Comparative morphology of terminal-instar larvae of Cynipoidea: phylogenetic implications

We describe the external morphology of the terminal-instar larvae of 30 species of Cynipoidea (Hymenoptera), with special reference to the head capsule and mouthparts. Twenty-five of the species belong to the Cynipidae and are gall inducers or phytophagous inquilines (guests) in galls, while five represent different insect-parasitic lineages of the Cynipoidea. Although we find only limited variation in body shape, the head sclerites and mandibles offer many characters of potential phylogenetic value. For instance, the mandibles of the parasitoids have one large pointed tooth, with several smaller dents along the inner margin in core figitids, whereas the phytophagous gall inducers and inquilines have mandibles with two or three blunt teeth of subequal size. The mandibles of inquiline larvae are unique in being covered by vertical striations and in having a dominating, broad second tooth. We summarize the qualitative variation among the studied terminal-instar larvae in terms of 33 morphological characters and one life-history trait and examine the phylogenetic implications of these data by running parsimony analyses under uniform character weights and under implied weights (Goloboff weights). The analysis under uniform weights is poorly resolved but the relationships suggested by the implied-weights analysis are largely congruent with previous analyses of adult morphology and molecular data. The larval data support inclusion of the genus Liposthenes in the Neaylax – Isocolus clade, in agreement with the molecular data but in weak conflict with adult morphology. However, the larval data agree with adult morphology and conflict with the molecular data in supporting monophyly of the inquilines.     

Head anatomy of adult Coniopteryx pygmaea : Effects of miniaturization and the systematic position of Coniopterygidae (Insecta: Neuroptera)

External and internal head structures of adult Coniopteryx pygmaea Enderlein, 1906, one of the smallest known lacewings, are described in detail for the first time. Possible effects of miniaturization and two hypotheses on the phylogenetic position of Coniopterygidae are evaluated and compared with data from literature. Several convergent modifications in C. pygmaea and other miniaturized insect species are outlined, e.g., a relative increase in the size of the brain, simplification of the tracheal system with respect to the number of tracheae, and reduction of the number of ommatidia and diameter of the facets. Further, the ocular ridge is bell-shaped and countersunk into the head capsule. The cuticle is weakly sclerotized and equipped with wax glands which are unique in Neuroptera. The total number of muscles is not affected by miniaturization. The phylogenetic analysis yields Coniopterygidae as sistergroup to the dilarid clade based on one larval character, the shape of the stylets. The enforced basal position of Coniopterygidae is supported by one disputable synapomorphy of the remaining Neuroptera, the presence of paraglossae in adults.     

A comparison of techniques used to study the morphology of final instar larvae of aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae)

Abstract. The morphology of the head sclerites of final instar larvae of aphid parasitoids prepared from cast larval skins and excised heads of whole larvae are compared, using both optical and scanning electron microscopy. Remarkable differences in the apparent shape of head sclerites, particularly mandibles, were found. Some of these differences are probably due to a consistent dorso-ventral flattening of the head during moulting, observed in cast skin preparations, whereas excised heads tend to flatten antero-posteriorly. The importance of these findings for understanding the three-dimensional shape of final instar mandibles and for the study of larval morphology and taxonomy of Hymenoptera is discussed     

The enigma of reversed asymmetry in lithodid crabs: absence of evidence for heritability or induction of morphological handedness in Lopholithodes foraminatus

SUMMARY Mutations or environmental factors that result in reversal of conspicuous left–right asymmetries provide an opportunity to study developmental mechanisms. They may also provide insight into evolutionary changes in asymmetry states within and between species. King crabs (family Lithodidae) have a larger right claw and females typically exhibit a dextrally offset abdomen. Nevertheless, I observed a high incidence of left handedness in laboratory reared box crabs ( Lopholithodes foraminatus ) and captured the first known egg-bearing female lithodid to exhibit reversed asymmetry. This provided a unique opportunity to characterize the reversed phenotype and to compare the incidence of reversed asymmetry in the offspring of normal and reversed females. Asymmetry of the chelae became apparent in the first postzoeal stage (glaucothoe) and handedness was maintained through subsequent instars. Females with larger left claws developed reversed abdominal asymmetry by the fourth crab stage. No reversed asymmetry was observed in the mandibles of zoea larvae or juveniles of either handedness. The incidence of reversed asymmetry in glaucothoe reared from one reversed and three normal females was high (between 20% and 30%), and independent of maternity ( P =0.67). Removal of the right cheliped of fourth stage zoeae, and the major cheliped of glaucothoe, did not reverse the direction of asymmetry. Elevated larval rearing temperature also did not affect the frequency of reversed individuals. This lack of evidence for either heritability or induction of handedness is enigmatic. Further investigation of reversed asymmetry in lithodid crabs may provide valuable insights into the development and evolution of bilateral asymmetries.     

