Skin carotenoid concentration correlates with male hunting skill and territory quality in the kestrel Falco tinnunculus

In the context of sexual selection carotenoid based signals are candidates for indicator traits: they have to be taken up in the diet by animals, they can indicate the ability of the bearer to accumulate a limited resource, and they help in maintaining the health status. We investigated the yellow-orange colouration of the tarsi of the kestrel Falco tinnunculus in relation to sex, diet and to different aspects of male reproductive behaviour. The colouration of the tarsi (estimated as hue) was more intense in males than in females. Among males, the tarsi hue was associated with the intake of invertebrates; this was true also if the population diet was based mainly on voles. Carotenoid based colouration was positively associated with the number of vertebrate preys delivered to the nest per time unit and with territory quality (calculated on the basis of home-range size, habitat extension and prey availability). These results are consistent with predictions derived from good-parent models of sexual selection, suggesting that in the common kestrel carotenoid based colouration is important as an indicator of male quality.     

The problem of the number of tarsomeres in the regenerated cockroach leg.

There are 5 tarsomeres in the normal cockroach leg, but this number is often reduced in regenerated legs. In order to examine this complicated situation, fore-, mid-, and hindlegs of German cockroaches were amputated at 11 different tarsal levels and at 18 different times during the last instar. When tarsi were amputated at or proximal to the 3rd tarsomere, 4-segmented tarsi regenerated. When legs were amputated distal to the 3rd tarsomere, the regenerated tarsi had 5 segments. Three-segmented tarsi rarely regenerated when legs were amputated proximal to 3rd tarsomere and in the latter half of the instar period. The lengths of all tarsomeres of regenerated tarsi were measured together with those of unoperated contralateral tarsomeres, and the ratios of the former to the latter were calculated. The ratios ranged from 28 to 138% for the various tarsomeres and levels of amputation. From a comparison of the ratios and morphological observations, it was suggested that the 3rd tarsomere of the normal 5-segmented tarsus has disappeared in the regenerated 4-segmented tarsus. Pads and disto-lateral spines of tarsomeres were observed on unoperated and regenerated tarsi. It was of interest that double spines were often found on the 4-segmented tarsi, mostly on the 2nd tarsomere, just proximal to the position of the missing 3rd tarsomere. This observation supported the idea that the 3rd tarsomere has not simply disappeared, but has probably fused with the 2nd tarsomere.     

Oviposition behaviour and the sensilla of the ovipositor of Chilo partellus and Spodoptera littoralis (Lepidoptera: Noctuidae).

Observations of oviposition behaviour by Chilo partellus revealed that antennae, ovipositor tip and possibly tarsi were all involved in the choice of oviposition site. Among a very large number of typical mechanoreceptor hairs on the tip of the ovipositor valves, two pairs of stouter, blunt-tipped hairs were discovered. These hairs alone took up silver nitrate stain and transmission electron microscope sections revealed that the hair shaft had a hollow lumen within which ran a group of four or five dendrites. It is concluded that these sensilla are contact chemoreceptors and they are positioned in such a way that, with the placing of each egg, they would contact the oviposition substrate. Similar hairs were found on the ovipositor of Spodoptera littoralis . It is suggested that the hairs may prevent oviposition on surfaces directly chemically harmful to the eggs.     

Ecological and morphological variation of Canary Island blue tits, Parus caeruleus (Aves: Paridae)

Canary Island blue tits have differentiated within the last million years. Compared with mainland populations they have short wings, large beaks and tarsi, and tend to be less variable in these parameters. Within the archipelago the eastern populations have the shortest wings and tarsi but the widest and deepest beaks. The eastern populations may have been derived from central island ones, following extinction of the original eastern populations during a warm period in the last Ice Age. New measurements have been used to test contrasting predictions of hypotheses that explain inter-island morphological variation in terms of either climatic or ecological influences. Neither hypothesis is sufficient to explain the variation. A model which combines mem is proposed as follows. Mild winter temperatures have selected for small body size and wing length, especially on the hot, dry, eastern islands. On the other pine-forested, islands, selection has favoured relatively long tarsi and long, slender, beaks for efficient foraging among pine foliage. The absence of die pine-dwelling coal tit from the islands has facilitated mis evolutionary shift. It is shown that blue tits in pines forage more in the needles and cones on the Canary Islands than they do on die North African mainland.     

