Breeding and bionomics of the British members of the Jaera albifrons group of species (Isopoda: Asellota)

A detailed 24 month study (1968–1970) of the breeding cycles and population structures of the four British members of the Jaera albifrons Leach group of species (Isopoda: Asellota) has been carried out in Milford Haven, Pembrokeshire. The species often show overlapping ecological distributions and they can only be identified on male secondary sexual characters. Sampling stations had therefore to be selected carefully to ensure that single species populations were studied. In this way virtually single-species populations of Jaera forsmani Bocquet, Jaera praehirsuta Forsman, and Jaera ischiosetosa Forsman have been studied but the Jaera albifrons Leach population was somewhat mixed with J. ischiosetosa , particularly when the stream normally inhabited by the last species dried up in the summer. Only one hybrid was found (between J. albifrons and J. ischiosetosa ) in a total of 6214 specimens collected. Gravid females were taken every month for each species with peaks of breeding occurring during spring and summer. Young were liberated throughout the year by J. albifrons and J. ischiosetosa , but with summer peaks. J. praehirsuta and J. forsmani had a particularly limited summer period for the release of juveniles. J. forsmani was consistently the largest species and has a restricted geographical distribution. J. praehirsuta also has a patchy distribution. The sex ratio of males to females was never 1:1 for any species. Females outnumbered males by up to 14:1, with the sex ratio varying throughout the year except in J. praehirsuta where it remained at about 1.5:1.     

Sexual behaviour and evolution of sexual dimorphism in body size in Jaera (Isopoda Asellota)

Individuals of the genus Jaera do not mate at random. In the species from the Mediterranean group, J. italica and. J. nordmanni, large males and medium sized females are at an advantage and their sizes are positively assorted. These effects are attributable to sexual competition between males. In the Ponlo-caspian species J. istri, no advantage of large males exists, but sexual selection could be the cause for a long passive phase prior to copulation and for normalizing selection upon female size at pairing. In the Atlantic species, J. albifrons, no selection can be ascertained.
Differential mating success in males appears as one of the causes of the evolution of sexual dimorphism in body size, which makes males larger, of equal size, or smaller than females according to the species. The reason for this reversal in dimorphism seems to differ in the two sexes. Sexual selection provides an explanation for the evolution of male size, while the interspecific changes in female length are more likely due to ecological factors.     

Disparity in population differentiation of sex-linked and autosomal variation in sibling species of the Jaera albifrons (Isopoda) complex

The genetic variation at four enzyme loci is described for 22 populations of three Jaera species--J. albifrons, J. ischiosetosa, and J. praehirsuta--in the J. albifrons complex (Crustacea, Isopoda) in Denmark. The variation at three of the loci is similar, with the allele frequency spectra close to each other in all three species. An evolutionary tree based on the variation at these three loci revealed that the populations from the different species are completely intermixed in the tree. This was supported by hierarchical F-statistics where the between-species component was zero. At a fourth locus, Gpi (glucose phosphate isomerase), the species differ substantially. This locus is sex linked in J. ischiosetosa, but in the two other species, J. albifrons and J. praehirsuta, it is either found on autosomes or is sex linked with a high recombination rate between the locus and the centromere. An evolutionary tree for this locus partitions the populations into separate groups and a hierarchical F-statistic has a between-species component of about 50%. The results are attributed to introgression with a higher rate for autosomes than for sex chromosomes.     

Sex and species differences in neuromodulatory input to a premotor nucleus: a comparative study of substance P and communication behavior in weakly electric fish

