The photomechanic infrared receptor for the detection of forest fires in the beetle Melanophila acuminata (Coleoptera: Buprestidae)

We recorded from single units of individual sensilla of the thoracic infrared (IR) pit organs of Melanophila acuminata. When the organ was stimulated with a thermal radiator whose emission spectrum was similar to that of a typical forest fire, units responded phasically with up to seven spikes within 30–40 ms at a radiation power of 24 mW cm⁻². In the experiments all wavelengths shorter than 1.6 μm were excluded by a longpass IR filter. Response latencies were about 4 ms and initial impulse frequencies were up to 250 impulses per second (ips). A single spike could be generated even when stimulus duration was only 2 ms. Reduction of total radiation power from 24 mW cm⁻² to 5 mW cm⁻² resulted in increased response latencies of 5–6 ms and the occurrence of only two to three spikes. Initial impulse frequencies decreased to 125 ips. According to our physiological results and calculations, Melanophila should be able to detect a 10-hectare fire from a distance of 12 km. Mechanical stimuli also evoked responses of the IR sensilla. All present morphological and physiological findings lead to the conclusion that the IR receptors of Melanophila must function by means of a hitherto undescribed photomechanic mechanism. Accepted: 1 November 1997     

Sensitivity threshold and response characteristics of infrared detection in the beetle Melanophila acuminata (Coleoptera: Buprestidae)

The minimum detection threshold of the infrared sensitive beetle, Melanophila acuminata, was measured with a helium-neon laser that emitted light at a wavelength of 3.39 microm. Extracellular recordings were taken both at the pit organ responsible for detection and at the interganglionic connectives in the thorax of the beetle. At the pit organ, generator and action potentials from single neurons were measured with a sharpened tungsten electrode. At the connectives that linked the fused second meso-/metathoracic and prothoracic ganglia, compound action potentials were measured with a tungsten hook electrode that encircled the connective. The latter recordings confirmed conveyance of infrared information through specific pathways to rostrally-situated sites in the nervous system of the beetle. The 50% probability irradiance threshold at which action potentials were elicited from the receptor and connectives occurred at 17.3 and 14.6 mW/cm(2), respectively. In addition to sensitivity threshold, several other characteristics of the response were quantified including dependence of generator potential latency, generator potential duration, spike frequency, and spike latency on irradiance, dependence of response strength (spike count) on exposure time, and flicker fusion frequency. The ability to detect infrared radiation is rare in nature, and these results provide valuable information necessary to understand this unique sensitivity.     

Afferent projections of infrared-sensitive sensilla in the beetle Melanophila acuminata (Coleoptera: Buprestidae)

Beetles of the genus Melanophila are able to detect infrared radiation by using specialized sensilla in their metathoracic pit organs. We describe the afferent projections of the infrared-sensitive neurons in the central nervous system. The axons primarily terminate in the central neuropil of the fused second thoracic ganglia where they establish putative contacts with ascending interneurons. Only a few collaterals appear to be involved in local (uniganglionic) circuits. About half of the neurons send their axons further anterior to the prothoracic ganglion. A subset of these ascend to the subesophageal ganglion, and about 10% project to the brain. Anatomical similarities suggest that the infrared-sensitive neurons are derived from neurons supplying mechanosensory sensilla. The arborization pattern of the infrared afferents suggests that infrared information is processed and integrated upstream from the thoracic ganglia.     

Electrophysiological characterization of the multipolar thermoreceptors in the "fire-beetle" Merimna atrata and comparison with the infrared sensilla of Melanophila acuminata (both Coleoptera,Buprestidae)

A thermosensitive multipolar neuron innervates each of the four abdominal receptors of the Australian buprestid beetle Merimna atrata. The neuron is spontaneously active within a broad range of body temperatures (tested between 10°C and 40°C). We heated the receptors with a red diode laser (=0.66 µm) at intensities ranging from 5.3 mW cm^–2 up to 1.3 W cm^–2. In general, warming caused an increase of receptor activity. Peak discharge frequencies were reached 100–300 ms after onset of irradiation. After peak frequencies were reached, distinct adaptation took place within seconds. A linear increase in irradiation intensity caused an exponential increase in peak frequencies. Lowest threshold was found to be at 40 mW cm^–2 where latencies were 47 ms. At the highest intensity tested (1.3 W cm^–2), peak frequencies increased up to about 300 Hz and latencies decreased to 24 ms. Considering the pyrophilous behaviour of Merimna and the morphological data from previous studies, our results support the hypothesis that the abdominal receptors are infrared receptors. We also recorded the responses of the photomechanic infrared sensilla of Melanophila acuminata under the same experimental conditions. These results show that the photomechanic sensillum of Melanophila has a higher sensitivity, and that the latencies are considerably shorter.     

