Morphological and physiological differences of the auditory system in three related bushcrickets (Orthoptera: Phaneropteridae, Poecilimon)

Abstract. The auditory system of three closely related bushcrickets was investigated with respect to morphological and physiological differences. The size of the acoustic vesicle in the prothorax cavity and the size of the acoustic spiracle were compared to differences in auditory tuning of the tympanic nerve and differences in the directionality. The results indicate that a small auditory vesicle and auditory spiracle provide reduced sensitivity in the high frequency range (above 10—15 kHz), but increase sensitivity at low frequencies (below 10 kHz). The directionality of the hearing system deteriorates at frequencies between 10 and 25 kHz in species with a small spiracle and trachea. The evolutionary implications of these differences of the auditory systems are discussed. They are considered to be influenced more by ecological factors than bioacoustical ones.     

中国特产雉类—褐马鸡,藏马鸡和蓝马鸡卵壳的电镜观察

迄今为止,在国内有关野生鸟类卵壳的超微结构报道尚少。我们对中国特产鸡类——三种马鸡卵壳的表皮、气孔、木栅层、锥体层、乳头结、壳膜和基帽进行了扫描巨镜观察,并对结果进行了比较、讨论。认为三种马鸡不同结构的形态,对研究它们的亲缘关系提供了新的价值。     

Functional morphology of the spiracles in two species of Argasidae (Metastigmata: Acarina), with particular reference to responses to desiccation

Spiracle morphology of Argas persicus (Oken) and Omithodorus acinus Whittick is described and the differences between the two species are related to biological factors. Argasid and ixodid spiracle structures and functions are compared.     

Do large bushcrickets have more sensitive ears? Natural variation in hearing thresholds within populations of the bushcricket Requena verticalis (Listroscelidinae: Tettigoniidae)

The auditory spiracle of tettigoniid Orthoptera influences hearing threshold and, for the most part, individuals with larger auditory spiracles have lower hearing thresholds; they are more sensitive. Hearing thresholds of both sexes of the bushcricket, Requena verticalis Walker (Orthoptera; Tettigoniidae; Listroscelidinae), were measured at the male call's carrier frequency and were found to correlate with spiracle dimension. In turn, spiracle dimension correlates with the size of the insect as measured by pronotum length. The best frequency of hearing is close to 16 kHz and this appears to be independent of size. Males show a higher variation in threshold than females and this was reflected in a trend toward lower variance in spiracle size in females.
To test the effects of size on sensitivity, spiracle size was manipulated by partially blocking it. Blocking the spiracle decreases sensitivity to high rather than low frequencies. As in other tettigoniids, the spiracle and associated auditory system act as a high-pass filter. Within and between sex differences in hearing sensitivity were compared with differences in male call intensity. It is argued that sensitivity to sounds associated with mating should be as much under sexual selection as the sexual calls of males.     

肥须亚麻蝇幼虫不同发育阶段气门形态学变化

为了满足实际工作中利用蝇类幼虫日龄推断尸体死后间隔时间的需求,本文采用显微镜观察不同龄期肥须亚麻蝇 Parasarcophaga crassipalpis (Macquart)幼虫气门的形态学变化,通过计算机图像分析系统对幼虫的前、后气门等可用于判断幼虫生长发育的形态学指标进行测量。结果表明：肥须亚麻蝇幼虫前、后气门随时间而发生规律性的变化,幼虫前气门色素管部的平均光密度及其长宽比,以及后气门的平均光密度这 3 项指标是推算幼虫日龄较理想的指标,对法医学中尸体死亡时间的推测具有重要的指示意义。此外,叠龄期作为幼虫发育的必经阶段,可为特殊时间点的推测提供可靠依据。     

纵纹腹小、鵰及长耳的卵壳超微结构研究（形目：鸱科）

本文首次报道纵纹腹小、、长耳卵壳的气孔、木栅层、锥体层、乳锥体、壳膜、基底帽等结构的扫描电镜观察,并对不同种类进行分析,初步探讨了它们的分类价值和生态意义。     

Spiracular transpiration in ticks: a passive diffusion barrier in three species of Ixodidae (Metastigmata: Acarina)

In ixodid ticks the morphology of the spiracle, and in particular that of the labyrinth, differs between species. A reduction in transpiration of water vapour by the 'mutual interference' effect previously described in Ixodes ricinus (L.) is not significant in many species. Instead, transpiration is retarded by very restricted air movement within the labyrinth and by a high resistance to diffusion generated by numerous small aeropyles. These adaptations may be particularly useful in arid environments.     

