Glands, muscles and genitalia. Morphological and phylogenetic implications of histological characters in the male genitalia of Fritillary butterflies (Lepidoptera: Nymphalidae: Argynnini)

The histology and internal morphology of the male valve in a broad sample of Fritillary butterflies are examined. A large internal ridge to which the extensor of the valve muscle fastens is probably an autapomorphy for the tribe Argynnini. A large glandular pocket is found in the valves of all examined Heliconiinae, but is missing in other examined Nymphalidae. The pocket shows considerable variation between the higher taxa. In the higher Fritillaries (subtribe Argynnina) the pocket probably has an effective closing mechanism allowing the male to store a considerable amount of glandular secretion. It has earlier been suggested that the secreta from the pocket glands are transferred to the female during copulation, where they probably have an antiaphrodisiac effect, preventing other males from mating a once mated female. All examined male Fritillaries also have glands in the anellus, the terminal membrane between the valves. In Boloria these glands are extremely well developed and abundant, a condition that is probably an autapomorphy for that genus. The obtained characters are tested against a known phylogeny of the tribe Argynnini.     

楔天牛属7种雄性外生殖器特征比较研究

本文研究了检疫截获的楔天牛属7种楔天牛八点楔天牛、白桦楔天牛、斑点楔天牛、柳楔天牛、青杨楔天牛、山杨楔天牛、十星楔天牛雄性外生殖器的形态结构,并提供其解剖图。研究表明,楔天牛属雄性外生殖器由阳茎基、中茎和内囊3部分组成。阳茎基的阳基侧突具有相对遗传稳定性,且在属内均成近似矩形,并首次测量了7种楔天牛的成虫雄性外生殖器阳基侧突的长宽比。得出了雄性外生殖器形态特征在楔天牛属种内差异小、种间差异大,并能用雄性外生殖器特征有效区分楔天牛属近似种的结论。     

Variation in the genitalia of the butterfly Maniola jurtina (Lepidoptera: Satyrinae)

The genilalia of animals, particularly insects, are extremely useful taxonomic characters, for they are highly conserved within species yet generally diverge rapidly during specialion: often the only reliable means of separating sibling species is via the morphology of the genilalia. A number of explanations have been proposed to explain this pattern of variation, the most prominent of which is the 'lock-and-key' hypothesis. This hypothesis and others are tested using the meadow brown butterfly, Maniola jurtina (L.), which exhibits variation in the male genitalia. A novel technique is described which enables dissection of copulating couples. Sampling and dissection of male butterflies from 14 populations in southern England was carried out to quantify variation in the genitalia. Mating success and the strength of the male-female bond during copulation was assessed in relation to the dimensions of the male genitalia.
The most variable portions of the male genitalia are the distal and dorsal margins of the paired valves. Contrary to the lock-and-key hypothesis, and the more recent sexual selection model, evidence is presented that the most variable portions of the genitalia have no apparent function during copulation, and therefore may not be subject to direct selection. I suggest that neutrality to selection was essential for the evolution of the wide variety in genital morphology currently found in the Lepidoptera.     

Comparison of phylogenetic signal between male genitalia and non-genital characters in insect systematics

It is generally accepted that male genitalia evolve more rapidly and divergently relative to non-genital traits due to sexual selection, but there is little quantitative comparison of the pattern of evolution between these character sets. Moreover, despite the fact that genitalia are still among the most widely used characters in insect systematics, there is an idea that the rate of evolution is too rapid for genital characters to be useful in forming clades. Based on standard measures of fit used in cladistic analyses, we compare levels of homoplasy and synapomorphy between genital and non-genital characters of published data sets and demonstrate that phylogenetic signal between these two character sets is statistically similar. This pattern is found consistently across different insect orders at different taxonomic hierarchical levels. We argue that the fact that male genitalia are under sexual selection and thus diverge rapidly does not necessarily equate with the lack of phylogenetic signal, because characters that evolve by descent with modification make appropriate characters for a phylogenetic analysis, regardless of the rate of evolution. We conclude that male genitalia are a composite character consisting of different components diverging separately, which make them ideal characters for phylogenetic analyses, providing information for resolving varying levels of hierarchy.
© The Willi Hennig Society 2009.     

