The peregrine falcon's rapid dive: on the adaptedness of the arm skeleton and shoulder girdle.

During a dive, peregrine falcons (Falco peregrinus) can reach a velocity of up to 320 km h- 1. Our computational fluid dynamics simulations show that the forces that pull on the wings of a diving peregrine can reach up to three times the falcon's body mass at a stoop velocity of 80 m s- 1 (288 km h- 1). Since the bones of the wings and the shoulder girdle of a diving peregrine falcon experience large mechanical forces, we investigated these bones. For comparison, we also investigated the corresponding bones in European kestrels (Falco tinnunculus), sparrow hawks (Accipiter nisus) and pigeons (Columba livia domestica). The normalized bone mass of the entire arm skeleton and the shoulder girdle (coracoid, scapula, furcula) was significantly higher in F. peregrinus than in the other three species investigated. The midshaft cross section of the humerus of F. peregrinus had the highest second moment of area. The mineral densities of the humerus, radius, ulna, and sternum were highest in F. peregrinus, indicating again a larger overall stability of these bones. Furthermore, the bones of the arm and shoulder girdle were strongest in peregrine falcons.

The impact of infrared radiation in flight control in the Australian "firebeetle" Merimna atrata.

Infrared (IR) receptors are rare in insects and have only been found in the small group of so-called pyrophilous insects, which approach forest fires. In previous work the morphology of the IR receptors and the physiology of the inherent sensory cells have been investigated. It was shown that receptors are located on the thorax and the abdomen respectively and show an astounding diversity with respect to structure and the presumed transduction mechanism. What is completely missing, however, is any behavioral evidence for the function of the IR receptors in pyrophilous insects. Here we describe the responses of the Australian "firebeetle", Merimna atrata to IR radiation. Beetles in a restrained flight were laterally stimulated with IR radiation of an intensity 20% above a previously determined electrophysiological threshold of the IR organs (40 mW/cm2). After exposure, beetles always showed an avoidance response away from the IR source. Reversible ablation experiments showed that the abdominal IR receptors are essential for the observed behavior. Tests with weaker IR radiation (11.4 mW/cm2) also induced avoidance reactions in some beetles pointing to a lower threshold. In contrast, beetles were never attracted by the IR source. Our results suggest that the IR receptors in Merimna atrata serve as an early warning system preventing an accidental landing on a hot surface. We also tested if another fire specific stimulus, the view of a large smoke plume, influenced the flight. However, due to an unexpected insensitivity of the flying beetles to most visual stimuli results were ambiguous.

Form-function relationships in dragonfly mandibles under an evolutionary perspective.

Functional requirements may constrain phenotypic diversification or foster it. For insect mouthparts, the quantification of the relationship between shape and function in an evolutionary framework remained largely unexplored. Here, the question of a functional influence on phenotypic diversification for dragonfly mandibles is assessed with a large-scale biomechanical analysis covering nearly all anisopteran families, using finite element analysis in combination with geometric morphometrics. A constraining effect of phylogeny could be found for shape, the mandibular mechanical advantage (MA), and certain mechanical joint parameters, while stresses and strains, the majority of joint parameters and size are influenced by shared ancestry. Furthermore, joint mechanics are correlated with neither strain nor mandibular MA and size effects have virtually play no role for shape or mechanical variation. The presence of mandibular strengthening ridges shows no phylogenetic signal except for one ridge peculiar to Libelluloidea, and ridge presence is also not correlated with each other. The results suggest that functional traits are more variable at this taxonomic level and that they are not influenced by shared ancestry. At the same time, the results contradict the widespread idea that mandibular morphology mainly reflects functional demands at least at this taxonomic level. The varying functional factors rather lead to the same mandibular performance as expressed by the MA, which suggests a many-to-one mapping of the investigated parameters onto the same narrow mandibular performance space.

Age-dependent male mating tactics in a spider mite-A life-history perspective.

