Cyclic release of carbon dioxide accompanied by abdominal telescoping movements in forager ants of Formica polyctena (Hymenoptera, Formicidae)

Abstract.  Discontinuous gas exchange and abdominal telescoping movements were investigated in intact forager ants of Formica polyctena Förster. For simultaneous recording of discontinuous gas exchange and abdominal movements, an electrolytic differential microrespirometer was combined with an infrared-optical device using an infrared-sensor diode commonly applied as a cardiograph in insect studies. The air in the insect chamber was saturated with water vapour, and an ant was contacted with a sliver of wet filter paper. Accordingly, the ants used in the experiments were regarded as being hydrated. The enforced immobility of the ant in the respirometry chamber (0.15 mL volume) induced continuous struggling during the first hours of measurement but, after 3–4 h, the activity periods alternated with quiescent periods when regular cycles of discontinuous gas exchange (8.97 mHz) were displayed. After remaining in the chamber for a day, the ants showed such cycles (8.15 mHz) without any interruption by periods of activity. The cyclic release of carbon dioxide (burst) was accompanied by a bout of telescoping movements of the abdominal segments (rapid contractions and slow relaxations), which were interpreted as active ventilation. The irregular and rare abdominal contractions, recorded during the interburst periods on the first day of measurements, were regarded as a symptom of stress because these disappeared on the second day. In approximately 20% of foragers, the telescoping movements during the interburst periods were characterized by very rapid (0.09 s) protractions and slow retractions of the abdominal segments, obviously acting as inspiration movements. The results do not support the hypothesis that discontinuous gas exchange is an adaptation for conserving water in ants.     

Discontinuous gas exchange in the fire ant,Solenopsos invicta Buren: Caste differences and temperature effects

The discontinuous gas exchange cycle (DGC), the cyclic release of CO(2) and uptake of O(2), were investigated in workers and female and male alates of the red imported fire ant, Solenopsis invicta Buren, using real-time CO(2) emission measurement by flow-through respirometry. All S. invicta castes displayed discontinuous emission of CO(2) in the temperature range of 15-25 degrees C, but only male alates and workers exhibited a DGC at 30 degrees C. The closed (C) and flutter (F) periods of the DGC were distinguishable in alates of both sexes at the lowest temperature, but not clearly differentiated in females at temperatures above 15 degrees C, in males above 20 degrees C, or workers at any temperature. DGC frequency increased for all castes as temperature increased, ranging from a low of 0.9+/-0.05 mHz (male alates at 15 degrees C) to 18+/-0.79 mHz (workers at 30 degrees C). O period (or burst) volumes of all castes decreased as temperature increased, and increased with body mass - this mass effect was most pronounced at lower temperatures. Q(10) values for DGC frequency (4.27, 5.81, and 5.62 for workers, female and male alates, respectively) were high compared with Q(10)'s for standard Vdot;(CO(2)). Differences in the salient characteristics of the DGC between castes are presented and discussed, and S. invicta DGC patterns are compared to known values for some other ant species.     

Respiratory physiology and water relations of three species of Pogonomyrmex harvester ants (Hymenoptera: Formicidae)

The respiratory physiology and water relations of three harvester ant species (Pogonomyrmex rugosus Emery, P. occidentalis[Cresson] and P. californicus[Buckley]) were examined at three temperatures (15, 25 and 35°C) using a flow-through respirometry system. As intact ants tended to be active during testing, we performed a parallel set of experiments on individuals rendered motionless by decapitation. Both intact and decapitated ants exhibited discontinuous ventilation. Decapitation caused metabolic rate (V˙CO₂) and burst frequency to decrease in all three species. Burst volume either remained constant or increased after removal of the head, though mass-specific V˙CO₂ was unaffected except in P. rugosus. Mass-specific V˙CO₂s of headless harvesters were comparable with published values derived from motionless specimens of other ant species. The mean Q₁₀ for intact ants of all three species was 2.37; for decapitated insects the mean was 2.32. Respiratory water constituted a small (< 5%) fraction of total loss, and we believe that discontinuous ventilation does not act to conserve water in these organisms, although it may serve other functions.     

Neural Control of Gas Exchange Patterns in Insects: Locust Density-Dependent Phases as a Test Case

The adaptive significance of discontinuous gas exchange cycles (DGC) in insects is contentious. Based on observations of DGC occurrence in insects of typically large brain size and often socially-complex life history, and spontaneous DGC in decapitated insects, the neural hypothesis for the evolution of DGC was recently proposed. It posits that DGC is a non-adaptive consequence of adaptive down-regulation of brain activity at rest, reverting ventilatory control to pattern-generating circuits in the thoracic ganglia. In line with the predictions of this new hypothesis, we expected a higher likelihood of DGC in the gregarious phase of the desert locust (Schistocerca gregaria, Orthoptera), which is characterized by a larger brain size and increased sensory sensitivity compared with the solitary phase. Furthermore, surgical severing of the neural connections between head and thoracic ganglia was expected to increase DGC prevalence in both phases, and to eliminate phase-dependent variation in gas exchange patterns. Using flow-through respirometry, we measured metabolic rates and gas exchange patterns in locusts at 30°C. In contrast to the predictions of the neural hypothesis, we found no phase-dependent differences in DGC expression. Likewise, surgically severing the descending regulation of thoracic ventilatory control did not increase DGC prevalence in either phase. Moreover, connective-cut solitary locusts abandoned DGC altogether, and employed a typical continuous gas exchange pattern despite maintaining metabolic rate levels of controls. These results are not consistent with the predictions of the neural hypothesis for the evolution of DGC in insects, and instead suggest neural plasticity of ventilatory control.     

