’ controversy. The genus is probably of South American origin and has repeatedly invaded Central and North America. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 14–53.     

The effects of fisheries management on harvest rates of native and non-native salmonid fish species

In central Europe, both brown trout Salmo trutta and European grayling Thymallus thymallus are threatened native salmonid species with high value in recreational angling and nature conservation. On the other hand, rainbow trout Oncorhynchus mykiss and brook trout Salvelinus fontinalis are intensively stocked non-native species of high angling value but no value for nature conservation. This study tested if harvest rates of native salmonids are negatively correlated to intensive stocking and harvest rates of non-native salmonids in inland freshwater recreational fisheries. Data were collected from 250 fishing sites (river and stream stretches) over 13 years using mandatory angling logbooks. Logbooks were collected from individual anglers by the Czech Fishing Union in the regions of Prague and Central Bohemia, Czechia (central Europe) and processed by the author of this study. In result, anglers harvested 200,000 salmonids with total weight of 80 tons over 13 years. Intensive stocking of multiple salmonid species lead to slightly lower harvests of native salmonids. Inversely, intensive harvests of multiple salmonid species lead to slightly higher harvest of native salmonids. Recapture rates of stocked salmonids were relatively low (0.6%–3.7%), proving fish stocking moderately ineffective. Since the effects of non-native salmonid stocking and harvest rates on native salmonid harvest were significant but not strong, it is suggested that rivers and streams that support fishing for non-native salmonids still support fishing for native salmonids. However, this idea does not apply for fishing sites with really high intensity of non-native salmonid stocking – harvest rates of natives were very low on these fishing sites.