Endocranial shape asymmetries in Pan paniscus, Pan troglodytes and Gorilla gorilla assessed via skull based landmark analysis

Brain shape asymmetries or petalias consist of the extension of one cerebral hemisphere beyond the other. A larger frontal or caudal projection is usually coupled with a larger lateral extent of the more projecting hemisphere relative to the other. The concurrence of these petalial components is characteristic of hominins. Studies aimed at quantifying petalial asymmetries in human and great ape endocasts rely on the definition of the midline of the endocranial surface. Studies of brain material show that, at least in humans, most of the medial surface of the left occipital lobe distorts along the midline and protrudes on to the right side, making it difficult for midline and corresponding left and right reference point identification. In order to accurately quantify and compare brain shape asymmetries in extant hominid species, we propose here a new protocol based on the objective definition of cranial landmarks. We describe and quantify for the first time in three dimensions the positions of frontal and occipital protrusions in large samples of Pan paniscus, Pan troglodytes and Gorilla gorilla. This study confirms the existence of frontal and occipital petalias in African apes. Moreover, the detailed analysis of the 3D structure of these petalias reveals shared features, as well as features that are unique to the different great ape species.     

Heterochrony and Early Left-Right Asymmetry in the Development of the Cardiorespiratory System of Snakes

Snake lungs show a remarkable diversity of organ asymmetries. The right lung is always fully developed, while the left lung is either absent, vestigial, or well-developed (but smaller than the right). A ‘tracheal lung’ is present in some taxa. These asymmetries are reflected in the pulmonary arteries. Lung asymmetry is known to appear at early stages of development in Thamnophis radix and Natrix natrix. Unfortunately, there is no developmental data on snakes with a well-developed or absent left lung. We examine the adult and developmental morphology of the lung and pulmonary arteries in the snakes Python curtus breitensteini, Pantherophis guttata guttata, Elaphe obsoleta spiloides, Calloselasma rhodostoma and Causus rhombeatus using gross dissection, MicroCT scanning and 3D reconstruction. We find that the right and tracheal lung develop similarly in these species. By contrast, the left lung either: (1) fails to develop; (2) elongates more slowly and aborts early without (2a) or with (2b) subsequent development of faveoli; (3) or develops normally. A right pulmonary artery always develops, but the left develops only if the left lung develops. No pulmonary artery develops in relation to the tracheal lung. We conclude that heterochrony in lung bud development contributes to lung asymmetry in several snake taxa. Secondly, the development of the pulmonary arteries is asymmetric at early stages, possibly because the splanchnic plexus fails to develop when the left lung is reduced. Finally, some changes in the topography of the pulmonary arteries are consequent on ontogenetic displacement of the heart down the body. Our findings show that the left-right asymmetry in the cardiorespiratory system of snakes is expressed early in development and may become phenotypically expressed through heterochronic shifts in growth, and changes in axial relations of organs and vessels. We propose a step-wise model for reduction of the left lung during snake evolution.     

The larval head of Exechia (Mycetophilidae) and Bibio (Bibionidae) (Diptera)

Exechia and Bibio have retained several plesiomorphic groundplan features of Diptera and Bibionomorpha, including a fully exposed and sclerotized head capsule, the transverse undivided labrum, the absence of movable premandibles, and undivided mandibles without combs. The fusion of the hypostomal bridge with the head capsule and largely reduced antennae are derived features shared by both taxa. The absence of teeth at the anterior hypostomal margin is a potential autapomorphy of Bibionomorpha. A basal position of Anisopodidae is suggested by a number of plesiomorphies retained in this family. Apomorphies of Bibionomorpha excluding Anisopodidae are the reduction of tentorial elements, the partial fusion of the labrum and clypeus, one-segmented antennae, the absence of a separate submental sclerite, the loss of the labial palpus, and the reduction of the pharyngeal filter apparatus. Head structures of Bibio are largely unmodified. The subprognathous orientation is one of few autapomorphic features. In contrast, the mouthparts of Exechia are highly modified in correlation with the specialized food uptake. The rasping counterrotating movements of maxillae and mandibles with teeth oriented in opposite directions are carried out by strongly developed extensors and flexors of the paired mouthparts. The modified labium mechanically supports the “drill head” formed by the mandibles und maxillae. The necessary stability of the head capsule is provided by the hypostomal bridge which also compensates the far-reaching reduction of the tentorium.     