Gas exchange pattern transitions in the workers of the harvester termite

The evolutionary genesis and the current adaptive significance of the use of the discontinuous gas exchange cycle (DGC) for respiration by insects is the subject of intense debate. Years of research have resulted in several leading hypotheses, one of which is the emergent-property hypothesis. This hypothesis states that DGC is an emergent property or consequence of interactions between the O₂ and CO₂ set points that regulate spiracular function, i.e. opening and closing. Workers of the harvester termite, Hodotermes mossambicus were selected as a model to test this hypothesis. The respiratory patterns of major workers, investigated using flow-through respirometry, were obtained at 100% relative humidity (RH) under varying temperature to evaluate the assumptions of the emergent-property hypothesis. Metabolic rate, measured as VCO₂ increased significantly after 15 °C. As VCO₂ increased in response to increasing temperature and activity, the gas exchange pattern displayed by workers transitioned to a continuous gas exchange. A true DGC, defined as showing all three phases and a coefficient of variation value close to 2, was not expressed under the experimental conditions. The conclusion drawn from this study of termite workers is that changes in respiratory patterns are most likely an emergent property of the insects’ nervous and respiratory system.     

Reactive oxygen species production and discontinuous gas exchange in insects

While biochemical mechanisms are typically used by animals to reduce oxidative damage, insects are suspected to employ a higher organizational level, discontinuous gas exchange mechanism to do so. Using a combination of real-time, flow-through respirometry and live-cell fluorescence microscopy, we show that spiracular control associated with the discontinuous gas exchange cycle (DGC) in Samia cynthia pupae is related to reactive oxygen species (ROS). Hyperoxia fails to increase mean ROS production, although minima are elevated above normoxic levels. Furthermore, a negative relationship between mean and mean ROS production indicates that higher ROS production is generally associated with lower . Our results, therefore, suggest a possible signalling role for ROS in DGC, rather than supporting the idea that DGC acts to reduce oxidative damage by regulating ROS production.     

Effects of Neem EC on gas exchange, tracheal ventilation, and water loss in diapausing pupae of Pieris brassicae

Effects of Neem EC (The Indian Neem Tree Company^TM, 1% azadirachtin) on gas exchange cycles, tracheal ventilation, and water loss in diapausing pupae of the large white butterfly, Pieris brassicae L. (Lepidoptera: Pieridae), were studied using a constant volume respirometer combined with an infrared probe actograph. The non‐treated pupae displayed discontinuous gas exchange cycles (DGC) with a trend coinciding with the bursts of carbon dioxide (CO₂) release, active tracheal ventilation, and the heartbeat periods. Two independent forms of tracheal ventilation were observed, relatively vigorous abdominal shaking movements and weak abdominal pulsations. The ability to respond to mechanical excitation with abdominal movements was entirely lost on the 2nd day after treatments with Neem EC, and also a reduced tendency to use a DGC was observed. During 2–3 days after treatments, the DGCs and gas exchange microcycles were entirely lost, as was active ventilation. Before treatments, body mass loss, that is, water loss, was 0.6–0.9 mg g^?1 day^?1. After the treatments, water loss increased to 3–5 mg g^?1 day^?1. The pupae remained alive for 10–15 days after the treatments and died after having lost about 50% of their initial body mass. The absence of heartbeats measured during at least 4–5 h was the main criterion for ascertaining death of pupae. The results suggested that respiratory failures, that is, the loss of cyclic gas exchange, evoked by Neem EC were the primary cause of lethal desiccation. Thus, the hypothesis that the cyclic gas exchange is an adaptation for restricting water losses in insects was supported.     

The use of the anaesthetic, enflurane, for determination of metabolic rates and respiratory parameters in insects, using the ant, Camponotus maculatus (Fabricius) as the model

This study investigated the effects of the anaesthetic, enflurane, on metabolic rates and ventilation patterns in the spotted sugar ant, Camponotus maculatus, using flow-through respirometry. The standard metabolic rate was not affected by the anaesthetic. While the ants were anaesthetised they exhibited a similar discontinuous gas exchange cycle to that observed when they were voluntarily motionless, but their spiracles remained open for a longer time during the open or burst phase even though the amount of CO(2) emitted during this phase remained constant. We discuss this finding in the context of the central nervous system control of the spiracle muscle. For both the determination of standard metabolic rate and ventilation patterns the individual ant has to be motionless. From this study we recommend the use of enflurane to ensure immobility in ants, and other small active insects, during the determination of standard metabolic rates, but the anaesthetic cannot be used to quantify the respiration pattern.     