Characters in the book lungs of Scorpiones (Chelicerata,Arachnida) revealed by scanning electron microscopy

The fine structure of the book lungs in 29 species representing ten monophyletic taxa of the Scorpiones (Arachnida) was investigated using scanning electron microscopy (SEM). Scorpion lungs are not homogeneous across the group. Here we describe and score three sets of phylogenetically informative characters: (1) the surface ornament of the lung lamellae, (2) the distal margins of the lamellae and (3) the fine structure of the spiracle margin. Provisional results suggest that reticulation on the surface of the lung lamellae is characteristic of the Buthidae. By contrast, non-buthid scorpions maintain the air space between adjacent lamellae using projecting trabeculae. Typically they are simple struts, but the trabeculae are distally branched in all investigated Scorpionidae, plus at least one species belonging to the Liochelidae. Simple thorns on the lamellar margins probably represent the plesiomorphic condition, while more complex, branched, arcuate morphologies appear to be homoplastic, occurring sporadically in numerous scorpion sub-groups. The tightly packed, hexagonal pillars around the posterior margin of the spiracle support a close relationship between Scorpionidae and Liochelidae, to the exclusion of the Urodacidae.     

The tettigoniid (Orthoptera : Tettigoniidae) ear: Multiple functions and structural diversity

The ears of bushcrickets (Orthoptera : Tettigoniidae) consist of a pair of membranes situated beneath the knee of the foretibia. These membranes, or tympana, are backed by a tracheal system dedicated to sound reception. In most species, a trachea opens through a hole, the auditory spiracle, in the pleuron of the prothorax adjacent to the ventilatory spiracle. Experimental evidence clearly shows that in those species possessing such an opening, sound entering through this route is amplified by the shape of the trachea, or at least is modified through resonances created by its tracheal cavities or tubes. In some species the external surface of the tympanic membrane is surrounded by cuticular folds, which may be formed into small resonating cavities or sound guiding pinnae. In other species the tympana are naked. Sound interacts with the external surface of the tympana, either through these cuticular cavities or directly at the surface of the membrane. As with auditory tracheae, the shape of these cavities may modify sound through resonances. This review examines the role of both external and internal sound-guides in sound reception. It discusses how different species may utilize different sound receiving systems and suggests how, in some cases, both external and internal routes may respond to different frequency bands. The maximum sensitivity of the ear invariably coincides with the male's call carrier frequency. Two Australian species of tettigoniid where the ear is clearly not tuned to the male's call are described. The biology of one undescribed species belonging to the subfamily, Zaprochilinae, where not only is the ear untuned, but there is marked sexual dimorphism in ear structure, is elaborated on. The review concludes with an examination of the routes selection may have taken to arrive at the extreme morphologies present in the Tettigoniidae.     

Rickettsiae-like structures in the larval salivary gland cells of Drosophila auraria

Rickettsiae-like structures were found in the salivary gland cells of Drosophila auraria during different larval and prepupal developmental stages, from the early 3rd instar up to 14 hr after spiracle inversion. These microorganisms are surrounded by a membrane, are constantly intracellular, and occur singly or in groups. Their widespread occurrence in various tissues of other Drosophila species indicates that they can be considered as symbionts, but their actual functional significance (if any) is unknown.     

Ecdysteroid levels and developmental events during larval moulting in the silkworm, Bombyx mori

In the silkworm, Bombyx mori, apolysis of the spiracle is the first visible sign of the initiation of a larval moult. After spiracular apolysis, the characteristic sequence of new spiracle formation can be recognized easily through the cuticle around the old spiracle. This sequence, called the spiracle index, was used as the basis for a precise developmental chronology of the larval moulting period. At certain spiracle-index stages, histological changes in the larval ventral abdominal integument were also examined. By allatectomy and abdominal ligations, the critical periods for the secretion of juvenile hormone and ecdysone respectively were found to be at the initiation of spiracular apolysis and at the time when general apolysis was complete. The haemolymph-ecdysteroid level as measured by radioimmunoassay was about 30–50 ng/ml during the first 2 days of the 4th instar, 60–70 ng/ml at the time of initiation of spiracular apolysis, and 200 ng/ml at the time of completion of apolysis of the general body surface. The maximal level of the hormone was about 290 ng/ml at the mid-moulting period. The relationship between the ecdysteroid titre and morphological events occurring during the larval moult was discussed.     