The Immediate Source of the Oviposition-Deterring Pheromone Produced by Larvae of Adalia bipunctata (L.) (Coleoptera, Coccinellidae)

As is the case for other insects ovipositing on or in resources that are limited in time and/or space, the two-spot ladybird beetle, Adalia bipunctata (L.) produces an oviposition-deterring pheromone (ODP), which is produced by the larval stages. Foraging larvae touch the substrate with their tarsi and the anal disk on the tenth abdominal segment. The aim of this paper was to determine whether the ODP produced by larvae was deposited by the tarsi or the anal disk. Fourth instar larvae either had their anal disk and tarsi, or anal disk, or tarsi coated with a water-soluble mounting medium. Larvae so treated were allowed to walk on filter paper that was subsequently presented to gravid females. The tracks of larvae that had both their tarsi and anal disk masked did not inhibit oviposition. However, the tracks of larvae that had only their tarsi masked significantly inhibited oviposition but those of larvae that had only their anal disk masked did not. It is concluded that the ODP is deposited on the substrate by the anal disk on the tenth abdominal segment of larvae.     

Insect walking techniques on thin stems

The attachment ability of insects on surfaces are associated not only with the micro- and nanostructure of the adhering part of an attachment device, but also with the global scale kinematics responsible for contact formation and release. In the present study, the locomotory techniques of several representatives of insects from four different orders (Orthoptera, Heteroptera, Coleoptera, and Hymenoptera), possessing different types of attachment structures, are described. The study is based on video recordings of insects walking on a flat surface and on cylindrical rods of various thickness, imitating plant stems. Attachment devices of tarsi and pretarsi were visualized using Scanning Electron Microscopy. The results show a different manner in the use of adhesive structures on substrates with various curvatures. Insects bearing attachment pads on proximal tarsomeres usually touch flat and curved substrates using all tarsomeres, whereas insects with their attachment devices on the distal tarsomeres usually walk on flat surfaces using the distal tarsomeres of the overextended tarsus. On substrates, with diameters comparable to or larger than the tarsus length, insects walk above the stem by clasping the stem with the bent tarsi. On thin stems, insects clasp the stem between their tarsi and hang under the stem. Thus, on thin and thick rods, forces applied to attachment organs act in opposite directions. There are two methods of leg positioning for walking on a rough flat substrate. In the first case, the tarsus is straightened and the rough substrate is gripped between the claws and the proximal complex of attachment devices (tarsal euplantulae, fossulae spongiosa, and terminal spurs of tibiae). In the second case the tibia does not touch the substrate; the insect is supported only by distal tarsomeres. The tarsus is in an overextended condition. On rods, with diameters comparable to or larger than the tarsus length, insects walk by clasping the stem with the bent tarsi. This posture is characteristic for the majority of insects independent of the tarsal position they normally use while walking on a plane. If the rod’s diameter is smaller than the tarsus length, walking insects usually clutch it between contralateral tarsi. Using such a posture they are supported by interlocking or by strong friction, generated by attachment devices of the proximal tarsomeres, and do not use attachment devices of the pretarsus. Contact with the substrate is reinforced due to the coordinated contralateral clutch using all supporting legs. It is concluded that the use of different types of attachment structures correlates with locomotory techniques. Handling Editor: Heikki Hokkanen     

Behavioural changes and the adaptive diversification of pigeons and doves

What factors determine the extent of evolutionary diversification remains a major question in evolutionary biology. Behavioural changes have long been suggested to be a major driver of phenotypic diversification by exposing animals to new selective pressures. Nevertheless, the role of behaviour in evolution remains controversial because behavioural changes can also retard evolutionary change by hiding genetic variation from selection. In the present study, we apply recently implemented Ornstein–Uhlenbeck evolutionary models to show that behavioural changes led to associated evolutionary responses in functionally relevant morphological traits of pigeons and doves (Columbiformes). Specifically, changes from terrestrial to arboreal foraging behaviour reconstructed in a set of phylogenies brought associated shorter tarsi and longer tails, consistent with functional predictions. Interestingly, the transition to arboreality accelerated the rates of evolutionary divergence, leading to an increased morphological specialization that seems to have subsequently constrained reversals to terrestrial foraging. Altogether, our results support the view that behaviour may drive evolutionary diversification, but they also highlight that its evolutionary consequences largely depend on the limits imposed by the functional demands of the adaptive zone.     