The infrared receptor of Melanophila acuminata De Geer (Coleoptera: Buprestidae): ultrastructural study of a unique insect thermoreceptor and its possible descent from a hair mechanoreceptor

The paired infrared organs of Melanophila acuminata consist of 50-100 sensilla situated at the bottom of a pit next to the coxae of the mesothoracic legs, where no exocuticle is developed. Each sensillum is accompanied by a wax gland and has a cuticular lens-like spherule (diameter 12-15 mum) bulging out with its upper hemisphere above the surface, covered only by a thin cuticle of about 1 mum. Distal processes of two enveloping cells surround the entire spherule in the form of a flattened protoplasmatic layer with the exception of a small apical stalk connecting the spherule to the outer cuticle. The spherule is innervated by a single sensory neuron of the ciliary type which is anchored ventrally with the distal tip of its cylindrical and unbranched DOS in the spherule. The insertion of the dendrite, which contains a well-developed tubular body, is always eccentric like in a hair mechanoreceptor (sensillum trichodeum) and there is no evidence of any optical function of the spherule. Three enveloping cells exist, but only one - probably the trichogen cell - forms a relatively small outer receptor lymph cavity. In the posterior wall of the pit - where exocuticle is developed - so-called suppressed systems can be found which remain completely below the cuticle with their otherwise well-developed spherules. Additionally, there is a tendency towards basally flattening and longitudinally stretching of spherules which are situated more peripherally. They strongly resemble the basal regions of hair mechanoreceptors (sensilla trichodea) in their immediate neighbourhood which are also accompanied by wax glands. Because of the existence of these transitional stages and the great ultrastructural resemblance between infrared receptors and hair mechanoreceptors concerning the bauplan of the sensory neurons and their mode of innervating the cuticular apparatus, we conclude that the infrared sensilla are probably derived from hair mechanoreceptors. Based on these results and transmission measurements of infrared radiation through the cuticular components of the organ, a model of the possible function of the infrared receptor is presented.     

Morphology of the prothoracic discs and associated sensilla of Acanthocnemus nigricans (Coleoptera, Acanthocnemidae)

The Australian ‘little ash beetle’ Acanthocnemus nigricans (Coleoptera, Cleroidea, Acanthocnemidae) is attracted by forest fires. A. nigricans has one pair of unique prothoracic sensory organs and it has been speculated that these organs may play a role in fire detection. Each organ consists of a cuticular disc, which is fixed over an air-filled cavity. On the outer surface of the disc, about 90 tiny cuticular sensilla are situated. The poreless outer peg of a sensillum is 3–5 μm long and is surrounded by a cuticular wall. One ciliary sensory cell innervates the peg. As a special feature, the outer dendritic segment is very short already terminating below the cuticle. A massive electron-dense cylindrical rod, which most probably represents the hypertrophied dendritic sheath, extends through the cuticular canal connecting the tip of the outer dendritic segment to the peg. The dendritic inner segment and the soma are fused indistinguishably. Thin, leaflike extensions of glial cells deeply extend into that conjoint and considerably enlarged compartment which also contains large numbers of mitochondria. In summary, the sensilla of the sensory disc of A. nigricans represent a new type of insect sensillum of hitherto unknown function. The possible role of the prothoracic sensory organ in fire detection is discussed.     

The analysis of the mechanosensory origin of the infrared sensilla in Melanophila acuminata (Coeloptera; Buprestidae) adduces new insight into the transduction mechanism

The thoracic infrared (IR) sensilla of the pyrophilous jewel beetle Melanophila acuminata most likely have evolved from hair mechanoreceptors (sensilla trichodea). To further elucidate the sensory transduction mechanism, the morphology of IR sensilla and of neighbouring hair mechanoreceptors was investigated by using conventional electron microscopical techniques (SEM, TEM) in combination with focused ion beam milling (FIB). It was assumed that any deviation from the bauplan of a sensillum trichodeum is of particular concern for the transduction of IR radiation into a mechanical stimulus. Thus, the structures supposed to be relevant for stimulus uptake and transduction were homologized. Compared to a hair mechanoreceptor, an IR sensillum shows the following special features: (i) the formation of a complex cuticular sphere instead of the bristle; the sphere consists of an outer exocuticular shell as well as of an inner porous mesocuticular part. (ii) The enclosure of the dendritic tip of the mechanosensitive neuron inside the sphere in a fluid-filled inner pressure chamber which is connected with a system of microcavities and nanocanals in the mesocuticular part. Hence we propose that an IR sensillum most probably acts as a microfluidic converter of infrared radiation into an increase in internal pressure inside the sphere which is measured by the mechanosensitive neuron.     

