ERRATUM. Because of an unfortunate error,the index to Volume 32 as it appeared in Volume 32, 4 is incorrect. The correct Index to Volume 32 appears at the back of this issue,Volume 33, 1.

Most Hepialidae have a single-layer wing covering of hollow scales. In the South American genera Callipielus and Dalaca, however, a very characteristic ‘type-2’ bilayer wing scale covering is present. As only Callipielus and Dalaca among 28 investigated hepialid genera have this type of wing scale covering, the two genera may be considered sister taxa.     

Comparative morphology and evolutionary aspects of the reflective under wing scale-pattern in Fritillary butterflies (Nymphalidae: Argynnini)

The wing scale ultrastructure of the reflective under wing pattern found in many Argynnini butterflies are examined for all recognised genera and subgenera and compared to that of some basal Heliconiini. A true reflective pattern probably evolved once within the Argynnini. But the phylogenetic information in these structures is limited due to a high degree of homoplasy in the scale ultrastructure related to the reflective patterns. The degree of specialisation is also homoplastic. The general morphological modification responsible for the reflective patterns seems to be a “closing” of the large windows, which generally occupy most of the inter-ridge space on the abwing surface in the scales of higher Lepidoptera. The fact that the Argynnis niobe morphs with a silvery pattern have scales with ‘closed windows’ whereas the Argynnis niobe morphs without a silvery pattern have typical non-reflective scales with very large windows supports this conclusion.The degrees of modification of the scales, including whether both cover and ground scales or only cover scales are modified, are to some extent correlated to the degree of reflectiveness in the wing pattern. Boloria eunomia has, as the only species, more modified ground scales than cover scales.     

Moth body size increases with elevation along a complete tropical elevational gradient for two hyperdiverse clades

The body size of an animal is probably its most important functional trait. For arthropods, environmental drivers of body size variation are still poorly documented and understood, especially in tropical regions. We use a unique dataset for two species‐rich, phylogenetically independent moth taxa (Lepidoptera: Geometridae; Arctiinae), collected along an extensive tropical elevational gradient in Costa Rica, to investigate the correlates and possible causes of body‐size variation. We studied 15 047 specimens (794 species) of Geometridae and 4167 specimens (308 species) of Arctiinae to test the following hypotheses: 1) body size increases with decreasing ambient temperature, as predicted by the temperature–size rule; 2) body size increases with increasing rainfall and primary productivity, as predicted from considerations of starvation resistance; and 3) body size scales allometrically with wing area, as elevation increases, such that wing loading (the ratio of body size to wing area) decreases with increasing elevation to compensate for lower air density. To test these hypotheses, we examined forewing length as a proxy for body size in relation to ambient temperature, rainfall, vegetation index and elevation as explanatory variables in linear and polynomial spatial regression models. We analysed our data separately for males and females using two principal approaches: mean forewing length of species at each site, and mean forewing length of complete local assemblages, weighted by abundance. Body size consistently increased with elevation in both taxa, both approaches, both sexes, and also within species. Temperature was the best predictor for this pattern (–0.98 < r < –0.74), whereas body size was uncorrelated or weakly correlated with rainfall and enhanced vegetation index. Wing loading increased with elevation. Our results support the temperature–size rule as an important mechanism for body size variation in arthropods along tropical elevational gradients, whereas starvation resistance and optimization of flight mechanics seem to be of minor importance.     

Ecological implications of the correlation of avian footprints with wing characteristics: a mathematical approach

The characteristics of avian wings that evolved for flying appear to show a distinct relationship to the shape of the pes and walking abilities as reflected in footprints. Wing area, wing span and body weight data of modern birds were collected and analysed in order to quantify the possible correlation, which was previously only inferred from empirical data. Discriminant analysis demonstrated that avian wings can be divided into three habitat groups, in a similar way to footprints. Multiple regression analyses revealed that the avian wing loading and aspect ratio were correlated with the parameters of footprint shape and can be expressed by a simple equation. The results may reflect the adaptation of avian locomotion to habitat. The relationships between wing area and wing span, and between wing area and footprint area, which are apparent in modern avians, were derived and used to estimate wing area and wing span from the footprints of extinct Cretaceous avian taxa. The estimated values of body weight, wing span and wing area suggest that the trackmakers of Archaeornithipus meijidei, Hwangsanipes choughi and Yacoraitichnus avis had bodies similar to herons (or cranes), large sandpipers (or small sea birds) and medium‐sized gull‐like birds, respectively.     