Transduction ion channels directly gated by sugars on the insect taste cell

Insects detect sugars and amino acids by a specialized taste cell, the sugar receptor cell, in the taste hairs located on their labela and tarsi. We patch-clamped sensory processes of taste cells regenerated from the cut end of the taste hairs on the labelum of the flashfly isolated from the pupa approximately 20 h before emergence. We recorded both single channel and ensemble currents of novel ion channels located on the distal membrane of the sensory process of the sugar receptor cell. In the stable outside-out patch membrane excised from the sensory processes, we could repeatedly record sucrose-induced currents for tens of minutes without appreciable decrease. An inhibitor of G-protein activation, GDP-beta-S, did not significantly decrease the sucrose response. These results strongly suggested that the channel is an ionotropic receptor (a receptor/channel complex), activated directly by sucrose without mediation by second messengers or G protein. The channel was shown to be a nonselective cation channel. Analyses of single channel currents showed that the sucrose-gated channel has a single channel conductance of approximately 30 pS and has a very short mean open time of approximately 0.23 ms. It is inhibited by external Ca(2+) and the dose-current amplitude relation could be described by a Michaelis-Menten curve with an apparent dissociation constant of approximately 270 mM. We also report transduction ion channels of the receptor/channel complex type directly gated by fructose and those gated by L-valine located on the sensory process.     

The chemosensilla on tarsi of Locusta migratoria (Orthoptera: Acrididae): Distribution,ultrastructure, expression of chemosensory proteins

The chemosensilla on the tarsi of Locusta migratoria were mapped using light microscopy, as well as scanning and transmission electron microscopy. Only chemosensilla chaetica were found on the tarsi. On the basis of their ultrastructure, these can be grouped into three main subtypes: short, long, and sunken sensilla chaetica. Short sensilla chaetica can be further divided into two groups containing 6 or 7 neurons. Long sensilla chaetica are mainly located on the lateral surface of the tarsi. Short sensilla chaetica were mainly found on the dorsal surface of the tarsi. Sunken sensilla chaetica were only found on the ventral surface, such as the pulvilli and arolium. Immunocytochemical localization of chemosensory protein (CSP) was performed on ultrathin sections of chemosensilla on tarsi. The antiserum against LmigCSP‐II intensively labeled all three types of sensilla chaetica. Gold granules were concentrated in the outer sensillum lymph surrounding the dendrite sheath, while the inner sensillum lymph containing dendrite branches was never labeled. Massive labeling with the anti‐LmigCSP‐II was also found in cuticle of the pulvilli on the ventral surface of tarsi. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Pollen‐feeding in hover‐flies (Diptera: Syrphidae)

Analyses of the pollen contents of the crop and intestine of 11 species of New Zealand Syrphidae . showed that small, sparsely haired hover‐flies with unbranched hairs, short, simple bristles, and a short proboscis had ingested at least 99% anemophilous pollens, and that larger, more hairy hover‐flies with pollen‐collecting hairs, long, spirally grooved bristles, and elongate mouthparts had ingested pollens almost exclusively from nectar‐bearing flowers. Pollen‐feeding behaviour was studied in one hairy species, the drone‐fly Eristalis tenax, and in one sparsely‐haired species, Melanostoma fasciatum. Using granulated charcoal as a substitute for pollen, it was found that in E. tenax particles trapped among the body hairs are combed off by the front and hind tibiae and transferred to pollen‐retaining bristles on the front and hind tarsi respectively. Particles retained among the front tarsal bristles are ingested directly from the bristles. Those retained by the hind tarsi are transferred in flight by leg‐scraping movements to the front tarsi, from which they are subsequently eaten. E. tenax also eats pollen directly from anthers. In M. fasciatum apparently all the pollen ingested is taken directly from anther lobes or stigmas. The few pollen grains that adhere to the body of this species are combed off by the front and hind tibiae and transferred to the front and hind tarsi, but are not retained there because the bristles are short and simple. The mouthparts, hairs, and bristles of E. tenax and M. fasciatum are illustrated. Drawings of leg movements associated with pollen collection and ingestion, and photographs showing leg scraping in E. tenax are included. Morphological similarities between drone‐flies and honey‐bees, previously regarded as the result of mimicry, can be explained by convergent evolution in response to similar food‐gathering behaviour. Probably the majority of Syrphidae, and also the related Acroceridae, collect pollen by means of branched or curly‐tipped hairs.     