Morph‐dependent form of asymmetry in mandibles of the stag beetle Prosopocoilus inclinatus (Coleoptera: Lucanidae)

Abstract 1. The form of asymmetry in bilateral organs usually follows the same pattern within single populations. However, some exceptions may occur when a population consists of different phenotypes that are from different ontogenic backgrounds and under different selective pressures. We investigated the asymmetric patterns of mandibles of larvae, females, and males in the stag beetle Prosopocoilus inclinatus. 2. Larval mandibles exhibited directional asymmetry both in length and cross direction, whereas female mandibles showed directional asymmetry in cross direction. These asymmetric structures might be more effective in cutting wood fibres. 3. For the relation of male mandible length to body size, a model with a switch point showed a better fit to the data than a convex curve model. This shows that the males are dimorphic with two distinct morphs. 4. The form of asymmetry in male mandible length differed between the morphs. The smaller males exhibited left‐biased directional asymmetry in common with larvae, whereas the larger males exhibited fluctuating asymmetry. 5. This is a novel finding of a morph‐dependent asymmetry. The morph‐dependent asymmetry in males may be as a result of different selection on each morph or a developmental constraint from larval mandibles to adult ones.     

Mouth asymmetry in the textbook example of scale-eating cichlid fish is not a discrete dimorphism after all

Individuals of the scale-eating cichlid fish, Perissodus microlepis, from Lake Tanganyika tend to have remarkably asymmetric heads that are either left-bending or right-bending. The ‘left’ morph opens its mouth markedly towards the left and preferentially feeds on the scales from the right-hand side of its victim fish, and the ‘right’ morph bites scales from the victims’ left-hand side. This striking dimorphism made these fish a textbook example of their astonishing degree of ecological specialization and as one of the few known incidences of negative frequency-dependent selection acting on an asymmetric morphological trait, where left and right forms are equally frequent within a species. We investigated the degree and the shape of the frequency distribution of head asymmetry in P. microlepis to test whether the variation conforms to a discrete dimorphism, as generally assumed. In both adult and juvenile fish, mouth asymmetry appeared to be continuously and unimodally distributed with no clear evidence for a discrete dimorphism. Mixture analyses did not reveal evidence of a discrete or even strong dimorphism. These results raise doubts about previous claims, as reported in textbooks, that head variation in P. microlepis represents a discrete dimorphism of left- and right-bending forms. Based on extensive field sampling that excluded ambiguous (i.e. symmetric or weakly asymmetric) individual adults, we found that left and right morphs occur in equal abundance in five populations. Moreover, mate pairing for 51 wild-caught pairs was random with regard to head laterality, calling into question reports that this laterality is maintained through disassortative mating.     

降香黄檀食叶害虫棕斑澳黄毒蛾的生物学特性及幼虫虫龄判断

【目的】棕斑澳黄毒蛾Orvasca subnotata Walker是于降香黄檀Dalbergia odorifera T. Chen人工林中新发现的一种食叶害虫,其生物学特性描述和龄期判定是虫害预测预报与林间防治的重要依据。【方法】本研究通过定期林间采样的方法,对棕斑澳黄毒蛾不同发育阶段的形态特征及生活史和生活习性进行调查; 并通过测量头宽、前胸毛瘤宽和体长3项分龄指标,测定棕斑澳黄毒蛾幼虫的龄期。【结果】本文报道了棕斑澳黄毒蛾不同发育阶段的形态特征、生活史和生活习性。研究结果表明,棕斑澳黄毒蛾幼虫随龄期增长,前胸逐渐变黑,腹部背线和翻缩腺颜色逐渐加深,特征逐渐变得明显。通过频次分析统计,测得棕斑澳黄毒蛾幼虫龄数为5龄。头宽和前胸毛瘤宽可作为分龄指标,头宽优于前胸毛瘤宽,且前胸毛瘤宽与头宽呈直线关系（y=0.994x-0.114,R=0.999）。体长经统计检验,符合Dyar氏法则,但区域重叠明显,不宜作为分龄指标。【结论】本研究确定了棕斑澳黄毒蛾的生物学特性及幼虫虫龄,可为林间防治提供参考依据。     