Males often fight with rival males for access to females. However, some males display nonfighting tactics such as sneaking, satellite behavior, or female mimicking. When these mating tactics comprise a conditional strategy, they are often thought to be explained by resource holding potential (RHP), that is, nonfighting tactics are displayed by less competitive males who are more likely to lose a fight. The alternative mating tactics, however, can also be explained by life-history theory, which predicts that young males avoid fighting, regardless of their RHP, if it pays off to wait for future reproduction. Here, we test whether the sneaking tactic displayed by young males of the two-spotted spider mite can be explained by life-history theory. We tested whether young sneaker males survive longer than young fighter males after a bout of mild or strong competition with old fighter males. We also investigated whether old males have a more protective outer skin-a possible proxy for RHP-by measuring cuticle hardness and elasticity using nanoindentation. We found that young sneaker males survived longer than young fighter males after mild male competition. This difference was not found after strong male competition, which suggests that induction of sneaking tactic is affected by male density. Hardness and elasticity of the skin did not vary with male age. Given that earlier work could also not detect morphometric differences between fighter and sneaker males, we conclude that there is no apparent increase in RHP with age in the mite and age-dependent male mating tactics in the mite can be explained only by life-history theory. Because it is likely that fighting incurs a survival cost, age-dependent alternative mating tactics may be explained by life-history theory in many species when reproduction of old males is a significant factor in fitness.

Morphological properties of the last primaries, the tail feathers, and the alulae of Accipiter nisus, Columba livia, Falco peregrinus, and Falco tinnunculus.

We investigated the mechanical properties (Young's modulus, bending stiffness, barb separation forces) of the tenth primary of the wings, of the alulae and of the middle tail feathers of Falco peregrinus. For comparison, we also investigated the corresponding feathers in pigeons (Columba livia), kestrels (Falco tinnunculus), and sparrowhawks (Accipiter nisus). In all four species, the Young's moduli of the feathers ranged from 5.9 to 8.4 GPa. The feather shafts of F. peregrinus had the largest cross-sections and the highest specific bending stiffness. When normalized with respect to body mass, the specific bending stiffness of primary number 10 was highest in F. tinnunculus, while that of the alula was highest in A. nisus. In comparison, the specific bending stiffness, measured at the base of the tail feathers and in dorso-ventral bending direction, was much higher in F. peregrinus than in the other three species. This seems to correlate with the flight styles of the birds: F. tinnunculus hovers and its primaries might therefore withstand large mechanical forces. A. nisus has often to change its flight directions during hunting and perhaps needs its alulae for this maneuvers, and in F. peregrinus, the base of the tail feathers might need a high stiffness during breaking after diving.

Concept of an Active Amplification Mechanism in the Infrared Organ of Pyrophilous Melanophila Beetles.

Jewel beetles of the genus Melanophila possess a pair of metathoracic infrared (IR) organs. These organs are used for forest fire detection because Melanophila larvae can only develop in fire killed trees. Several reports in the literature and a modeling of a historic oil tank fire suggest that beetles may be able to detect large fires by means of their IR organs from distances of more than 100 km. In contrast, the highest sensitivity of the IR organs, so far determined by behavioral and physiological experiments, allows a detection of large fires from distances up to 12 km only. Sensitivity thresholds, however, have always been determined in non-flying beetles. Therefore, the complete micromechanical environment of the IR organs in flying beetles has not been taken into consideration. Because the so-called photomechanic sensilla housed in the IR organs respond bimodally to mechanical as well as to IR stimuli, it is proposed that flying beetles make use of muscular energy coupled out of the flight motor to considerably increase the sensitivity of their IR sensilla during intermittent search flight sequences. In a search flight the beetle performs signal scanning with wing beat frequency while the inputs of the IR organs on both body sides are compared. By this procedure the detection of weak IR signals could be possible even if the signals are hidden in the thermal noise. If this proposed mechanism really exists in Melanophila beetles, their IR organs could even compete with cooled IR quantum detectors. The theoretical concept of an active amplification mechanism in a photon receptor innervated by highly sensitive mechanoreceptors is presented in this article.

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.

A model for µ-biomimetic thermal infrared sensors based on the infrared receptors of Melanophila acuminata.

Beetles of the genus Melanophila acuminata detect forest fires from distances as far as 130 km with infrared-sensing organs. Inspired by this extremely sensitive biological device, we are developing an IR sensor that operates at ambient temperature using MEMS technology. The sensor consists of two liquid-filled chambers that are connected by a micro-fluidic system. Absorption of IR radiation by one of these chambers leads to heating and expansion of a liquid. The increasing pressure deflects a membrane covered by one electrode of a plate capacitor. The micro-fluidic system and the second chamber represent a fluidic low-pass filter, preventing slow, but large pressure changes. However, the strong frequency dependence of the filter demands a precise characterization of its properties. Here, we present a theoretical model that describes the frequency-dependent response of the sensor based on material properties and geometrical dimensions. Our model is divided into four distinct parts that address different aspects of the sensor. The model describes the frequency-dependent behaviour of the fluidic filter and a thermal low-pass filter as well as saturation effects at low frequencies. This model allows the calculation of optimal design parameters, and thereby provides the foundation for the development of such a sensor.