Carotenoids and bile pigments in Danaid and swallowtail butterflies

The carotenoids found in adult kite swallowtails ( Graphium spp.) and the Danaid glassy tigers ( Ideopsis, Tirumala and Parantica ), butterflies with exposed blue/green pigmented wing membranes (which contain pterobilins) and a modified scale vestiture, were compared with those of related species of Papilios and Danaids with concealed wing membranes an an entire scale vestiture. It was found that both the kite swallowtails contained only lutein, and even lacked β-carotene, the first known instance among butterflies. This was presumably due to selective storage by the larva since the food-plant was found to be rich in varied carotenoids. A possible link between the high concentration of pterobilins in the wing membrane and the lack of carotenoids is discussed.     

Microsculpture of the wing surface in Odonata: evidence for cuticular wax covering

The insect wing membrane is usually covered by scales, hairs, and acanthae, which serve diverse functions, such as species-specific coloration pattern, decrease of wind resistance during flight or decrease of wing wettability. Representatives of Palaeoptera (Odonata and Ephemeroptera) have no hairy structures on the wing membrane, but both its sides are fine-sculptured. In this study, the nature of the wing covering was studied using acoustic microscopy, scanning- and transmission electron microscopy followed by a variety of chemical treatments. It was shown that wing microsculptures are not cuticular outgrowths, but a wax covering, which is similar to pruinosity, which has been previously described in several odonate taxa. Data from scanning acoustic microscopy revealed that scratches on the wax covering have material density different from the surrounding material. Various functions of the wax covering are discussed.     

Evolutionary relationships of wing venation and wing size and shape in Aphidiinae (Hymenoptera: Braconidae)

We explored evolutionary changes in wing venation and wing size and shape in Aphidiinae, one of the well-known groups of parasitic wasps from the family Braconidae. Forewings of 53 species from 12 genera were examined, for which a molecular phylogeny was constructed on the basis of the mitochondrial barcoding gene COI. By covering all types of wing venation within the subfamily Aphidiinae and by using landmark-based geometric morphometrics and phylogenetic comparative methods, we tested whether evolutionary changes in wing shape correlate to the changes in wing venation and if both changes relate to wing size. The relationship between wing morphology and host specificity has been also investigated. We found that six types of wing venation, with different degree of vein reduction, could be recognized. Wing venation type is largely genus specific, except in the case of maximal reduction of wing venation which could be found across examined Aphidiinae taxa. The reconstruction of evolutionary changes in wing venation indicates that evolutionary changes in wing shape are related to the changes in wing size, indicating that miniaturization play a role in evolution of wing morphology while host specialization does not affect the wing shape within the subfamily Aphidiinae.     

Evolution of insect wings and development – new details from Palaeozoic nymphs

The nymphal stages of Palaeozoic insects differ significantly in morphology from those of their modern counterparts. Morphological details for some previously reported species have recently been called into question. Palaeozoic insect nymphs are important, however – their study could provide key insights into the evolution of wings, and complete metamorphosis. Here we review past work on these topics and juvenile insects in the fossil record, and then present both novel and previously described nymphs, documented using new imaging methods. Our results demonstrate that some Carboniferous nymphs – those of Palaeodictyopteroidea – possessed movable wing pads and appear to have been able to perform simple flapping flight. It remains unclear whether this feature is ancestral for Pterygota or an autapomorphy of Palaeodictyopteroidea. Further characters of nymphal development which were probably in the ground pattern of Pterygota can be reconstructed. Wing development was very gradual (archimetaboly). Wing pads did not protrude from the tergum postero‐laterally as in most modern nymphs, but laterally, and had well‐developed venation. The modern orientation of wing pads and the delay of wing development into later developmental stages (condensation) appears to have evolved several times independently within Pterygota: in Ephemeroptera, Odonatoptera, Eumetabola, and probably several times within Polyneoptera. Selective pressure appears to have favoured a more pronounced metamorphosis between the last nymphal and adult stage, ultimately reducing exploitation competition between the two. We caution, however, that the results presented herein remain preliminary, and the reconstructed evolutionary scenario contains gaps and uncertainties. Additional comparative data need to be collected. The present study is thus seen as a starting point for this enterprise.     

