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.     

Location of SH-1 and SH-2 in the heavy chain segment of heavy meromyosin.

The two essential thiol groups of myosin, SH-1 and SH-2, have been localized in an ~ 20K segment of the heavy chain by analysis of the distribution of radioactivity after tryptic digestion of tryptic heavy meromyosin (HMM) or papain-HMM subfragment-1, both labeled at SH-1 and SH-2 with [¹⁴C]iodoacetamide and [¹⁴C]N-ethyl maleimide, respectively. The results are discussed in the framework of earlier work (Bálint, M., Sréter, F. A., Wolf, I., Nagy, B., and Gergely, J. (1975) J. Biol. Chem. 250, 6168–6177) on the tryptic fragmentation of myosin heavy chain and in the light of more recent work on the location of a fragment that reacts with a photoaffinity analog of ATP (Szilágyi, L., Bálint, M., Sréter, F. A., and Gergely, J. (1978) Fed. Proc. 37, 1695) and of suggestions concerning the binding of ATP in the region containing the SH-1 and SH-2 (Elzinga, M., and Collins, J. H. (1977) Proc. Nat. Acad. Sci. USA74, 4281–4284).     

Biogeography and systematics of Aricia butterflies (Lepidoptera,Lycaenidae)

Butterflies of the Aricia species group represent a paradigm of unresolved taxonomy, both at the genus and species levels. We studied phylogenetic relationships, biogeography, and systematics based on genetic – nuclear and mitochondrial – and morphometric – external (wings) and internal (genitalia) – data. We show that Aricia is a monophyletic genus comprising the taxa Pseudoaricia, Ultraaricia and Umpria, which are here considered junior synonyms of Aricia. The taxa allous, inhonora, issekutzi, mandzhuriana, myrmecias and transalaica, which have often been raised to species rank, are shown to probably represent subspecies or synonyms. We show that montensis is likely a good species that is sister to all A. artaxerxes populations across the Palearctic region. The species A. anteros and A. morronensis are shown to display deep intraspecific divergences and they may harbor cryptic species. We also discovered that A. cramera and A. agestis exhibit a pattern of mutual exclusion on islands, and a parapatric distribution in mainland with a narrow contact zone where potential hybrids were detected. The lack of a prezygotic barrier that prevents their coexistence could explain this phenomenon. This study will hopefully contribute to the stability of the systematics of Aricia, a group with potential for the study of the link between speciation and biogeography.     

Taxonomy of the genus Dianous (Coleoptera: Staphylinidae: Steninae) in China and zoogeographic patterns of its distribution

Abstract Ninety‐six species of the rove beetle genus Dianous Leach, 1819 have been recorded in China. In this paper, we describe a new species, D. poecilus n. sp. from Yunnan, and record D. viriditinctus ( Champion, 1920 ) for the first time from China. To facilitate identification of species, species groups and/or species complexes, we compiled a new version of keys, including all species of Dianous hitherto recorded in the territory of China. Moreover, main patterns of geographical distribution are outlined for the world Dianous fauna of this genus.     

Two new species of the ground-beetle genus <Emphasis Type="Italic">Harpalus</Emphasis> Latr. (Coleoptera,Carabidae) from Turkey and China

Two new species of the genus Harpalus Latreille, 1802 are described: H. stevensi sp. n. from China (Sichuan) and H. dudkoi sp. n. from Turkey (Adana). Both species belong to the nominotypical subgenus and are included in the tardus species-group.     