Bimodal innervation of the infrared organ of Merimna atrata (Coleoptera, Buprestidae) by thermo- and mechanosensory units.

The pyrophilous Australian "fire-beetle"Merimna atrata approaches forest fires and possesses abdominal infrared (IR) organs. Each round IR organ is centrally innervated by a sensory complex showing two different units: one thermoreceptive multipolar neuron and one mechanosensitive chordotonal organ (CO) consisting of two scolopidia. We investigated the CO and found that the scolopidia are mononematic (the scolopale cap remains below the cuticle) and monodynal (one sensory cell per scolopidium). The dendrites of the scolopidia extend anteriorly and are attached by their caps to the cuticle about in the middle of the absorbing area. Structural features at the site of innervation suggest that the CO measures minute thermal deformations caused by IR absorption. Therefore, an additional photomechanic component which has been described for the IR receptors of pyrophilous jewel beetles of the genus Melanophila can be proposed for the IR organ of Merimna. Because scolopidia can measure displacements in the subnanometer range, the CO may enhance the sensitivity of the IR organ. The sensory complex of the Merimna IR organ shows the same units and similar cuticular modifications as the tympanal organs of some noctuid moths. Therefore, a parallel evolution of insect ears and the Merimna IR organ is discussed.

Material properties of photomechanical infrared receptors in pyrophilous Melanophila beetles and Aradus bugs.

Jewel beetles of the genus Melanophila and some pyrophilous species of the flat bugs genus Aradus show a pyrophilous behaviour and have developed so-called photomechanical infrared (IR) receptors. In a spherical photomechanical IR sensillum incoming IR radiation is converted into micromechanical action, finally stimulating the dendritic tip of a mechanosensitive sensory cell. The tip is located inside a tiny cuticular sphere with a diameter of about 12 µm. The material properties of the different cuticular components of this sphere are of great importance for stimulus generation. We measured the modulus and hardness of the outer exocuticular shell of the sphere and the mesocuticle inside the core. Measurements were made by nanoindentation at sensilla which were partly cut open under dry as well as under rewetted (i.e. quasi-natural) conditions. We found that in the rewetted sensilla the outer exocuticular shell of the sphere in the Melanophila sensillum is about 50% harder and 20% stiffer than reference exocuticle, and that in both species especially the rewetted mesocuticle of the inner core of the IR sensilla is significantly softer (about 80% in Melanophila) and more compliant (about 90% also in Melanophila) than the reference mesocuticle. The findings can be interpreted as special adaptations of the cuticular microdomains of photomechanical infrared sensilla to enhance thermomechanical performance and, thereby, sensitivity.

Modelling a historic oil-tank fire allows an estimation of the sensitivity of the infrared receptors in pyrophilous Melanophila beetles.

Pyrophilous jewel beetles of the genus Melanophila approach forest fires and there is considerable evidence that these beetles can detect fires from great distances of more than 60 km. Because Melanophila beetles are equipped with infrared receptors and are also attracted by hot surfaces it can be concluded that these infrared receptors are used for fire detection.The sensitivity of the IR receptors is still unknown. The lowest threshold published so far is 0.6 W/m(2) which, however, cannot explain the detection of forest fires by IR radiation from distances larger than approximately 10 km. To investigate the possible sensitivity of the IR receptors we assumed that beetles use IR radiation for remote fire detection and we made use of a historic report about a big oil-tank fire in Coalinga, California, in 1924. IR emission of an oil-tank fire can be calculated by "pool fire" simulations which now are used for fire safety and risk analysis. Assuming that beetles were lured to the fire from the nearest forests 25 and 130 km away, our results show that detection from a distance of 25 km requires a threshold of the IR receptors of at least 3×10(-2) W/m(2). According to our investigations most beetles became aware of the fire from a distance of 130 km. In this case the threshold has to be 1.3×10(-4) W/m(2). Because such low IR intensities are buried in thermal noise we suggest that the infrared sensory system of Melanophila beetles utilizes stochastic resonance for the detection of weak IR radiation. Our simulations also suggest that the biological IR receptors might be even more sensitive than uncooled technical IR sensors. Thus a closer look into the mode of operation of the Melanophila IR receptors seems promising for the development of novel IR sensors.