Wing-scales of Pseudoleptocerus chirindensis Kimmins (Trichoptera: Leptoceridae)

The African species Pseudoleptocerus chirindensis belongs to a small group of Trichoptera most unusual in having scaly wings. Electron microscope studies reveal 13 structurally distinct kinds of cuticular process on the wings, including several types of squamiform and hair-like macrotrichia. These are described in detail and their possible functions inferred. The optical properties of the scales forming the colour pattern of the forewings are related to ultrastructural elements including diffraction and thin film interference systems. Trirhopteran scale structure is compared with that of the Lepidoptera, the sister-group in the Amphiesmenoptera. Differences are found and it is tentatively concluded that wing-scales have evolved independently in the two orders.     

Combined molecular and morphological evidence on the phylogeny of the earliest lepidopteran lineages

Agreement among recent morphological and molecular phylogenetic analyses has strengthened estimates of the relationships among the earliest lineages of the holometabolan order Lepidoptera. For a few major groups, evidence for monophyly and basal relationships remains relatively weak or contradictory — chiefly within the clades of basal Glossata and Heteroneura. Here we assess the support for these controversial areas of lepidopteran classification through molecular systematic investigation of 18S rDNA sequence variation. Parsimony and maximum likelihood analyses are presented for 1379 alignable sites of 18S. These data are then combined with 61 morphological features scored for major lineages of basal Glossata and Heteroneura. Our 18S rDNA data support recent hypotheses for the placement of Micropterigidae and Agathiphagidae as the basal-most lineages of Lepidoptera, and support the monophyly of the groups Neolepidoptera and Exoporia. 18S data alone are shown to be insufficient for resolving the monophyly and relationships of the Glossata, and for specifying relationships above the Neolepidoptera. Combination of the 18S data with published morphological ground-plan scorings improves overall support for the morphology-based hypothesis for basal glossatans, but phylogenetic resolution among published alternatives for the basal Heteroneura remains a major question for lepidopteran systematics.     

Lateral line scale types and review of their taxonomic distribution

The trunk canal of fishes is contained within a series of lateral line (LL) scales. To categorise LL scale structural types, and determine their distribution, an analysis of original data was undertaken using light and scanning electron microscopy in combination with a literature survey from over 1,000 species representative of most orders of bony fishes. Our categorisation of LL scales is based on the relationship between the tube, or ossified trunk canal segment, and associated scale. Tubular‐Scalar LL scales consist of a distinguishable tube and elasmoid scale in scale pockets. Four types occur only in species with elasmoid scales. Integrated LL scales do not develop in scale pockets, and their tube is enclosed or extended by a non‐elasmoid scale or spines. Integrated 1 and 2 LL scales co‐occur with ganoid and calcidermoid scales, and Integrated 3 LL scales occur when common scales are absent or elasmoid. Tubular LL scales are tubes only, occurring mainly in scaleless species or with calcidermoid and elasmoid scales. Non‐Tubular LL scales are composed only of a scale, co‐occurring mainly with cycloid scales. There is consistency of LL scale type in many orders, families and genera and the presence of different types within taxa can be meaningful.     

Pyramimonas tychotreta, sp. nov. (prasinophyceae), a new marine species from antarctica: light and electron microscopy of the motile stage and notes on growth rates

An undescribed marine prasinophyte, Pyramimonas tychotreta, sp. nov., was isolated from a water sample collected near the ice edge in the Weddell Sea (Antarctica) and is characterized by means of light and electron microscopy. This is the second described Antarctic species in the genus and it possesses a cell ultrastructure typical for members of the subgenus Vestigifera McFadden. The quadriflagellated cells measure 8–12 μm in length and 6–7 μm in width and are equipped with seven types of organic scales that cover the flagella and cell body. The scale floor of the box scales is ornamented by quadrants of parallel striations running perpendicular to one another. The scale floor is further characterized by a number of randomly positioned perforations. The wall of the box scales may be solid or possess up to five perforations. The base of the crown scales is square with rounded corners. It is formed of two crossed ribs, the extremities of which are interconnected by a peripheral rib. Four upright arms, attached to the peripheral rib in positions slightly offset from its junction with the cross ribs, join up with the distal extremity of a central upright strut. Each arm possesses two spines. The limuloid scales are cross‐striped by 10–12 ribs. Some details of the flagellar apparatus are briefly reported. Pyramimonas tychotreta is compared with other species of the genus. Experiments were conducted to study the response of growth rate to variations in temperature and salinity in the clonal culture. The best growth rate (0.45 divisions·24 h ^{? 1}) was found at 4.6° C; growth ceased at temperatures in excess of 12° C. Growth in salinities ranging from 15 to 35 psu was similar, but was arrested at 10 psu. These studies suggest that P. tychotreta as a cold stenotherm and euryhaline taxon. New observations are presented on the geographic distribution of previously described species of Pyramimonas Schmarda from the Northern Foxe Basin, Canada.     