Motor patterns during kicking movements in the locust

Locusts (Schistocerca gregaria) use a distinctive motor pattern to extend the tibia of a hind leg rapidly in a kick. The necessary force is generated by an almost isometric contraction of the extensor tibiae muscle restrained by the co-contraction of the flexor tibiae (co-contraction phase) and aided by the mechanics of the femoro-tibial joint. The stored energy is delivered suddenly when the flexor muscle is inhibited. This paper analyses the activity of motor neurons to the major hind leg muscles during kicking, and relates it to tibial movements and the resultant forces.During the co-contraction phase flexor tibiae motor neurons are driven by apparently common sources of synaptic inputs to depolarized plateaus at which they spike. The two excitatory extensor motor neurons are also depolarized by similar patterns of synaptic inputs, but with the slow producing more spikes at higher frequencies than the fast. Trochanteral depressors spike at high frequency, the single levator tarsi at low frequency, and common inhibitors 2 and 3 spike sporadically. Trochanteral levators, depressor tarsi, and a retractor unguis motor neuron are hyperpolarized.Before the tibia extends all flexor motor neurons are hyperpolarized simultaneously, two common inhibitors, and the levator trochanter and depressor tarsi motor neurons are depolarized. Later, but still before the tibial movement starts, the extensor tibiae and levator tarsi motor neurons are hyperpolarized. After the movement has started, the extensor motor neurons are hyperpolarized further and the depressor trochanteris motor neurons are also hyperpolarized, indicating a contribution of both central and sensory feedback pathways.Variations in the duration of the co-contraction of almost twenty-fold, and in the number of spikes in the fast extensor tibiae motor neuron from 2–50 produce a spectrum of tibial extensions ranging from slow and weak, to rapid and powerful. Flexibility in the networks producing the motor pattern therefore results in a range of movements suited to the fluctuating requirements of the animal.     

Identification and genomic structure of chemosensory proteins (CSP) and odorant binding proteins (OBP) genes expressed in foreleg tarsi of the swallowtail butterfly Papilio xuthus

Chemoreception is a key feature for selection of host plants by phytophagous insects. Female swallowtail butterflies recognize their host plants using chemosensilla present on foreleg tarsi. We constructed a cDNA library of female tarsi and a genome library of Papilio xuthus. We identified 11 chemosensory protein (CSP) genes and three odorant binding proteins (OBP) genes from the cDNA library and eight additional CSP genes from the genome library using the ESTs as probes. A sequence similarity tree of insect CSPs showed that lepidopteran CSPs constructed big branches of the order. Small numbers of CSPs have been identified from the whole genomes of several insect orders which belong to branches separated from those of Lepidoptera. The CSP gene family of Lepidoptera may have diverged in at least two steps, the first on a small scale and the second on a large scale before and after the diversification of insect orders, respectively. Seventeen of 19 CSP genes of P. xuthus clustered in a specific region of the genome, suggesting that they were diversified by gene duplication from a common ancestral gene.     