Many electric fish species modulate their electric organ discharges (EODs) to produce transient social signals that vary in number and structure. In Apteronotus leptorhynchus, males modulate their EOD more often than females, whereas in Apteronotus albifrons, males and females produce similar numbers of modulations. Sex differences in the number of EOD modulations in A. leptorhynchus are associated with sex differences in substance P in the diencephalic nucleus that controls transient EOD modulations, the CP/PPn. These sex differences in substance P have been hypothesized to regulate sex differences in the production of EOD modulations. To comparatively test this hypothesis, we examined substance P immunoreactivity in the CP/PPn of male and female A. leptorhynchus and A. albifrons. Because the number of EOD modulations is sexually monomorphic in A. albifrons, we predicted no sex difference in substance P in the CP/PPn of this species. Contrary to this prediction, male A. albifrons had significantly more substance P in the CP/PPn than females. This suggests that sex differences in substance P are not sufficient for controlling sex differences in the number of EOD modulations. Modulation structure (frequency excursion and/or duration), however, is also sexually dimorphic in A. leptorhynchus and is another possible behavioral correlate of the sexually dimorphic distribution of substance P. The present study found pronounced sex differences in the structure of EOD modulations in A. albifrons similar to those in A. leptorhynchus. Thus, sex differences in substance P may influence sex differences in the structure, rather than the number, of EOD modulations.     

The morphology of mouthparts,wings and genitalia of Paleozoic insect families Protohymenidae and Scytohymenidae reveals new details and supposed function

Megasecoptera is an extinct group of insects with specialized rostrum-like mouthparts, which is a synapomorphy shared with all members of the Late Paleozoic Palaeodictyopterida, and markedly slender wings that are unable to flex backwards. Here we describe the close up morphology of Protohymenidae and Scytohymenidae and uncover new aspects of the endoskeleton (tentorium) of the head, structure of the mouthparts with discernible proximal part of stylets controlled by muscles, surface of compound eyes that consist of a hexagonal pattern of large facets, structure and microstructures on the wings and reconstruct male and female external genitalia using ESEM and light stereomicroscopy. Furthermore, we describe Protohymen novokshonovi sp. n. based on an exceptionally well preserved fossil from the early Permian at Tshekarda in Russia, which shows crucial details, and the earliest species of Protohymenidae, Carbohymen testai gen. et sp. n. from a late Carboniferous siderite nodule at Mazon Creek in Illinois, USA. Our comparative study confirmed a set of structural and microstructural details on their wings, such as the composite anterior wing margin, development of an apical cell and the previously unknown external genitalia. Based on the results and comparison of homologous structures known primarily for extant relatives, such as mayflies and dragonflies, we outline for the first time the function of the mouthparts, in particular, the stylets, structure of the tentorium, vision provided by large hexagonal ommatidia and male copulatory structures bearing curved claspers for holding a female during copulation and penial lobes with seminal grooves.     

柳二十斑叶甲两亚种比较形态学研究(鞘翅目,叶甲科,叶甲亚科)

柳二十斑叶甲包括两个亚种,分别为指名亚种和高山亚种.两者可以通过头部、前胸背板、足和鞘翅的斑纹区分开来.本文对这两个亚种的外部和内部的形态结构进行了研究,主要包括触角、口器、中胸、后胸、后翅、足、雌雄外生殖器、成虫和幼虫的相关结构.并对确定亚种的形态特征进行了探讨.     

The phylogenetic information carried by a new set of morphological characters in planthoppers: the internal mouthpart structures and test in the Cixiidae model (Hemiptera: Fulgoromorpha)

Internal morphological structures of Cixiidae mouthparts are described and compared in various representatives of the Cixiidae and several other representatives of hemipterans. The morphological study shows that the mouthpart structures have not evolved uniformly and reveals the great disparity of these structures. Particularly, the connecting system of the mouthparts, localisation of salivary canal and shape of the mandibular and maxillar stylets provide together a new set of 17 new characters. A parsimonious analysis to evaluate the phylogenetic interest carried by these 17 selected characters shows that mouthpart structures have not evolved anarchically, but that they indeed carry some phylogenetic information that will be useful to be included in further morphological phylogenetic analysis.     

Systematics of the dobsonfly subfamily Corydalinae (Megaloptera: Corydalidae)

Abstract. The genera of Corydalinae are redefined, and representative characters are figured for each genus. New character sources, such as mouthparts and internal female genitalia, are investigated, as well as traditional male genitalia and wings. Allohermes is synonymized with Protohermes, Doeringia with Platyneuromus . Intergeneric relationships are hypothesized on the basis of a cladistic analysis. Acanthacorydalis and the New World genera form a monophyletic group, as do Protohermes and Neurhermes , and Neuromus and Neoneuromus. Chloroniella belongs in the Acanthacorydalis - New World lineage, but exact placement is uncertain. A phyletic sequence classification is proposed on the basis of the cladistic analysis.     