Evidence for lock-and-key mechanisms in the internal genitalia of the Apamea moths (Lepidoptera, Noctuidae)

Abstract. Fifty of the fifty-six species of the genus Apamea known from North America and three Palaearctic species were analysed for lock-and-key characters in their internal genitalia, mainly in the male vesica and the female bursa copulatrix. There were an average of 4.5 such characters per species, structurally corresponding in the two sexes. Anatomically they form a postcopulatory but prezygotic isolation mechanism. In some closely related species, the internal genitalia are very similar, but these species have a precopulatory isolation mechanism in the presence or absence of male abdominal coremata. Closely related species did not have more lock-and-key characters than unrelated species, which is taken to indicate absence of character displacement. The anatomical distribution of the lock-and-key characters was examined and the organs of eight species are illustrated.
The lock-and-key hypothesis has been abandoned by several earlier authors but mainly on consideration of external genitalia. In Apamea the invariable functional correspondence between sexes in the sperm transferring organs, and the overall species-specifity of characters but non-existence of interspecific differences under a precopulatory mechanism indicate that (a) lock-and-keys are functioning and (b) they act as isolation mechanisms. Alternative hypotheses of genitalic evolution are reviewed.     

Female genitalia of Pero obtusaria Prout, 1928 (Lepidoptera: Geometridae)

The female genitalia of Pero obtusaria Prout, 1928 (Lepidoptera: Geometridae) are described and illustrated for the first time and compared to congenerics. The antrum with the dorsal part sculptured with two sinuous longitudinal stripes enables the identification of this species.     

鳞翅目昆虫化石研究进展

迄今为止,世界古昆虫学家共发现鳞翅目昆虫化石42科、176属、254种。本文回顾了鳞翅目化石研究简史,通过图表列举了鳞翅目化石昆虫名录及其分布和年代,概述了鳞翅目昆虫的地质历史、分类系统和系统发育,初步分析了鳞翅目昆虫化石研究中存在的问题,并对今后的研究前景进行了展望。     

Congruence between comparative morphology and molecular phylogenies: the final stage of evolution of the skeletal characters of male genitalia in the subtribe Polyommatina (Lepidoptera,Lycaenidae). Part 2. Uncus and subunci

The morphology of the male terminalia (uncus and subunci) was studied in 39 species of the subtribe Polyommatina (Lycaenidae). Traces of nine established characters were mapped on the molecular phylogeny of the subtribe proposed by Talavera et al. (2013). The ancestral character states for these morphological structures were established in Plebejus s. l. and Polyommatus s. l. According to the time calibration of the proposed molecular tree, the studied skeletal structures (uncus and subunci) evolved very slowly within the last million years. It was shown that the similar shape of the uncus and subuncus branches arose independently and underwent parallel morphological transformations. In particular, wide and, as a rule, ventrally directed uncus lobes arose independently in Polyommatus s. str., Alpherakya, Glabroculus, and Aricia. Yet, in various evolutionary lines of Polyommatina similar changes in the length ratio of the proximal and distal parts of the subunci as well as in the shape of their apical hooks were observed. The character states marking the groups Polyommatus s. str. and Aricia were established. The evolutionary changes in the uncus morphology were highly non-uniform in the various monophyletic lines of Polyommatina. As a result, representatives of some “old” lines have more specialized features of the uncus morphology in comparison with those belonging to “young” lines that retain ancestral character states. In Polyommatina, the transformation of the male terminalia was directed toward development of the most effective mechanism of female fixation during copulation. This goal was reached in similar ways in various evolutionary lines of Polyommatina, which explains numerous parallelisms in morphology of these structures.     