Phylogeny,landmark analysis and the use of wing venation to study the evolution of social wasps (Hymenoptera: Vespidae: Vespinae)

Wing venation provides useful characters with which to classify extant and fossil insects. Recently, quantification of its shape using landmarks has increased the potential of wing venation to distinguish taxa. However, the use of wing landmarks in phylogenetic analyses remains largely unexplored. Here, we tested landmark analysis under parsimony (LAUP) to include wing shape data in a phylogenetic analysis of hornets and yellow jackets. Using 68 morphological characters, nine genes and wing landmarks, we produced the first total‐evidence phylogeny of Vespinae. We also tested the influence of LAUP parameters using simulated landmarks. Our data confirmed that optimization parameters, alignment method, landmark number and, under low optimization parameters, the initial orientation of aligned shapes can influence LAUP results. Furthermore, single landmark configurations never accurately reflected the topology used for data simulation, but results were significantly close when compared to random topologies. Thus, wing landmark configurations were unreliable phylogenetic characters when treated independently, but provided some useful insights when combined with other data. Our phylogeny corroborated the monophyly of most groups proposed on the basis of morphology and showed the fossil Palaeovespa is distantly related to extant genera. Unstable relationships among genera suggest that rapid radiations occurred in the early history of the Vespinae.     

Sex‐ and age‐dependent morphology and selection on wing shape in the barn swallow Hirundo rustica

Wings have evolved in phylogenetically distant organisms with morphologies that depend on the combined effects of diverse, potentially contrasting selective forces. In birds, long pointed wings boost speed and energetic efficiency during cruising flight but reduce manoeuvrability. Migratory behavior is believed to lead to the evolution of more pointed wings, but selection on pointedness has never been estimated. Because annual routines of migrants are tightly scheduled, wing pointedness may be selected for because it allows for earlier arrival to the breeding grounds. In long‐distance migratory barn swallows Hirundo rustica we showed that selection via breeding date and thus annual fecundity operates on wing pointedness, but not on other wing traits, among yearling females but not among older females or males. Selection on wing pointedness specifically in yearling females may result from climatic effects, which favour earlier arrival from migration, and from yearling females being the sex‐by‐age class with the latest migration and the smallest wing pointedness. Wing morphology differed between sexes and age classes because of change in size of the outermost but not the innermost wing feathers. Hence, sex‐ and age‐specific selection on wing pointedness operates in a species with sex‐ and age‐dependent variation in phenology and wing morphology.     

Effect of non‐lethal sampling on life‐history traits of the protected moth Graellsia isabelae (Lepidoptera: Saturniidae)

Abstract 1. Non‐lethal genetic surveys in insects usually extract DNA from a leg or a piece of wing. Although preferable to lethal sampling, little is known about the effect of leg/wing non‐lethal sampling on fitness‐related traits. 2. Graellsia isabelae (Graells, 1849) is a European moth protected by the Habitats Directive and the Bern Convention. Conservation genetics surveys on this species should therefore use non‐lethal sampling. 3. The present study aimed to (1) quantify the effects of both leg and hind‐wing tail sampling on survivorship and reproductive behaviour of adult males and females, and (2) assess the quality and quantity of DNA obtained from those tissues. 4. Both hind‐wing tails and mid‐legs proved to be good sources of high quality nuclear and mitochondrial DNA. DNA concentration was significantly higher when extracted from a large (130 mm²) piece of the hind‐wing tails than from about half of the mid‐leg. Using mark–release–recapture experiments with adults, it was found that neither mid‐leg nor hind‐wing tail sampling significantly reduced male survivorship or total number of matings. However, although mid‐leg sampling did not significantly affect female survivorship, it had a negative effect on female mating success. 5. Wing‐tail clipping on males appeared to be the best non‐lethal sampling procedure for G. isabelae. It is a fast procedure, similar to natural wing impairment, and did not significantly affect survival or mating behaviour.     