Mandibulate chironomids: primitive or derived? (Diptera: Chironomidae)

Abstract Mandibulate functional mouthparts are reported in males and females of the two Early Cretaceous Chironomidae (Diptera): Wadelius libanicus Veltz et al., 2007 (in Tanypodinae) and Libanochlites Brundin, 1976 (transferred from the Podonominae to the Tanypodinae). Females of Haematotanypus libanicus gen.n. et sp.n. (subfamily Tanypodinae) have mandibulate mouthparts. Although currently considered as plesiomorphic structures, the presence of such mandibulate mouthparts in these Tanypodinae and in the recent Podonominae genera Archaeochlus and Austrochlus could correspond to reversals, based on a parsimony argument after the current chironomid phylogeny. On the contrary, similar mandibulate mouthparts probably are plesiomorphic in the Early Cretaceous Cretaenne kobeyssii gen.n. et sp.n. and Cretaenne inexpectata sp.n. (Aenneinae or stem group of recent Chironomidae).     

The effects of stylopisation in two species of planthoppers in the Krian District, West Malaysia (Homoptera: Delphacidae)

Nilaparvata lugens (Stål) and Sogatellafurcifera (Horvath) found in light trap samples at Titi Serong Padi Experimental Station, Parit Buntar, Perak, West Malaysia, between 1972–74, were stylopised by Elenchus sp. The types of changes caused by stylopisation to the secondary and tertiary sexual organs/characters are discussed. The sex of the specimens with extremely reduced secondary sexual organs is analyzed. The development of the male secondary sexual organ from the late fifth instar to a fully developed male is traced. The reasons why the secondary sexual organs of stylopised male Delphacidae are more reduced than females are outlined.     

Niche separation in Varecia variegata rubra and Eulemur fulvus albifrons: II. Intraspecific patterns

Based on a year-long field study in northeastern Madagascar, I summarize annual patterns of niche use (food patch size, diet, forest height, and forest site) in two sympatric lemurs, Varecia variegata rubra and Eulemur fulvus albifrons. Furthermore, I examine intraspecific patterns of niche use according to sex, season, and reproductive stage in these two lemurs that differ in terms of energetic investment in reproduction. Lemurs as a group provide a special opportunity to test hypotheses concerning sex differences in niche use. Due to their body size monomorphism and seasonal, synchronous pattern of breeding, it is possible to directly evaluate whether sex differences in diet reflect high energetic investment in reproduction by females. Results confirm the hypothesis that intraspecific variation in niche use (e.g., sex differences, seasonal differences) would be more pronounced in V. v. rubra than in E. f. albifrons, due in large measure to the former's relatively high energetic investment in reproduction: 1a) Dietary sex differences in V. v. rubra are most pronounced during costly reproductive stages and involve acquisition of low-fiber, high-protein plant foods. Females of both species consume more seasonally available low-fiber protein (young leaves, flowers) relative to conspecific males during the hot dry season, but only in V. v. rubra females is this pattern also evident during gestation and lactation. 1b) The diets of female V. v. rubra and female E. f. albifrons are more similar to each other than are the diets of conspecific males and females in the case of V. v. rubra. This is not uniformly the case for female E. f. albifrons. This finding confirms a hypothesis put forward in Vasey ([2000] Am J Phys Anthropol 112:411-431) that energetic requirements of reproductive females drive niche separation more than do the energetic requirements of males. 1c) Both species synchronize most or all of lactation with seasonal food abundance and diversity. E. f. albifrons shows a more protracted period of synchrony, and this may contribute to its wide biogeographic distribution in Madagascar. 2) Sex differences and seasonal differences in microhabitat use reflect intraspecific patterns of thermoregulation, predator avoidance, and in the case of V. v. rubra, reproduction. One important factor selecting for body size monomorphism in lemurs appears to be the tight synchrony between lactation and periods of food abundance afforded by annual, seasonal breeding.     