A combinatorial enhancer recognized by Mad, TCF and Brinker first activates then represses dpp expression in the posterior spiracles of Drosophila

A previous genetic analysis of a reporter gene carrying a 375-bp region from a dpp intron (dppMX-lacZ) revealed that the Wingless and Dpp pathways are required to activate dpp expression in posterior spiracle formation. Here we report that within the dppMX region there is an enhancer with binding sites for TCF and Mad that are essential for activating dppMX expression in posterior spiracles. There is also a binding site for Brinker likely employed to repress dppMX expression. This combinatorial enhancer may be the first identified with the ability to integrate temporally distinct positive (TCF and Mad) and negative (Brinker) inputs in the same cells. Cuticle studies on a unique dpp mutant lacking this enhancer showed that it is required for viability and that the Filzkorper are U-shaped rather than straight. Together with gene expression data from these mutants and from brk mutants, our results suggest that there are two rounds of Dpp signaling in posterior spiracle development. The first round is associated with dorsal-ventral patterning and is necessary for designating the posterior spiracle field. The second is governed by the combinatorial enhancer and begins during germ band retraction. The second round appears necessary for proper spiracle internal morphology and fusion with the remainder of the tracheal system. Intriguingly, several aspects of dpp posterior spiracle expression and function are similar to demonstrated roles for Wnt and BMP signaling in proximal-distal outgrowth of the mammalian embryonic lung.     

The spiracle of Ixodes ricinus (L.) (Ixodidae: Metastigmata: Acarina): a passive diffusion barrier for water vapour

The structure of the spiracle in the adult female of Ixodes ricinus (L.) is assessed. Gaseous exchange between the tracheal system and the external atmosphere is shown to occur only via the aeropyles in the surface of the sieve plate and is regulated by the atrial valve. The elaborate structure is shown to be an effective passive mechanism to retard the transpiration of water vapour from the tracheal system. The pedicels reduce airflow within the labyrinth to a minimum, while the size and arrangement of the aeropyles allow the formation of a cloud of water vapour above the surface of the spiracle. It is from this cloud which diffusion of water vapour takes place, with no significant changes in water vapour pressure occurring within the tracheal system or the spiracle itself. The mechanics of the system are analysed.     

A new biophysical method to determine the gain of the acoustic trachea in bushcrickets

A method is described for measuring the gain (i.e., the change of amplitude and phase angle) for sounds that propagate to the internal surface of the tympana in ears working as pressure difference receivers. The gain of the acoustic trachea has been measured in two similarly sized and closely related species of bushcrickets, in which the acoustic spiracles and tracheae differ markedly in size. The amplitude part of the gain is much larger in the species with the larger acoustic spiracle, whereas the phase part is very similar in the two species. The method is compared with other methods, which in the past have been used for estimating the gain of sound pathways inside animal bodies.     

二种尸食性蝇类幼虫形态学变化及在法医学上的意义

为了满足实际工作中利用蝇类幼虫日龄推断尸体死后间隔时间的需求,采用形态观察与图像分析相结合的方法对巨尾阿丽蝇Aldrichina grahami(Aldrich)、大头金蝇Chrysomya megacephala(Fabricius)幼虫前气门、后气门、头咽骨进行了研究,筛选适合日龄推断的指标。结果显示:2种尸食性蝇类幼虫前气门、后气门、头咽骨均随着时间的延长发生规律性的变化。其中,咽骨的骨化面积、平均光密度,后气门的平均光密度是最理想的判断幼虫日龄的指标,为法医昆虫学精确推断死后间隔时间提供重要的指导意义。     

Morphometric study of third-instar larvae from five morphotypes of the Anastrepha fraterculus cryptic species complex (Diptera,Tephritidae)