STUDIES ON ULTRASTRUCTURE AND THREE‐GENE PHYLOGENY OF THE GENUS MALLOMONAS (SYNUROPHYCEAE)1

The genus Mallomonas, a common and often abundant member of the planktic community in many freshwater habitats worldwide, consists of 180 species divided into 19 sections and 23 series. Classification of species is based largely on ultrastructural characteristics of the siliceous scales and bristles that collectively form a highly organized covering over the cell. However, the relative importance of the different siliceous features of the scales, such as the dome, V rib, and secondary structures, as well as the different types of scales, in understanding the evolution and phylogeny of the genus is little known. In this study, we investigated the scale and bristle ultrastructure, along with sequences of three genes, for 19 isolates (18 species) of Mallomonas (18 isolates were from Korean habitats). The isolates represented nine of the 19 sections. Sequences for both the nuclear SSU and LSU rDNA and plastid LSU of RUBISCO (rbcL) genes for each of the 19 Mallomonas isolates and four outgroups were determined. Bayesian and maximum‐likelihood (ML) analyses of the data revealed that Mallomonas consists of two strongly supported clades. Mallomonas bangladeshica (E. Takah. et T. Hayak.) Siver et A. P. Wolfe was at the base of the first clade that included taxa from the sections Planae and Heterospinae, both of which lack a V rib on the shield of the scales. Our results indicated that the sections Planae and Heterospinae should be combined. The second clade, with Mallomonas insignis Penard and Mallomonas punctifera Korshikov at the base, contained taxa from the sections Mallomonas, Striatae, Akrokomae, Annulatae, Torquatae, Punctiferae, and Insignes, all of which have V ribs or well‐developed marginal ribs on the scales. Sister relationships between Mallomonas and Striatae were strongly supported, but interrelations among the remaining sections were not resolved, probably due to inclusion of too few species. Our results suggest that the current classification of the genus Mallomonas at the section level will require some revision. Additional species will need to be added in future analyses.     

Phylogenetic relationships of Salix (Salicaceae) based on rbcL sequence data

Nucleotide sequences of the chloroplast-encoded rbcL gene were used to examine phylogenetic relationships of the genus Salix together with other allied genera of the family Salicaceae. Phylogenetic analyses of rbcL sequences strongly suggest the monophyly of three commonly recognized genera (Chosenia, Salix, and Toisusu). Two monophyletic groups are recognized within the larger monophyletic group. They do not correspond with any infrageneric taxa proposed so far. With regard to character evolution, it is thought that the reduction of stamen number from more than two stamens to two might occur in at least three lineages and that fused bud scales evolved several times and/or the reverse evolution occurred from fused to free. Some types of pollen surfaces are considered to have evolved independently.     

Two new species of Mallomonas (Synurophyceae) from the Ocala National Forest, Florida, U.S.A.

Two new species of Mallomonas, M. ocalensis and M. caerula , are described from a clearwater, acidic and oligotrophic waterbody in the Ocala National Forest, Florida, U.S.A. Mallomonas ocalensis , in the Series Doignonianae of the Section Torquatae, consists of relatively small cells with scales that have a series of parallel shield ribs with alternating thicknesses, collar scales with very short bristles and posterior scales with small protruding spines. Mallomonas ocalensis is most similar to Mallomonas dickii , but is distinguished from the latter species based on the highly variable thicknesses of the shield ribs and significantly larger scales. Mallomonas caerula , in the Series Mallomonas of the Section Mallomonas, consists of relatively large cells that bear large domed and dorneless scales, and long, robust and ribbed bristles. Bristles with and without helmets can be found. The shield, especially the distal portion, and the posterior flange of scales consists of series of more or less parallel ribs. Mallomonas caerula appears to be most closely related to Mallomonas acaroides var. acaroides and Mallomonas acaroides var. muskokana , but can be distinguished from the latter taxa on the basis of scale type, scale structure and bristle morphology. Both new species were found in the plankton and surface sediments of Blue Sink Pond.