Infrared receptors in pyrophilous ("fire loving") insects as model for new un-cooled infrared sensors.

Beetles of the genus Melanophila and certain flat bugs of the genus Aradus actually approach forest fires. For the detection of fires and of hot surfaces the pyrophilous species of both genera have developed infrared (IR) receptors, which have developed from common hair mechanoreceptors. Thus, this type of insect IR receptor has been termed photomechanic and shows the following two special features: (i) The formation of a complex cuticular sphere consisting of an outer exocuticular shell as well as of a cavernous microfluidic core and (ii) the enclosure of the dendritic tip of the mechanosensitive neuron inside the core in a liquid-filled chamber. Most probably a photomechanic IR sensillum acts as a microfluidic converter of infrared radiation which leads to an increase in internal pressure inside the sphere, which is measured by a mechanosensitive neuron.A simple model for this biological IR sensor is a modified Golay sensor in which the gas has been replaced by a liquid. Here, the absorbed IR radiation results in a pressure increase of the liquid and the deflection of a thin membrane. For the evaluation of this model analytical formulas are presented, which permits the calculation of the pressure increase in the cavity, the deformation of the membrane and the time constant of an artificial leak to compensate ambient temperature changes. Some organic liquids with high thermal expansion coefficients may improve the deflection of the membrane compared to water.

Water as a major modulator of the mechanical properties of insect cuticle.

The mechanical properties of the sternal cuticle of the locust were investigated by nanoindentation. Modulus and hardness of the exo-, meso-, and endocuticular layers were locally measured under dry and fully wetted conditions in the normal (i.e. perpendicular to the outer surface) as well as in the transverse direction (i.e. parallel to the alignment of the respective layers). The results show that water has a major impact on the mechanical properties of all layers. After drying the endocuticle, in particular, became harder by a factor of up to 9 and stiffer by a factor of up to 7.4. Additionally the gradual decrease in hardness and Young's modulus from the outer exo- to the inner endocuticle, characteristic of native cuticle, was eliminated or even reversed in dried cuticle. A pronounced anisotropy was revealed in all layers when comparing data obtained by probing in the normal (lower values) vs. probing in the transverse direction (higher values). Cyclic drying and rewetting of the endocuticle showed that the mechanical properties can be reproducibly changed by altering the water content. Based on our results we propose a new role of the epicuticle: fine-tuning of the mechanical properties of the different cuticular layers can be accomplished by setting the local cuticular transpiration.

Distribution and functional morphology of photomechanic infrared sensilla in flat bugs of the genus Aradus (Heteroptera, Aradidae).

Globally the flat bug genus Aradus comprises about 200 species. About half a dozen Aradus species can be primarily found on burnt areas and, therefore, have been called pyrophilous. Bugs and their offspring feed on fungi growing on burnt wood. Recently, prothoracic infrared (IR) receptors have been described in the pyrophilous Australian species Aradus albicornis. In our study we investigated 10 Aradus species, once again including A. albicornis, and found prothoracic as well as hitherto unknown mesothoracic IR sensilla in A. albicornis, Aradus lugubris and Aradus fuscicornis. In Aradus flavicornis only prothoracic IR receptors were found. Currently the latter two species are not known as pyrophilous. However, there is considerable evidence that these flat bugs also approach forest fires. In all four species where IR receptors were identified, the dome-shaped IR sensilla look very similar. An IR sensillum consists of an internal exocuticular sphere reinforced by consecutive layers of chitin fibres. In the center of the sphere, a microfluidic core is located which consists of a cup-shaped plug of cuticle and an underlying fluid filled annular channel surrounding the tip of the dendrite of a mechanosensitive neuron. Like the IR receptors of buprestid beetles of the genus Melanophila, the IR sensilla found in Aradus species can be classified as photomechanic IR receptors.

Micromechanical properties of consecutive layers in specialized insect cuticle: the gula of Pachnoda marginata (Coleoptera, Scarabaeidae) and the infrared sensilla of Melanophila acuminata (Coleoptera, Buprestidae).