Taxonomy of Selaginella: a study of characters, techniques, and classification in the Hong Kong species

This study of 11 Hung Kong species of Selaginella indicates that sufficient characters exist by which they may be conveniently, consistently, and conclusively distinguished, and implies that the same is true for the rest of the genus, which suffers from chronic taxonomic confusion. Characters assessed are: leaf size, shape, epidermal cell patterns and mesophyll structure; ligule shape; patterns seen in stem transections; sporophyll shape; microsporangium shape; sporangial distribution; and spore surface ornamentation. Of these, the shape of median leaves, the maximum size of lateral leaves and epidermal cell patterns are the most significant and distinctive foliar features. These, together with spore surface ornamentation, are the most useful for distinguishing species. Correlations between the shapes of ligules, sporophylls, microsporangia, and vascular bundles indicate natural subgeneric groups.
These studies suggest that there are two ways by which taxonomic investigations of Selaginella can be improved. First leaves and spores should be prepared in permanent mountant (e.g. Hoyer's solution) for accurate observation of size, shape, and epidermal patterns. Secondly, data should be collected on a wide range of features.     

An illustrated guide to the identification of the known species of Diatraea Guilding (Lepidoptera,Crambidae, Crambinae) based on genitalia

The genus Diatraea Guilding is one of the most economically important groups of moths in the Western Hemisphere. The larvae are stem borers that feed on species of Poaceae, or grasses, such as sugarcane, corn, rice, and sorghum, as well as many other native grasses. Interest in this group has risen considerably since sugarcane and other grasses have been utilized and/or investigated as biofuels. This is the first modern study to treat all 41 valid described species. Most type specimens were examined and we provide a checklist with 19 new synonyms. We provide keys for the identification of most species in this genus based on morphology of the male and female genitalia and modern illustrations of male and female genitalia. We also provide an updated table of species distribution by country.     

Inter- and intraspecific variation in the genitalia of the ‘Melitaea phoebe group’ (Lepidoptera, Nymphalidae)

The ecology and phylogeny of the Melitaeini butterflies have received considerable attention in the last few years. Several publications have been based on research using molecular methods. Genital morphometry has already been published on two sibling species but without any accompanying taxonomic revision of related species or their intraspecific taxonomy. In this study the morphology of genitalia was analysed by landmark-based geometric morphometrics in males and by traditional morphometrics in females. Based on these morphological characters, the whole M. phoebe group (subg. Cinclidia) was revised. According to our results M. telona Fruhstorfer, 1908 is conspecific with M. ‘phoebe’ ornata Christoph, 1893. Thus, we consider M. telona a junior synonym of M. ornata, and we propose new combinations: M. ornata ornata Christoph, 1893, M. ornata telona Fruhstorfer, 1908, M. ornata amanica Rebel, 1917, M. ornata capreola Varga, 1967, M. ornata emipunica Verity, 1919, M. ornata enoch Higgins, 1941, M. ornata kovacsi Varga, 1967, M. ornata nigrogygia Verity, 1938, M. ornata ogygia Fruhstorfer, 1907 and M. ornata totila Stauder, 1914 based on the rule of priority. The validity of the species status of M. punica and M. scotosia was confirmed, whereas M. sibina and M. pseudosibina could not be separated from M. phoebe using morphometrics. A new species from Iran (Zagros Mts.) was described as M. zagrosi sp. n. In addition, the whole geographical range of M. ornata was reconsidered and some biogeographical connections were discussed. In M. ornata four morphotypes were found based on genitalia: 1. ‘Western type’: including the Italian, Hungarian and SW Ukrainian populations; 2. ‘Eastern type’: with a much extended distribution ranging from the Balkan Peninsula through Asia Minor to Kazakhstan; 3. ‘The Levant type’ (=M. ornata telona comb. nov.); 4. ‘Northern Iranian type’ (Kopeth Dagh and Elburz Mts.). The distribution of these morphotypes can be attributed to historic separation in distinct glacial refugia.     