Variation in number and differentiation of the abdominal infrared receptors in the Australian 'fire-beetle' Merimna atrata (Coleoptera, Buprestidae)

Most individuals of the Australian ‘fire-beetle’ Merimna atrata have two pairs of IR receptors which are located ventrolaterally on the second and third abdominal sternite. An IR receptor consists of a specialized IR absorbing area, which is innervated by a neural complex. This complex contains one thermoreceptive multipolar neuron with a unique terminal dendritic mass (TDM) and two scolopidia and was termed ‘sensory complex’. However, also individuals with one pair of IR receptors on the second sternite and beetles with three pairs on the second, third, and fourth sternites were found. Additionally, beetles having one or two pairs of IR receptors may have preliminary stages of IR receptors on the third and fourth sternite, respectively. We found two kinds of preliminary stages, both of which are characterized by a much less pronounced absorbing area. In all five abdominal sternites segmental nerves are attached to the cuticle with a neural complex. Investigation of complexes of non-IR sternites suggests that the sensory cells inside the sensory complex of an IR receptor have developed from common internal stretch receptors. From our results it can be hypothesized that the IR sensory system in Merimna atrata has not yet reached a stage, which can be regarded as evolutionary stable.     

Thermomechanical properties of the stimulus transducing cuticle in the infrared organ of Merimna atrata (Coleoptera,Buprestidae)

The pyrophilous Australian “fire‐beetle” Merimna atrata strongly depends on the occurrence and localization of forest fires for its reproduction. As a special adaptation to its unusual biology, elaborate infrared (IR) organs have evolved in this species. The IR‐organs consist of a specialized cuticular portion, the absorbing area, innervated by a sensory complex. The sensory complex contains a thermosensitive multipolar neuron with a specialized dendritic region, the terminal dendritic mass, and a mechanosensitive unit represented by a chordotonal organ (CO). Evidence for the IR‐receptive function so far has only been provided for the multipolar neuron. Based on morphological data, it has been hypothesized that the CO could also be involved in IR‐reception by measuring minute thermal deformations of the absorbing area. To test this hypothesis, we investigated structural features like cuticle thickness, reduced Young's modulus and hardness of the absorbing area. The results were used in finite element simulations to analyze the thermomechanical behavior and performance of the IR‐organ. Our findings indicate that considerable thermal deformation of the absorbing area occurs, supporting the hypothesis that the CO could function as photomechanical IR‐receptor. Interestingly, at the innervation site of the CO the lowest relative displacements of the absorbing area were found. This may indicate that the CO as putative photomechanic IR‐receptor has not been adapted according to the requirements of highest sensitivity. Probable benefits of the bimodal innervation by a thermosensory and a mechanosensory unit and their possible interaction for an improved performance of the IR‐organ are discussed. J. Morphol. 275:991–1003, 2014. © 2014 Wiley Periodicals, Inc.     

Space allocation in Melanophila knoteki knoteki (Reitt.) var. hellenica (Obenberger) (Col., Buprestidae) in the attack of Greek fir [Abies cephalonica Loud. var. graeca (Fraas) Liu]: a pattern to process approach

The phloeo‐cambiophagous buprestid Melanophila knoteki knoteki (Reitt.) var. hellenica (Obenberger.) is not a primary factor of fir decline problem although the beetle substantially contributes to Greek fir Abies cephalonica Loud. var. graeca (Fraas) Liu mortality. By using mapping depiction of the exit holes of the insect on a set of fir trees located on a line transect in a randomized point‐centred quarter scheme and employing pattern analysis techniques we were able to reveal various scales of the infestation pattern. Four scales were recognized, two of them corresponding to the pattern of microsite selection on the bark of a fir tree. While the dispersed exit holes exhibited a statistically significant random dispersion on the bark, within each aggregation the pattern was uniform. The area of compartments created by Dirichlet partition approximated very well the sizes of the actual larval galleries. The Dirichlet tessellation of the bark space and the analysis of the parameter of the resulting partitions showed the predominance of the hexagonal conformation of the larval spaces when space was limited. When some exit holes were positioned close together it was found that they were directed away from each other so the resulting galleries were well separated. Several hypotheses are presented as to the mechanisms underpinning the observed patterns. The allocation of space is in accordance with the widely accepted ‘central place theory’ of W. Christaller, a general theory of pattern generated in the geographical dispersion of human settlements. The revealed pattern was also in accordance with the predictions of the theory of ‘central place foraging’ of R. H. MacArthur and the theory of ‘resource concentration hypothesis’ of R. Root.     