Spatio-ecological niche segregation of two sympatric species of<Emphasis Type="Italic">Clidemia (Melastomataceae)</Emphasis> in Western Amazonian non-flooded rainforests

The frequent occurrence of sympatric series of closely related plant species in tropical rainforests has evoked claims for and against the application of the competitive exclusion principle in these ecosystems. Narrow niche limits defined by biotic as well as abiotic specialization have been reported for sympatric species of the same genus or family. In Amazonian lowland rainforests this question deserves renewed attention because: (1) the existence of edaphically defined community types has recently been well established, and (2) spatio-ecological niche segregation of congeneric species may help explain not only the maintenance of the high Amazonian alpha-diversity, but also its origin through sympatric ecological speciation. In this study, the morphology, ecology, and distribution patterns of two species,Clidemia epiphytica andC. longifolia (Melastomataceae), from western Amazonia, were analyzed. The aims were to find out whether they really are two distinct taxonomic species and if so, whether they also can be considered biological species; if the species are sympatric; and if they are ecologically specialized. The results showed that the morphological variation of the species seems continuous, but that they exhibit opposite morphological responses to variation in soil cation concentration, which suggests that they also are separate biological species. Furthermore, the species occur sympatrically but in different habitats. It is suggested that a part of the enigma of sympatric congeners in rainforests may be explainable by spatial segregation stemming from ecological specialization in relation to subtle environmental variation. It is hypothesized that the studied species are a good candidate case of sympatric speciation driven by ecological specialization.     

Ecogeographic variation in body size and shape of Cape sparrows (Passer melanurus) in southern Africa

Canonical discriminant analyses were used to assess whether four populations of Cape sparrows varied in body size and shape according to predictions from Bergmann's Rule. In accordance with Bergmann's Rule, birds from two hot, arid Namib desert sites (Namib 1 and Namib 2) were smaller than birds from two cool, mesic Transvaal sites. If heat dissipation through extremities (tarsi) is important to reduce water lost from evaporative cooling, birds under hot conditions in dry environments (Namib 2) should have longer tarsi than those in more humid hot environments (Namib 1). Contrary to this, males at Namib 2 had relatively longer wings hut shorter tarsi than at Namib 1, and female relative tarsus length did not vary between desert sites     

Sensing the Plant Surface Prior to Feeding and Oviposition: Differences in External Ultrastructure and Function Among Tarsi of <Emphasis Type="Italic">Heliconius erato</Emphasis>

Adult foretarsi of Heliconius erato Linnaeus (Lepidoptera, Nymphalidae) are reduced in size and are not used for walking. Foretarsi of the female have specialized sensilla that are presumably used to identify the host plant, by drumming. The mid- and hind tarsi also bear sensilla in both sexes, but these have not been described in detail, nor has their chemosensory function been determined. We described and compared the tarsi of H. erato under light and scanning electron microscopy. Behavioral experiments showed that differences in the shape, number, and size of sensilla were related to feeding and oviposition behaviors. Two types of sensillum (chaeticum and trichodeum) were found in similar numbers and size on the mid- and hind tarsi of both sexes. Sensilla on the female foretarsi act in host-plant site selection, strongly affecting oviposition rates when isolated. Male foretarsi lack sensilla, which may have been selected against due to the absence of function and thus lost. Sensilla on the mid- and hind tarsi are involved in sugar detection in both sexes, responding to an effective dose of sucrose (ED₅₀) near 0.01 M, and therefore might be used to identify food resources when the butterflies settle on flowers.     

Mouthpart grooming behavior in honeybees: Kinematics and sectionalized friction between foreleg tarsi and proboscises

The mouthpart of a honeybee is prone to contamination by granular particles such as pollen or dirt from the field. To clean the contaminated mouthparts, a honeybee swings its foreleg tarsi forward and backward to brush the proboscis continuously, sweeping the contaminant from the surfaces of the labial palpi, galeae, and bushy haired tongue (glossa). This grooming behavior has been documented but the dynamic characteristics therein have not been investigated yet. We quantified the grooming behavior of a honeybee from the perspective of kinematic and tribological properties. We captured high-speed videos that recorded the mouthpart grooming patterns of honeybees from the front and side views and measured the friction on the grooming surfaces using a precision dynamometer. During grooming, a honeybee first positions the mouthpart and then places a pair of foreleg tarsi to the tubular-folded galea. The tarsi press the galea and labial palpi and slide downward while keeping close contact with the galea. Then, the hairy glossa stretches out of the temporary tube with the glossa setae erected. The tarsi slowly slide down when grooming the glossa. In the return stroke of grooming, the foreleg tarsi detach from the mouthpart and retreat swiftly. Friction analysis shows that the honeybees can coordinate the velocity of the foreleg tarsi to the sectionalized tribological property of the tarsus–mouthpart interface. The specific grooming pattern enables honeybees to save energy and resist wear, resulting in a possible highly evolved grooming strategy. These findings lead to further understanding of the honeybee’s grooming behavior facilitated by the special motion kinematics and friction characteristics.     