Comparative chiasma analysis using a computerised optical digitiser

A new computerised technique has been devised for measuring the distribution of chiasmata along diplotene bivalents. The method involves the introduction into the field of view of the microscope, of a fine light spot which can be accurately manipulated along the chromosomes of each bivalent. The data recorded include (a) the positions of the chiasmata along the bivalent in terms of their relative distances from the centromere and (b) the individual bivalent and cellular chiasma frequencies. — The method has been applied to the analysis of chiasma distribution patterns in the two known species of the genus Caledia, C. species nova 1 and C. captiva and in two chromosomal races of the latter. Statistical tests indicate that within bivalents at least 40% of the comparative distribution patterns of chiasmata between races and species are significantly different. Similar comparisons between populations within races reveal only 18% significant differences. — The observed distribution patterns of chiasmata in this genus suggest that chiasma formation is sequential from centromere to telomere. — The variation in the frequency and distribution of chiasmata between races and species suggests that the interference distances between successive chiasmata are, at least partially, independent of chiasma frequency and position. — The interracial and interspecific differences in chromosome structure are correlated with changes in chiasma pattern.     

A new genus and species of monostiliferous hoplonemertean (Nemertea: Enopla: Monostilifera) from Japan

A new genus and species of monostiliferous hoplonemertean, Diopsonemertes acanthocephala gen. et sp. nov., is described from Otsuchi Bay, Japan. Significant anatomical features of the new form include a body wall longitudinal musculature anteriorly divided into inner and outer layers by connective tissue, no pre-cerebral septum, the presence of a thin coat of diagonal muscle fibres between the body wall longitudinal and circular muscle layers in the foregut body region, cephalic retractor muscles derived only from the inner portion of the divided longitudinal muscles and a rhynchocoel more than half the body length.     

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.     

Boromorphus saudicus sp. n., a new record of the genus from the Arabian Peninsula (Tenebrionidae: Pimeliinae: Boromorphini)

Boromophus saudicus sp. n. (subfamily Pimeliinae Latreille, 1802, tribe Boromorphini Skopin, 1978) is described from the Kingdom of Saudi Arabia (Rawdhat Khorim, Riyadh Province). This finding represents the first record of the genus Boromorphus Wollaston, 1854 from the Arabian Peninsula. The species was collected from Acacia trees and from pitfall traps.     

On Podocotyle scorpaenae,Poracanthium furcatum and Derogenes latus,three poorly known digenean parasites of western Mediterranean teleosts

Three little-known digeneans, two opecoelids and a derogenid, from marine fishes off Corsica are redescribed and their systematics discussed. Distoma scorpaenae Rudolphi, 1819, from Scorpaena porcus, S. scrofa and S. notata, is transferred to the genus Podocotyle as P. scorpaenae (Rudolphi, 1819) n. comb. The authority for the taxon Poracanthium furcatum is amended to Dollfus (1948). This species is described from Mullus surmuletus and distinguished from Opecoeloides furcatus (Bremser in Rudolphi, 1819). Derogenes latus Janiszewska, 1953 is redescribed from Mullus surmuletus and discussed in relation to D. ruber Lühe, 1900.     

Musculature of the penial complex: A new criterion in unravelling the phylogeny of Hygrophila (Gastropoda: Pulmonata)

The taxonomy of freshwater pulmonates (Hygrophila) has been in a fluid state warranting the search for new morphological criteria that may show congruence with molecular phylogenetic data. We examined the muscle arrangement in the penial complex (penis and penis sheath) of most major groups of freshwater pulmonates to explore to which extent the copulatory musculature can serve as a source of phylogenetic information for Hygrophila. The penises of Acroloxus lacustris (Acroloxidae), Radix auricularia (Lymnaeidae), and Physella acuta (Physidae) posses inner and outer layers of circular muscles and an intermediate layer of longitudinal muscles. The inner and outer muscle layers in the penis of Biomphalaria glabrata consist of circular muscles, but this species has two intermediate longitudinal layers separated by a lacunar space, which is crossed by radial and transverse fibers. The muscular wall of the penis of Planorbella duryi is composed of transverse and longitudinal fibers, with circular muscles as the outer layer. In Planorbidae, the penial musculature consists of inner and outer layers of longitudinal muscles and an intermediate layer of radial muscles. The penis sheath shows more variation in muscle patterns: its muscular wall has two layers in A. lacustris, P. acuta, and P. duryi, three layers in R. auricularia and Planorbinae and four layers in B. glabrata. To trace the evolution of the penial musculature, we mapped the muscle characters on a molecular phylogeny constructed from the concatenated 18S and mtCOI data set. The most convincing synapomorphies were found for Planorbinae (inner and outer penis layers of longitudinal muscles, three-layered wall of the penis sheath). A larger clade coinciding with Planorbidae is defined by the presence of radial muscles and two longitudinal layers in the penis. The comparative analysis of the penial musculature appears to be a promising tool in unraveling the phylogeny of Hygrophila.     