红缘吉丁（鞘翅目： 吉丁虫科）触角感器的扫描电镜观察

对红缘吉丁Buprestis fairmairei Thery触角感器的超微结构、类型和分布进行了研究。结果表明,红缘吉丁触角具有3种刺形感器、2种毛形感器、4种锥形感器和1种芽孢感器。触角感器在每节上的数量和分布不同。     

Revision of the aerea group of Chrysobothris (Coleoptera: Buprestidae)

Abstract. The aerea group of the genus Chrysobothris Eschscholtz, as characterized for the first time, includes eight species which occur in the south-western United States and in Mexico. The complicated taxonomic history and confused identity of some species is discussed in depth. Four Mexican species are redescribed in detail from type material: aerea Chevrolat, quadriplagiata Water-house, simplex Waterhouse, stellifera Waterhouse. Lectotypes are designated for the last two species. C.costifrons Waterhouse is considered to comprise three subspecies; two, C.c.rubiterga and C.c.baja are newly described. The remaining species in the group are C.bispinosa, chalcophoroides and serripes . Illustrations and keys to species and subspecies are provided.     

Lehr's fields of campaniform sensilla in beetles (Coleoptera): Functional morphology. II. Wing reduction and the sensory field

Loss of the flight ability and wing reduction has been reported for many taxa of Coleoptera. If elytra are closed, their roots are clenched between the tergum and the pleuron, forces applied to the elytra can not be transmitted to the field of campaniform sensilla situated on the root. That is why it is plausible to assume that the field becomes redundant in non-flying beetles. We examined the relationships between the hind wing reduction and characters of this mechanosensory field in beetles of six families. We measured the size of the elytron, that of the hind wing and counted the number of sensilla in the sensory field. Mesopterous non-flying beetles retain one half to one third of sensilla present in macropterous species of the same body size. Further reduction of the sensory field in brachypterous species is obvious, but sensilla are still present in insects with strongly reduced wings, as long as their elytra are separable and mesothoracic axillaries are present. Complete loss of sensilla coincides with the existence of a permanent sutural lock. However, some beetles with permanently locked elytra and absence of axillaries still retain few campaniform sensilla. A very special case of an extreme wing modification in feather-wing beetles is considered. No sensilla were revealed either on the root of the elytron or on the basal segment of such fringed wings in flying ptiliid species.     

柑桔爆皮虫卵巢发育动态

本研究利用室内饲养成虫,对柑桔爆皮虫的蛹及正常取食和交配、正常取食但未交配以及正常交配但未取食三种处理的雌成虫的卵巢发育进行了系统观察。结果显示：该虫具有1对卵巢,每侧有5根卵巢管。前两种处理的雌虫卵巢管均能正常发育,成虫寿命在30天左右; 没有取食的雌虫寿命只有7天左右,在其卵巢管的生长区和成熟区均无卵形成。根据卵巢的形状、卵的产生过程、卵巢萼内有无卵粒以及卵黄沉积情况等将卵巢发育程度分为6个级别,即发育初期（0级）、卵黄沉积前期(Ⅰ级)、卵黄沉积期(Ⅱ级)、成熟待产期(Ⅲ级)、产卵盛期(Ⅳ级)和产卵末期(Ⅴ级)。每头雌虫最高怀卵量在140粒左右,根据雌虫怀卵量变化趋势,推测正常取食和交配的柑桔爆皮虫雌虫在出孔后10天左右开始产卵,产卵历期可达22天左右。据此提出该虫卵巢管发育到成熟待产期(Ⅲ级)之前（即羽化出孔后10天之内）为出孔成虫的防治适期。     

Lehr's fields of campaniform sensilla in beetles (Coleoptera): Functional morphology. I. General part and allometry