Mechanoresponsive neurones in the suboesophageal ganglion of the locust

Abstract. A preparation is described for intracellular recording from the neur-opile of the sub-oesophageal ganglion (SOG) of the locust, while stimulating the labial and maxillary palps with plant material in such a way as to mimic the palpation behaviour which precedes and continues throughout feeding. Twelve neurones responding to simulated palpation were recorded from and stained in the SOG. Axons of three neurones ascended to the brain, six had descending axons and three had all of their processes confined to the suboesophageal ganglion. The major regions of arborization were in the ventrolateral and mediolateral neuropiles of the maxillary and labial neuromeres. All twelve neurones were solely mechanoresponsive. In addition to responding to palpation of one or more of the four palps, five also responded to stimulation of the labrum, one to touching each antenna, and one to mechanical stimulation of each of the six tarsi. In the context of what is known about the role of mechano-stimulation in the control of feeding, and given their particular patterns of input and arborizations, it is suggested that the neurones may be active during food selection and ingestion.     

Cellular aspects of tolerance. VI. The effect of age on responsiveness and tolerance induceability of SJL mice

New born and 3-week-old SJL mice but not 8–12-week-old animals could be rendered tolerant to rabbit γ-globulin. Animals reconstituted with thymus cells from 12-week-old donors and bone marrow cells from 3-week-old donors showed resistance to tolerance induction. Animals reconstituted with bone marrow cells from 12-week-old animals and thymus cells from 3-week-old donors could be rendered tolerant. Earlier work has shown that tolerance could be induced in older animals, if they were deprived of competent accessory cells. It was suggested that a lesion in the thymus cell population is expressed through the mediation of accessory cells. The possibility of a relation between resistance to tolerance induction and lymphoid malignancies was discussed.     

Comparative morphology of the prothoracic leg in heliconian butterflies: Tracing size allometry,podite fusions and losses in ontogeny and phylogeny

Prothoracic legs of heliconian butterflies (Nymphalidae, Heliconiinae, Heliconiini) are reduced in size compared to mesothoracic and metathoracic legs. They have no apparent function in males, but are used by females for drumming on host plants, a behavior related to oviposition site selection. Here, taking into account all recognized lineages of heliconian butterflies, we described their tarsi using optical and scanning electron microscopy and searched for podite fusions and losses, and analyzed allometry at the static, ontogenetic and phylogenetic levels. Female tarsi were similar, club-shaped, showing from four to five tarsomeres, each bearing sensilla chaetica and trichodea. Male tarsi were cylindrical, formed from five (early diverging lineages) to one (descendant lineages) either partially or totally fused tarsomeres, all deprived of sensilla. Pretarsi were reduced in both sexes, in some species being either vestigial or absent. Tarsal lengths were smaller for males in almost all species. An abrupt decrease in size was detected for the prothoracic legs during molting to the last larval instar at both histological and morphometric levels. In both sexes, most allometric coefficients found at the population level for the prothoracic legs were negative compared to the mesothoracic leg and also to wings. Prothoracic tarsi decreased proportionally in size over evolutionary time; the largest and smallest values being found for nodes of the oldest and youngest lineages, respectively. Our results demonstrate that evolution of the prothoracic leg in heliconian butterflies has been based on losses and fusions of podites, in association with negative size allometry at static, ontogenetic and phylogenetic levels. These processes have been more pronounced in males. Our study provided further support to the hypothesis that evolution of these leg structures is driven by females, by changing their use from walking to drumming during oviposition site selection. In males the leg would have been selected against due to absence of function and thus progressively reduced in size, in association with podites fusions and lost.