The morphological and histological characters of the male external genitalia of the house musk shrew, Suncus murinus

External genitalia are the reproductive organs necessary for efficient copulation and internal fertilization in various mammalian species. Their morphogeneses display significant morphological and developmental differences among species. The house musk shrew, Suncus murinus (hereafter described as suncus) is a species of the order Insectivora, which has been considered as primitive and one of the earliest eutheria phylogenetically. Comparative anatomical analyses of phylogenetically different mammals will contribute to the better understanding of morphological diversity of external genitalia. This study performed various anatomical and histological analyses concerning the organization of the external genitalia of male suncus. It was shown that the external genitalia of suncus possessed a muscular structure, which we proposed as musculus ischiocavernosus dorsalis of suncus. The musculus ischiocavernosus dorsalis is originated from the inner surface of the tuber ischiadicum and was allocated adjacent to the corpus cavernosum penis. In addition, a pair of alpha-smooth muscle actin positive muscles was located bilaterally to the urethra. This unique morphology of the external genitalia of suncus males may provide a unique model system to investigate genital morphogenesis.     

Effect of husk characters on resistance to corn earworm (Lepidoptera: Noctuidae) in high-maysin maize populations

Two maize (Zea mays L.) breeding populations with very high concentrations of maysin, a silk-expressed flavone glycoside, were tested for their ability to resist ear damage by the corn earworm, Helicoverpa zea Boddie, under field conditions. Tests were conducted in 2000 and 2001 at multiple locations in Georgia. The high maysin populations, EPM6 and SIM6, as well as resistant and susceptible checks, were scored for silk-maysin content, H. zea damage, and husk characters. In 2000, there was a negative correlation between husk tightness and earworm damage at three of five locations, while there was no significant correlation between damage and maysin content at any location. In 2001, EPM6 and SIM6 had approximately ten times the maysin content of the low-maysin control genotypes; nevertheless, earworm damage to EPM6 and SIM6 was either greater than or not significantly different from the low-maysin genotypes at all locations. The resistant control genotype, Zapalote Chico, had significantly less earworm damage than EPM6 and SIM6 for both years at all locations. The results of this study highlight the importance of identifying and quantifying husk and ear traits that are essential to H. zea resistance in maize.     

Alternative respiratory oxidases to study the animal electron transport chain

Oxidative phosphorylation is a common process to most organisms in which the main function is to generate an electrochemical gradient across the inner mitochondrial membrane (IMM) and to make energy available to the cell. However, plants, many fungi and some animals maintain non-energy conserving oxidases which serve as a bypass to coupled respiration. Namely, the alternative NADH:ubiquinone oxidoreductase NDI1, present in the complex I (CI)-lacking Saccharomyces cerevisiae, and the alternative oxidase, ubiquinol:oxygen oxidoreductase AOX, present in many organisms across different kingdoms. In the last few years, these alternative oxidases have been used to dissect previously indivisible processes in bioenergetics and have helped to discover, understand, and corroborate important processes in mitochondria. Here, we review how the use of alternative oxidases have contributed to the knowledge in CI stability, bioenergetics, redox biology, and the implications of their use in current and future research.     

A new approach to the study of genetic variability of complex characters

A new approach to multivariate genetic analysis of complex organismal traits is developed. It is based on examination of the distribution of parental strains and the F1 and F2 hybrids in a multidimensional space, and the determination of the directions corresponding to heterozygosity, epistatic and additive gene effects. The effect of heterozygosity includes variability produced by interaction between and within heterozygous loci. The additive gene effects and the remaining epistatic interactions between the homozygous loci can be visualized separately from the effects of heterozygosity by an appropriate projection of the points in multidimensional space. In all, 20 morphological, physiological and behavioural characters and 21 craniometric measures were studied in crosses between two laboratory rat strains. Linear combinations of craniometric and of morphophysiological characters with a high narrow-sense heritability could be identified. These combinations characterized the organismal stress response, which had been selected for in one of the strains. The prospects for the practical application of the new approach and also for the evaluation of the contribution of the genetic diversity to phenotypic variability in animals in natural populations are discussed.     