Setiferous structures of male coleoptera

A scanning electron microscopy study was made of the male setiferous sex patches and analogous structures in 11 families of Coleoptera (Anthribidae, Bruchidae, Ciidae, Cleridae, Coccinellidae, Dermestidae, Leiodidae, Ptinidae, Staphylinidae, Tenebrionidae, and Ostomatidae). These secondary sexual characters appear to have several features in common including relatively long, often ridged, setae, cuticular ducts (frequently cribriform pore plates), and the production of a secretion. It is suggested that these structures may all be concerned with the production, release, and dissemination of pheromones.     

Patterns of fluctuating asymmetry in sexual ornaments predict female choice

Extravagant secondary sexual characters are assumed to have arisen and be maintained by sexual selection. While traits like horns, antlers and spurs can be ascribed to intrasexual competition, other traits such as extravagant feather ornaments, displays and pheromones have to be ascribed to mate choice. A number of studies have tested whether females exert selection on the size of male ornaments, but only some of these have recorded female preferences for the most extravagantly ornamented males. Here I demonstrate that female choice can be directly predicted from the relationship between the degree of fluctuating asymmetry and the size of a secondary sexual character. Fluctuating asymmetry is an epigenetic measure of the ability of individuals to cope with stress, and it occurs when an individual is unable to undergo identical development of an otherwise bilaterally symmetric trait on both sides of its body. There is a negative relationship between the degree of fluctuating asymmetry and the absolute size of an ornament in those bird species with a female preference for the largest male sex trait, while there is a flat or U-shaped relationship among species without a female preference. These results suggest that females prefer exaggerated secondary sexual characters if they reliably demonstrate the ability of males to cope with genetic and environmental stress. Some species may demonstrate a flat or U-shaped relationship between the degree of fluctuating asymmetry and the absolute size of an ornament because (i) the genetic variance in viability signalled by the secondary sex trait has been depleted; (ii) the secondary sex trait is not particularly costly and therefore does not demonstrate condition dependence; or because (iii) the sex traits can be considered arbitrary traits rather than characters reflecting good genes.     

Effect of the hereditary characteristics of male blue foxes Alopex lagopus L. on the sex ratio of their offspring

Family analysis of a commercial population of the blue fox (the Pushkinskoe Breeding Fur Farm, Moscow oblast) with respect to secondary sex ratio has been performed. The offspring of each individual male or female involved in crossing between 1984 and 1988 was analyzed. The study of all families formed by every male and every female has made it possible to determine a group of "outstanding" fathers (23 out of 287 males), whose offspring was predominantly male (62.1% of the offspring were males, versus 53.9% in the total population). The results of subsequent detailed study on the pedigrees of male blue foxes in whose offspring the sex ratio significantly deviates from 1:1 indicate that this character is transmitted from fathers to sons without the deterioration of other commercially valuable characters. It is presumed that the significant deviation of sex ratio from 1:1 in the offspring of some male blue foxes is determined by genetic factors.     

Phylogenomic analysis resolves the relationships among net‐winged beetles (Coleoptera: Lycidae) and reveals the parallel evolution of morphological traits

Net‐winged beetles (Coleoptera: Lycidae) are a diverse group of elateroids known for aposematism and neoteny. Phylogenetic analyses of morphological and molecular data have revealed different results with respect to within‐group relationships. In this study, we recovered a highly supported phylogenomic phylogeny and identified seven subfamilies: Dexorinae stat.n. , Calochrominae stat.n. , Erotinae, Ateliinae, Lycinae, Lyropaeinae stat.n. and Metriorrhynchinae stat.n. Our results suggest that female neoteny evolved multiple times. Therefore, the development of similar morphological modifications in neotenics may be linked and may have produced characteristics such as body miniaturization, structural simplification, i.e. reduction of mouthparts, fewer antennomeres and palpomeres, uniquely shaped terminal palpomeres, shortened elytra, the loss of coadaptation between the elytra and pronotum, and others. Additional traits evolved in parallel due to similarities in biology, function and sexual selection. These characteristics include mimetic similarities, the presence of the rostrum, pronotal carinae and elytral costae, and the structure of male genitalia. By comparing the phylogenomic topology with the evolution of morphological characters, we were able to identify evolutionary trends in lycids and compare them with analogues for other neotenic elateroids. These traits have not been accepted as homoplasies due to the ambiguous phylogenetic signal from Sanger sequencing markers.     