Juvenile hormone mediates developmental integration between exaggerated traits and supportive traits in the horned flour beetle Gnatocerus cornutus

Sexually selected exaggerated traits are often coupled with modifications in other nontarget traits. In insects with weapons, enlargements of nontarget characters that functionally support the weapon often occur (i.e. supportive traits). The support of sexual traits requires developmental coordination among functionally related multiple traits—an explicit example of morphological integration. The genetic theory predicts that developmental integration among different body modules, for which development is regulated via different sets of genes, is likely to be coordinated by pleiotropic factors. However, the developmental backgrounds of morphological integrations are largely unknown. We tested the hypothesis that the juvenile hormone (JH), as a pleiotropic factor, mediates the integration between exaggerated and supportive traits in an armed beetle Gnatocerus cornutus. During combat, males of this beetle use exaggerated mandibles to lift up their opponents with the supportive traits, that is, the head and prothoracic body parts. Application of methoprene, a JH analog (JHA), during the larval to prepupal period, induced the formation of large mandibles relative to the body sizes in males. Morphometric examination of nontarget traits elucidated an increase in the relative sizes of supportive traits, including the head and prothoracic body parts. In addition, reductions in the hind wing area and elytra length, which correspond to flight and reproductive abilities, were detected. Our findings are consistent with the genetic theory and support the idea that JH is a key pleiotropic factor that coordinates the developmental integration of exaggerated traits and supportive characters, as well as resource allocation trade‐offs.     

The morphology of the larval head of Tipulidae (Diptera, Insecta) - The dipteran groundplan and evolutionary trends

External and internal head structures of the larva of Tipula montium are described in detail. The results are compared to conditions found in other representatives of Tipuloidea and other dipteran and antliophoran lineages. Despite of the conceivably basal position of Tipulomorpha within Diptera, the larvae are mainly characterised by derived features. The partially retracted head, the specific hemicephalic condition and several other derived character states support the monophyly of Tipuloidea. A clade comprising Tipuloidea excluding Pediciidae is suggested by the strongly retracted head, by deep dorsolateral incisions of the head capsule, by a distinctly toothed anterior premental margin, by the loss of the second extrinsic maxillary muscle, and possibly by the loss of the pharyngeal filter. Eriopterinae and Hexatominae are characterised by a tendency towards an extreme reduction of the head capsule. Limoniinae, Cylindrotomidae, and Tipulidae form a clade supported by the presence of a premaxillary suture. This implies the non-monophyly of Limoniidae. A feature shared by Cylindrotomidae and Tipulidae is the presence of a movable lacinia mobilis. However, this is arguably a plesiomorphic feature, as it also occurs in Nannochoristidae. Features of the larval head of Trichoceridae, which were included in Tipulomorpha, do not show affinities with those of Tipuloidea. Trichocerid larvae share a specialised subdivided mandible with larvae of psychodomorph groups. Tipuloidea are a highly specialised group. The characters examined did not reveal plesiomorphic features supporting a basal position, and features suggesting closer affinities with Brachycera are vague. The evolution of dipteran larval head structures was apparently strongly affected by the loss of legs and the tendency to live in cryptic habitats. Diptera are the group of Endopterygota with the highest number of apomorphic features of the larval head. The appendages are generally simplified and the muscular apparatus is strongly reduced. Specialised features evolving within dipteran lineages include specifically arranged brushes of hairs on the labrum and epipharynx, movable messores, subdivided mandibles, different mandibular brushes, and a far-reaching reduction of labial parts.     

Analysis of the shortvein cis-regulatory region of the decapentaplegic gene of Drosophila melanogaster

In mammals, the Transforming Growth Factor-beta (TGF-beta) superfamily controls a variety of developmental processes. In Drosophila, by contrast, a single member of the superfamily, decapentaplegic (dpp) performs most TGF-beta developmental functions. The complexity of dpp functions is reflected in the complex cis-regulatory sequences that flank the gene. Dpp is divided into three regions: Hin, including the protein-coding exons; disk, including 3' cis-regulatory sequences; and shortvein (shv), including noncoding exons and 5' cis-regulatory sequences. We analyzed the cis-regulatory structure of the shortvein region using a nested series of rearrangement breakpoints and rescue constructs. We delimit the molecular regions responsible for three mutant phenotypes: larval lethality, wing venation defects, and head capsule defects. Multiple overlapping elements are responsible for larval lethality and wing venation defects. However, the area regulating head capsule formation is distinct, and resides 5' to these elements. We have demonstrated this by isolating and describing two novel dpp alleles, which affect only the adult head capsule.     