In this first of three articles we show the construction of the articular part of the elytron, the root. The root bears a conspicuous field of campaniform sensilla. This field was studied using light and scanning electron microscopes. The diversity of shape of the field among beetles, types of orientation of elongated sensilla within the field, individual variability of their number among conspecifics are demonstrated. Elongated sensilla point to the junction of the elytron with the second axillary plate. Presumably, they monitor twist movement in this junction, which is possible if the elytron is open. The goal of the whole project is to reveal the effect of both structure and function of the hind wings and elytra on the morphology of this mechanosensory field. Our data on allometric relationships between the animal size and quantitative characteristics of the field in normally flying beetles provide an important background for further functional analysis of this sensory organ.We selected 14 series of several species belonging to the same taxon but differing in size from big to small. It is revealed that the area of the sensory field is directly proportional to the elytral area, whereas the number of sensilla is proportional to the square root of the elytral area. Despite the great range in the elytral area (1500 times) in series of selected species the area of an external pit or cap of a single sensillum varies only 25-fold. The density of sensilla per unit area of the sensory field increases with decrease of the elytral area.     

Revision of the umbellatarum species group of Anthaxia (Coleoptera: Buprestidae)

Abstract. The umbellatarum species group of the genus Anthaxia Eschscholtz of the Mediterranean region is characterized. A key is provided to distinguish this group from some superficially similar groups occurring in the same region, and to the five species and one subspecies referable to the group. Two new species, scylla and hamata are described. A.inculta ab. boissyi Obenberger is recognized as a distinct species and a lectotype is designated. A.domina Abeille is made a subspecies of umbellatarum (Fabricius). A.inculta ab. karamani Obenberger is found to be a colour form of cichorii (Olivier), not of umbellatarum as previously regarded. The other species belonging to the group is lusitanica Obenberger. All species belonging to the group are described.     

Morphology of a thermosensitive multipolar neuron in the infrared organ of Merimna atrata (Coleoptera, Buprestidae)

Two pairs of infrared (IR) organs are situated ventrolaterally on the second and third abdominal sternites of the Australian fire beetle Merimna atrata (Buprestidae). In ventral view, each IR organ has a round IR absorbing area under which a sensory complex is attached to the epidermis. The main component of the complex is a single large multipolar neuron and its mass of highly branched dendrites. All parts of this neuron are enveloped in glial cells. The proximal primary dendrites, which arise from the soma, finally branch into several hundred tightly packed terminal dendrites, which contain many mitochondria. We term this unusual morphology of the dendritic region a terminal dendritic mass (TDM). Additionally, two chordotonal organs were found in each sensory complex. Their somata are integrated in the complex and the dendrites extend to the periphery of the absorbing area. The bauplan of the dendritic region is reminiscent of the thermosensitive trigeminal nerve fibers innervating the absorbing structures in the IR receptors in boid and crotalid snakes. Because this multipolar neuron also functions as a thermoreceptor, another example of a functional analogy between insect and vertebrate sensory systems could be demonstrated.     

Contribution to the knowledge of the Jewel Beetles (Coleoptera: Buprestidae) of the Near East

Abstract

This paper provides faunistic information about almost a hundred different species and subspecies of Jewel Beetles (Coleoptera: Buprestidae) including the subfamilies Julodinae, Acmaeoderinae, Polycestinae, Chalcophorinae and Buprestinae (except the genus Anthaxia) which have been found in the Near East. It also gives a survey of the literature concerning the Jewel Beetles of this region. There are photographs of l8 species. As the author intends to continue the list, he is interested in further information and offers assistance with identifications.     

Attachment forces of the hemelytra-locking mechanisms in aquatic bugs (Heteroptera: Belostomatidae)

Combined hemelytra-locking system of Heteroptera, consisting of several locking mechanisms, aids the mechanical stabilisation of the body at rest, resists external loads, and keeps air stored with the option to easily unlock hemelytra prior to flight. The resistance to unlocking of the hemelytron was measured (in mN) with the aid of a load cell force transducer combined with a three-axial micromanipulator. It is shown that macro- and microstructural features of several submechanisms are responsible for their directionality. The highest resistance to unlocking was measured in lateral and dorsal directions. Summarised force of separately measured submechanisms was considerably lower than the force measured in the combined mechanism. Each submechanism is optimised for achieving high resistance to the hemelytron uncoupling in particular direction(s) and to be easily unlocked in another direction. It was demonstrated in the high-speed videorecordings that hemelytra uncoupling is promoted by their short anterior displacement.