Demographic aspects of interspecific hybridization

Summary A demographic study was carried out on two closely related species of the isopods Jaera (albifrons) ischiosetosa and J. (a.) albifrons and their F ₁ hybrids. The results from the 16 possible combinations of crosses have permitted an analysis of the nature of the mechanisms assuring the isolation of the species studied. Although intraspecific crosses yield an immediate success, interspecific crosses in the absence of choice of mates progress only slowly during the course of weeks. The results of both crosses between hybrids and back-crosses turn out to be intermediate between those of intra- and interspecific crosses. The hybrids of the first generation are perfectly viable and their survival curves are identical to those of the parents. The fertility of parents in intra- and interspecific crosses is comparable, with the exception of the fragility of female descendants (heterogametic sex) in one direction of crossing. The fertility of the F ₁ hybrids, however, crossed either among themselves or with their parents, is quite noticeably decreased: the time needed to double the size of the population is 2.5 times longer for the hybrids than for the parents. This hybrid breakdown completes the pre-fertilization isolating mechanisms: partial ecological isolation, and especially ethological isolation, is practically total when a choice of mates exists. The two species studied, for which demographic parameters are quite close, were raised together for ten generations and yielded only exceptional hybrids with a frequency which does not exceed that found under natural conditions.     

The cost of secondary sexual characters and the evolution of cost-reducing traits

Secondary sexual traits are characterized by their exaggerated expression relative to homologous nonsexual characters in other species. All models of sexual selection assume that sex traits are costly to produce and maintain, and individuals with reduced costs of production and maintenance of secondary sexual characters would be at a selective advantage. A number of morphological, physiological and behavioural traits may have evolved as a result of their cost-reducing properties: (1) body size, which does not change throughout life, that allows certain individuals to develop exaggerated sex traits, (2) cost-reducing traits, such as muscle size, that improve with practice and (3) actual cost-reducing traits, such as wing size in birds with song flight, which are produced in advance of or simultaneously with the sex trait. Cost-reducing traits may coevolve with secondary sexual characters and allow more extreme sexual signalling than would otherwise have been possible in their absence or in reduced versions.     

The cost of secondary sexual characters and the evolution of cost-reducing traits

Secondary sexual traits are characterized by their exaggerated expression relative to homologous nonsexual characters in other species. All models of sexual selection assume that sex traits are costly to produce and maintain, and individuals with reduced costs of production and maintenance of secondary sexual characters would be at a selective advantage. A number of morphological, physiological and behavioural traits may have evolved as a result of their cost-reducing properties: (1) body size, which does not change throughout life, that allows certain individuals to develop exaggerated sex traits, (2) cost-reducing traits, such as muscle size, that improve with practice and (3) actual cost-reducing traits, such as wing size in birds with song flight, which are produced in advance of or simultaneously with the sex trait. Cost-reducing traits may coevolve with secondary sexual characters and allow more extreme sexual signalling than would otherwise have been possible in their absence or in reduced versions.     

Intraspecific variation and intersexual correlation in secondary sexual characters of three diving beetles (Coleoptera: Dytiscidae)

Male diving beetles of the subfamily Dytiscinae possess tarsi with adhesive discs that they strike on the female dorsum during mating interactions. Females of many species are dimorphic, being either smooth or structured dorsally. Darwin suggested the female structures were an aid for the male but in this study we investigate these characteristics in the light of sexual conflicts. The intraspecific variation in the numbers and size distribution of male tarsal discs, and in body measurements were recorded for three dytiscine species, all with dimorphic females. The number of protarsal discs in the two Dytiscus species varied much more than previously reported. In addition, only a small part of the variation could be explained by body size. In Graphoderus we found highly significant differences in male secondary sexual characters among populations. A multivariate analysis significantly correlated male secondary sexual characters with the proportion of granulate females in the populations. These observations are consistent with the theory of arms races and female counter adaptations. Covariation between male and female characters is predicted from a framework of sexual conflict over mating rate. At the same time our study gives a new perspective on the function of dytiscine female dorsal irregularities debated ever since Darwin.