Testis asymmetry in birds: the influences of sexual and natural selection

Gonad size and shape asymmetries are particularly common in birds. Although some obvious size and shape differences between the left and right testes in birds were first documented more than a century ago, little is known about what influences the variation across species in either the degree or the direction of these asymmetries. Here we show that a left bias in size is the most likely ancestral state in most orders and families, and that there is a weak but significant negative relation between the degree of size and shape asymmetries. In extant species, testis size and shape symmetries increase with the degree of sperm competition (relative testes mass), but those relations are significant only in those species with left bias in each of these traits. When space is particularly constrained on the left side of the body cavity due to a large gizzard (e.g. in granivores), the left testis is more elongated and the degree of size symmetry is larger. Despite significant patterns, relative testes mass and gizzard complexity together explain < 10% of the variation in testis asymmetry in species with left biases. Thus our analyses suggest that some other factor is largely responsible for the evolution of gonad asymmetry in birds: 1) that a left bias in testis size might be a correlated response to selection for a left bias in the development of a single (left) ovary in females of most bird species, and/or 2) that physiological efficiency due to the dramatic and rapid increase in testis size of most species during the breeding season might favour enlargement of one testis. Our conclusions highlight the need to rethink and improve our understanding of the physiological processes underlying the investment in gonads by male birds.     

Progenitor properties of symmetrically dividing Drosophila neuroblasts during embryonic and larval development

Asymmetric cell division generates two daughter cells of differential gene expression and/or cell shape. Drosophila neuroblasts undergo typical asymmetric divisions with regard to both features; this is achieved by asymmetric segregation of cell fate determinants (such as Prospero) and also by asymmetric spindle formation. The loss of genes involved in these individual asymmetric processes has revealed the roles of each asymmetric feature in neurogenesis, yet little is known about the fate of the neuroblast progeny when asymmetric processes are blocked and the cells divide symmetrically. We genetically created such neuroblasts, and found that in embryos, they were initially mitotic and then gradually differentiated into neurons, frequently forming a clone of cells homogeneous in temporal identity. By contrast, larval neuroblasts with the same genotype continued to proliferate without differentiation. Our results indicate that asymmetric divisions govern lineage length and progeny fate, consequently generating neural diversity, while the progeny fate of symmetrically dividing neuroblasts depends on developmental stages, presumably reflecting differential activities of Prospero in the nucleus.     

Size and shape regulation during larval growth in the lepidopteran Pieris brassicae

By adopting a longitudinal study design and through geometric morphometrics methods, we investigated size and shape regulation in the head capsule during the larval development of the cabbage butterfly Pieris brassicae under laboratory conditions. We found evidence of size regulation by compensatory growth, although not equally effective in all larval stages. Size compensation is not attained through the regulation of developmental timing, but rather through the modulation of per‐time growth rate. As for the shape, neither the variance of the symmetric component of shape, nor the level of fluctuating asymmetry show any evidence of increase across stages, either at the population or individual level, which is interpreted as a mark of ontogenetic shape regulation. In addition, also the geometry of individual asymmetry is basically conserved across stages. While providing specific documentation on the ontogeny of size and shape variation in this insect, this study may contribute to a more general understanding of developmental regulation and its influence on phenotypic evolution.     

二种萤火虫幼虫捕食行为及头部功能性结构观察比较

对短角窗萤Diaphanes sp.和胸窗萤Pyrocoelia pectoralis Olivier幼虫的捕食行为进行观察。发现短角窗萤捕食蚯蚓而胸窗萤则捕食蜗牛。利用扫描电镜对短角窗萤和胸窗萤幼虫的头部及口器进行观察比较。发现2种幼虫均具有1对侧单眼,1对发达的、左右对称的3节触角。2种幼虫口器非常发达,具1对锋利的、中空的镰刀状上颚,1对下颚须,1对下唇须,1对内颚叶。2种萤火虫口器最具特别的结构在于:短角窗萤上颚的2/3处内侧着生1个球形结构,上面着生1层齿状结构,而胸窗萤则在上颚基部内侧弯曲成齿状突起。