A combined genetic-morphometric analysis unravels the complex biogeographical history of Polyommatus icarus and Polyommatus celina common blue butterflies

Widespread species have the potential to reveal large‐scale biogeographical patterns, as well as responses to environmental changes possibly unique to habitat generalists. This study presents a continental‐scale phylogeographical analysis of Polyommatus icarus, one of the most common Palaearctic butterflies, and the morphologically and ecologically similar Polyommatus celina, a recently discovered cryptic species. By combining data from mitochondrial [cytochrome c oxidase subunit I (COI)] and nuclear [internal transcribed spacer (ITS2)] molecular markers with geometric morphometrics, we document a complex phylogeographical history for the two species. Despite morphological similarities, the genetic divergence between these two species is high (more than 5% at COI) and they are not sister species. For the first time, we show that P. celina occurs not only in North Africa but also in Europe, where it inhabits several west Mediterranean islands, as well as large parts of Iberia, where it occurs in parapatry with P. icarus. The two species appear to completely exclude each other on islands, but we provide morphological and molecular evidence that introgression occurred in the Iberian Peninsula. We discovered strongly diverged lineages that seem to represent relict populations produced by past range expansions and contractions: Crete and Iberian isolates for P. icarus, Balearics–Sardinia and Sicily–Lipari for P. celina. This study shows that a combined genetic‐morphometric approach can shed light on cryptic diversity while providing the necessary resolution to reconstruct a fine‐scale phylogeographical history of species at both spatial and temporal levels.     

A review of the genus Chironomus (Diptera,Chironomidae)

Analysis of the banding pattern of the salivary gland chromosomes of Chironomus tepperi indicates that, despite a somewhat modified male hypopygium, the relationships of this species are close to the other Australian species of the genus, particularly to Ch. oppositus. No inversion polymorphism has been found in Ch. tepperi and this, together with the relatively high chiasma frequency as measured at metaphase I, would appear to be an adaptation to provide genetic variability necessary for its colonizing ability.     

Muscle fibers in the central nervous system of nemerteans: Spatial organization and functional role

The system of muscle fibers associated with the brain and lateral nerve cords is present in all major groups of enoplan nemerteans. Unfortunately, very little is known about the functional role and spatial arrangement of these muscles of the central nervous system. This article examines the architecture of the musculature of the central nervous system in two species of monostiliferous nemerteans (Emplectonema gracile and Tetrastemma cf. candidum) using phalloidin staining and confocal microscopy. The article also briefly discusses the body‐wall musculature and the muscles of the cephalic region. In both species, the lateral nerve cords possess two pairs of cardinal muscles that run the length of the nerve cords and pass through the ventral cerebral ganglia. A system of peripheral muscles forms a meshwork around the lateral nerve cords in E. gracile. The actin‐rich processes that ramify within the nerve cords in E. gracile (transverse fibers) might represent a separate population of glia‐like cells or sarcoplasmic projections of the peripheral muscles of the central nervous system. The lateral nerve cords in T. cf. candidum lack peripheral muscles but have muscles similar in their position and orientation to the transverse fibers. The musculature of the central nervous system is hypothesized to function as a support system for the lateral nerve cords and brain, preventing rupturing and herniation of the nervous tissue during locomotion. The occurrence of muscles of the central nervous system in nemerteans and other groups and their possible relevance in taxonomy are discussed. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Rotifer muscles as revealed by phalloidin-TRITC staining and confocal scanning laser microscopy