The occurrence of cryptic species among economically important fruit flies strongly affects the development of management tactics for these pests. Tools for studying cryptic species not only facilitate evolutionary and systematic studies, but they also provide support for fruit fly management and quarantine activities. Previous studies have shown that the South American fruit fly, Anastrepha fraterculus, is a complex of cryptic species, but few studies have been performed on the morphology of its immature stages. An analysis of mandible shape and linear morphometric variability was applied to third-instar larvae of five morphotypes of the Anastrepha fraterculus complex: Mexican, Andean, Ecuadorian, Peruvian and Brazilian-1. Outline geometric morphometry was used to study the mouth hook shape and linear morphometry analysis was performed using 24 linear measurements of the body, cephalopharyngeal skeleton, mouth hook and hypopharyngeal sclerite. Different morphotypes were grouped accurately using canonical discriminant analyses of both the geometric and linear morphometry. The shape of the mandible differed among the morphotypes, and the anterior spiracle length, number of tubules of the anterior spiracle, length and height of the mouth hook and length of the cephalopharyngeal skeleton were the most significant variables in the linear morphometric analysis. Third-instar larvae provide useful characters for studies of cryptic species in the Anastrepha fraterculus complex.     

Hearing dimorphism, trait variation and conflicts over space in the thorax of the bushcricket Requena verticalis (Listroscelidinae: Tettigoniidae: Orthoptera)

The hearing system of Requena verticalis is sexually dimorphic. Previous work has shown size of the auditory spiracle determines absolute threshold and as female spiracles are, on average, larger than males, females are more sensitive to the main energy of the male call. In all measured traits in morphology and physiology, females showed lower coefficient of variation than males. This difference was significant for bulla volume and hearing threshold. In addition, female ear size covaries with thorax dimensions but this is not so in males. Such a finding suggests stabilising selection on ear size in females, perhaps explained by the requirement of females to recognise and locate the male. As the auditory bulla is larger in females than males, so occupying thoracic space, we suggest a possible trade-off in this brachypterous species between hearing sensitivity and sound production. Finally, we examine relative growth of body structures not associated with hearing and those that influence hearing sensitivity. Scaling, where traits are under strong selection, may result in allometry. Female hearing traits show positive allometry with absolute size and while the relationship between bulla volume and spiracle area was positively allometric in females this was not the case for males.     

Larval morphology of the lesser housefly, Fannia canicularis

The morphology of all larval instars of Fannia canicularis (Linnaeus) (Diptera: Fanniidae) is documented using a combination of light and scanning electron microscopy. The following structures are documented for all instars: antennal complex; maxillary palpus; facial mask; cephaloskeleton; ventral organ; anterior spiracle; Keilin's organ; posterior spiracle; fleshy processes, and anal pad. Structures reported for the first time for all instars include: two pairs of lateral prominences on the prothoracic segment; additional ventrolateral prominences on the second thoracic segment, and a papilla at the base of the posterior spiracle. Other structures reported for the first time are anterior spiracles in the first instar and a serrated tip on the mouthhook in the second instar. A trichoid sensillum on the posterior spiracular plate, representing a sensory organ otherwise unknown in the Calyptratae, is described in the second and third instars. Results are discussed and compared with existing knowledge on dipteran larval morphology.     

Spiracular closing mechanisms in the firebrat, Thermobia domestica (packard) (Thysanura)

Mechanisms for regulating the degree of opening of its spiracles are present in Thermobia. That of the mesothoracic spiracle is of the external type with a flap-like hood guarding the spiracular aperture. Contraction of muscles open the spiracle by raising the hood. Closure is brought about by muscular relaxation and elastic cuticular recoil. Opening is either partial, with small-scale oscillatory movements ('fluttering'), or complete ('wide-opening'). Wide-opening follows bouts of muscular activity. Carbon dioxide anaesthesia relaxes the opener muscles causing the spiracles to close by elastic recoil. This explains continued low tracheal water loss during anaesthesia, and also in death. The control mechanisms of the metathoracic and 8 pairs of abdominal spiracles are of the internal type, with a crypt-like atrium leading into the slit-like neck region of the spiracular pit, one side of which has an elastic cuticular rod running along it. Muscles inserted on the opposite side widen the aperture. As with the mesothoracic spiracle, closure is brought about by muscular relaxation and elastic cuticular recoil.