Insect cuticle is a highly adaptive material that fulfils a wide spectrum of different functions. Cuticle does not only build the exoskeleton with diverse moveable parts but is also an important component of a stunning variety of mechanosensory receptors. Therefore, the mechanical properties of these specialized cuticular systems are of crucial importance. We studied the different cuticular layers of the head part (gula) of the head-to-neck ball articulation of Pachnoda marginata and of the photomechanic infrared (IR) sensilla of Melanophila acuminata on the basis of cross sections. In our study, we combined histological methods (i.e. detection of the different types of cuticle by specific staining) with measurements of hardness (H) and reduced elastic modulus (E(r)) by nanoindentation technique. In the gula of Pachnoda we found an unusual aberrance from the well-known layering. Between the epi- and exocuticle, two meso- and one endocuticular layers are deposited which are softer and more elastic than the underlying exo- and mesocuticular layers. The hardest of all examined materials is the cuticle of the exocuticular shell of the internal sphere of the Melanophila IR sensillum with H=0.53GPa whereas the inner mesocuticular core of the sensillum represents the most elastic and softest layer with values of H=0.29GPa and E(r)=4.8GPa. Results are discussed with regard to the proposed functions.

Microfluidic photomechanic infrared receptors in a pyrophilous flat bug.

Infrared (IR) receptors are so far known only in boid and crotalid snakes and in three genera of pyrophilous beetles that seek out forest fires. Pyrophilous insects can also be found in other orders, however, so it can be hypothesised that IR receptors also occur in some of these species. We investigated the pyrophilous Australian flat bug Aradus albicornis and found a small number of dome-shaped sensilla (diameter 13 microm) on the prothorax, which have previously not been described. Ultrastructural investigations revealed that the sensilla are characterised by a fluid-filled inner compartment enclosed in a round cuticular shell. The cuticular apparatus is innervated by the dendrite of a ciliary mechanoreceptor, which is fluidically coupled to the inner compartment. Electrophysiological recordings demonstrated that the sensilla respond to brief warming by red laser light or to broadband IR radiation. Depending on the radiation intensity (4.4-549 mW/cm(2) tested, threshold measured as 11.3 mW/cm(2)), first spike latencies varied between 3.4 and 7.5 ms. Thus, our findings demonstrate that A. albicornis most probably possesses photomechanic IR sensilla resembling the metathoracic IR sensilla of buprestid beetles of the genus Melanophila. In the Melanophila sensillum, IR radiation causes thermal expansion of a fluid, which rapidly deforms the dendritic membrane of a mechanosensory cell. The existence of photomechanic IR receptors in both beetles and bugs demonstrates a remarkable convergent evolution towards this particular biophysical transduction mechanism and suggests that it provides selective advantages over other possible solutions.

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.

Electrophysiological characterisation of the infrared organ of the Australian "Little Ash Beetle" Acanthocnemus nigricans (Coleoptera, Acanthocnemidae).

This study characterises the response properties of the sensilla located on the prothoracic disc organ of the beetle Acanthocnemus nigricans, such as intensity response functions and temporal coding properties. Warming the sensilla by a red laser accelerated their ongoing spiking activity, cessation of the stimulus suppressed their firing as revealed by extracellular recordings. Convective heat sources also increased sensillum activity, but stimuli of other modalities failed to elicit responses. The response threshold was between 11 and 25 mW/cm2 and latencies ranged between 20 and 40 ms. Repeating stimuli with frequencies between 5 and 20 Hz were reliably resolved by the sensilla. This temporal resolution enables the disc sensilla to represent behaviourally relevant changes in heat stimuli in a thermally patchy environment. These findings complement our knowledge on the sensory physiology of pyrophilous insects by hinting at two different, elementary orientation strategies evolved in the three pyrophilous beetle species described. A. nigricans seems to be best adapted to short-range orientation on freshly burnt areas.

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.

A new type of insect infrared organ of low thermal mass.

The Australian beetle Acanthocnemus nigricans is attracted by forest fires and has a pair of complex infrared (IR) receptor organs on the first thoracic segment. Each organ consists of a tiny sensory disc (diameter 120-130 microm) which serves as an absorbing structure for IR radiation. The disc is arranged above an air-filled cavity which is located just anteriorly to the coxae of the prothoracic legs. Inside the disc, about 30 multipolar thermoreceptors (warmth receptors) are tightly attached to the cuticle which is directed to the outside. The many dendrites of each multipolar neuron are tightly wrapped around the soma and contain a large number of mitochondria. Absorption of IR radiation by the disc causes an increase in temperature which is measured by the warmth receptors. Therefore, the IR receptors of A. nigricans can be classified as microbolometers with reduced thermal mass and in principle can be compared to the IR organs of pit vipers.