By combining phalloidin‐TRITC staining with confocal scanning laser microscopy (CSLM), the pattern of the musculature in two species of Rotifera, Euchlanis dilatata unisetata and Brachionus quadridentatus is revealed. The same general muscle pattern prevails in both species. The major components of the body wall musculature are: 1. retractor muscles (5 pairs in E. dilatata unisetata and 3 pairs in B. quadridentatus); 2. Two pairs of dorso‐ventral muscles; 3. Two pairs of perpendicular muscles (in E. dilatata unisetata); 4. retractors of the corona (median, lateral and ventral); 5. Foot retractors. In addition, three pairs of cutaneo‐visceral muscles and visceral muscles (including mastax muscles) are described. The sphincter of the corona was found only in B. quadridentatus. The high degree of muscle differentiation points to a high level of development of rotifer muscular system.     

Comparative myology of the forelimb of Liolaemus sand lizards (Liolaemidae)

The lizard genus Liolaemus includes numerous constituent clusters of putatively related taxa, one of which is the Liolaemus boulengeri group, which in turn includes the sand lizards (of the Liolaemus wiegmannii subgroup). Members of the sand lizard group exhibit three different modes of burying into sand. The general morphology of the forelimb muscles of those Liolaemus species is analysed. Herein, we present a study of the forelimb musculature of all species considered by Halloy et al. (1998). This study has three principal goals. First, we are seeking myological characters that will be useful in formulating phylogenetic hypothesis about the species of Liolaemus. With these characters, we also wish to compile morphological data that represent the morphological space implied in the diverse locomotor behaviours of these animals. Second, we are looking for derived features that reflect functional changes in the use of forelimb. Third, we wish to provide a cladistic analysis that can be used to test phylogenetic hypothesis derived from other sources of data. We present 48 characters in a data set and analyse it cladistically. We obtained a hypothesis of relationships of the Liolaemus species and compared this with previous hypotheses based on other characters. The trees obtained are not congruent with previously proposed phylogenies. We were unable to identify in our trees nodes that are based on structures reflecting functional changes in the use of the forelimb. The morphological similarities in the forelimb musculature of all species analysed seems to conform a very conservative general anatomical pattern with which Liolaemus sand lizards perform most of their locomotor behaviours.     

Revision of the genus Amadotrogus Reitter, 1902 (n. stat.) (Coleoptera: Scarabaeidae: Melolonthinae)

In this paper, the subgenus Amadotrogus Reitter, 1902 and its taxonomic status as a subordinate taxon of genus Amphimallon are reviewed. Two kinds of character sets are discussed; those relative to the external morphology of the adult and those of the male and female genitalia. Subgenus Amadotrogus is compared with its recognized allied genera in the same clade (Coca-Abia 1995) Amphimallon Berthold, 1827; Geotrogus Guérin, 1842; Monotropus Erichson, 1847; Pseudoapeterogyna Escalera, 1914 and Rhizotrogus Berthold, 1827 to asses the realtionships of these taxa. Phylogenetic analysis discloses that Amadotrogus, considered to be a subgenus of Amphimallon, has synapomorphic characters which justify its elevation to generic rank. Thus, genus Amadotrogus includes seven species distributed across the Northern Mediterranean basin. The type species, Amadotrogus quercanus (Burmeister, 1855), and six others, transferred from genus Rhizotrogus: Amadotrogus grassii (Mainardi, 1902), Amadotrogus insubricus (Burmeister, 1855), Amadotrogus oertzeni (Brenske, 1886), Amadotrogus patru- elis (Reiche, 1862), Amadotrougs truncatus (Brenske, 1886) and Amadotrogus vicinus (Mulsant, 1842) (Coca-Abia & Martín-Piera 1998). In addition, Rhizotrogus rugifrons Burmeister, 1855 is considered a new synonym of Amadotrogus vicinus. Rhizotrogus bolivari Martínez y Sáez, 1873, Amphimallon cata- launicum Báguena, 1956 and Rhizotrogus lajonquierei Baraud, 1970 are synonymized with Amadotrogus patruelis.