广州地区女性下生殖道感染致病微生物分析

目的 对广州地区女性下生殖道感染致病微生物状况进行调查,用以指导临床正确诊断治疗。方法 对以外阴瘙痒及自带增多为主诉的妇女患者进行临床检查,先取阴道分泌物进行pH、胺试验及假丝酵母菌、滴虫和线索细胞检查,同时采用特制密封棉拭子宫颈管内取样,然后采用重量差减法,求出分泌物的重量,采用荧光定量PCR测定解脲脲原体、淋球菌、沙眼衣原体,可疑部位取材作HSV和HPV检测及梅毒血清学检查。结果 在533例入选患者中,解脲脲原体感染345例占64．7％,沙眼衣原体感染142例占26．6％,阴道假丝酵母菌病117例占22％,非特异性阴道炎81例占15％,细菌性阴道病17例占0．03％。尖锐湿疣12例占0．02％。淋病5例,梅毒4例,生殖器疱疹2例,滴虫阴道炎2例。结论 解脲脲原体、沙眼衣原体、阴道假丝酵母菌病是当前广州地区女性下生殖道感染的主要致病微生物,在选用抗生素时要兼顾解脲脲原体、沙眼衣原体。     

Congruence between comparative morphology and molecular phylogenies: the final stage of evolution of the skeletal characters of male genitalia in the subtribe polyommatina (Lepidoptera,Lycaenidae). Part 1. Valvae (distal part), aedeagus,and juxta

The evolution of the aedeagus, juxta, and microstructures on the valvae in the subtribe Polyommatina (Lycaenidae) during the last 5 MY is considered, and the chronology of morphological changes in these structures is established. The morphological characters of 47 species of the subtribe were tested by molecular analysis. Numerous cases of parallel evolution were discovered in the taxa of various taxonomic levels within Polyommatina. The principal shapes of the aedeagus apex developed independently in different clades. Multiple independent cases of an increase in the number of marginal denticles on the dorsal valvar clasps were observed in the genera Polyommatus, Plebejus, and Kretania. Parallel cases of a decrease in the number of these denticles were observed less frequently, in Aricia, Plebejus (P. argus), and Agriades (A. glandon). The juxta evolved from V- to U-shaped almost in all the clades. The rates of evolution of microcuticular and skeletal structures were shown to be different. For example, the number of cuticular denticles may change in 1 MY, while changes in the functionally important cuticular structures took as long as 2–3.5 MY. However, some structural changes were relatively fast; for instance, the club-shaped apex of the aedeagus was modified into a more typical conical form in less than 1 MY in species of the genus Agriades. Skeletal changes occurred in different moments of the evolutionary time frames of monophyletic species groups.     

Evolution of the skeleton and musculature of the male genitalia in the family lycaenidae (Lepidoptera): II. Infratribe Polyommatina Swainson, 1827

Skeleton and musculature of valvae were examined in 19 species from the section Polyommatus (Lepidoptera, Lycaenidae) revised by Bálint and Johnson (1997). Functional organization of male genitalia in the section Polyommatus was shown to be relatively constant. Significant interspecific variation of the skeleton and muscles was found only in the structure of valvae. During their evolution, the valvae became elongated, the costal torsion and longitudinal medial fold appeared on their medial wall, the subcostal groove was formed, and the intravalval muscle differentiated into two muscles forming a chiasma. This muscular chiasma was formed at least twice in the evolution of the group, in Polyommatus Latreille, 1804 + Aricia [Reichenbach], 1819 and in Plebeius Kluk, 1802. The genus Chilades Moore, [1881] is the least advanced as concerns the functional organization of the valvae. The genera Polyommatus and Aricia are the most advanced and constitute a sister pair. The intermediate position is occupied by Plebeius Kluk, 1802 and probably Madeleinea Bálint, 1993. The genus Albulina Tutt, 1809 is a compound taxon, its species occupying an intermediate position between Polyommatus and Aricia on the one hand, and Plebeius on the other hand. With a sole exception, they are characterized by a plesiomorphic state of the valval musculature that does not allow their mutual phylogenetic relations to be evaluated consistently. The only examined species of the genus Agriades Hübner, [1819], A. glandon (De Prunner, 1798), also possesses undifferentiated musculature, but according to the characters of the vaginal area, its females are close to the branch Polyommatus + Aricia.