Monitoring of gas exchange cycles and ventilatory movements in the pine weevil Hylobius abietis: respiratory failures evoked by a botanical insecticide

The large pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae), is the most important insect pest of young coniferous plants. The implementation of new control methods requires not only a profound knowledge of the ecology and behaviour of the pest, but particularly of its physiology. Standard metabolic rate (SMR) and discontinuous gas exchange cycles (DGCs) were recorded in parallel with abdominal ventilation movements in adults of H. abietis using a differential electrolytic respirometer‐actograph. Quiescent weevils displayed DGCs of the constriction, flutter, and ventilation phases of the CFV type, while bursts of carbon dioxide were always accompanied by abdominal pumping movements, i.e., muscular ventilation in the closed subelytral cavity (SEC). In some beetles the C phase was absent and thus (C)FV cycles were recorded. In addition, at the beginning and often at the end of a burst, the SEC was rhythmically opened and closed by movements of the last abdominal segments. Continuous pumping movements and an absence of DGCs were signs of stress imposed by handling or by a new environment, even if the beetle was not moving. All individuals showed clear DGCs after recovering from handling and apparatus stress lasting 2–3 h. The results show that in the monitoring of DGCs, it is essential to determine whether they are of the constriction, flutter, and open phases (CFO), or the CFV subtype of the constriction, flutter, and burst (CFB) cycles. Use of our simple closed‐system respirometer enables non‐invasive simultaneous recording of SMR, oxygen uptake, DGCs, and active ventilation in H. abietis and other beetles. The topical application of adult H. abietis with sublethal doses of a botanical insecticide, NeemAzal T/S, caused essential respiratory failures: cyclic gas exchange was lost and irregular pumping movements appeared. In the treated beetles normal DGCs did not resume.     

Pump out the volume—The effect of tracheal and subelytral pressure pulses on convective gas exchange in a dung beetle, Circellium bacchus (Fabricus)

Many flightless beetles like the large apterous dung beetle Circellium bacchus, possess a subelytral cavity (SEC) providing an extra air space below the elytra which connects to the tracheal system (TS) via metathoracic and abdominal spiracles. By measuring subelytral and intratracheal pressure as well as body movements and gas exchange simultaneously in a flow-through setup, we investigated the contribution of convection on Circellium respiratory gas exchange.No constriction phase was observed. TS and SEC pressures were always around atmospheric values. During interburst phase open abdominal spiracles and a leaky SEC led to small CO₂-peaks on a continuous CO₂ baseline, driven by intermittent positive tracheal pressure peaks in anti-phase with small negative subelytral pressure peaks caused by dorso-ventral tergite action.Spiracle opening was accompanied by two types of body movements. Higher frequency telescoping body movements at the beginning of opening resulted in high amplitude SEC and TS pressure peaks. High frequency tergite movements caused subelytral pressure peaks and led to a saw tooth like CO₂ release pattern in a burst. We propose that during the burst open mesothoracic spiracles increase the compliance of the subelytral cavity allowing big volumes of tracheal air being pulled out by convection.     

Periodic abdominal pumping supports leg development during metamorphosis in tenebrionid beetle Zophobas atratus

Rhythmic abdominal pumping movements in a pupa of giant mealworm beetle Zophobas atratus caused large hemolymph pressure pulses of approximately 20 mmHg. The abdominal pumping movements were completely blocked by transecting the ventral nerve cord (VNC) between the first and second abdominal ganglia. Transection of the VNC until 2 days after pupation caused a developmental defect of adult legs: morphogenesis of the tibial and tarsal segments was severely retarded, and the segments remained covered with a thick pupal cuticle. The developmental defect was rescued by artificially inducing rhythmic abdominal bending for 3 days after transection of VNC. Blocking of the abdominal pump did not increase the amount of water loss during the pupal period. The transplanted tibial segments lacking active tracheal ventilation could form a thick adult cuticle. The results suggest that abdominal pumping movements during the pupal period support the development of adult legs by facilitating hemolymph circulation.     

Body movements and their role as triggers of heartbeats in pupae of the Colorado potato beetle Leptinotarsa decemlineata

Rhythmic body movements and their role as triggers of intermittent heartbeats were studied in pupae of Leptinotarsa decemlineata Say. Heartbeats and body movements were recorded simultaneously by means of an optical method (infra‐red cardiography) combined with respirometry. IR‐cardiography allows heartbeats and body movements to be distinguished on the basis of their different rates (40–80 min^?1 and 4–8 min^?1, respectively) and amplitude. In the mid‐interecdysial period, abdominal movements in the pupae were always accompanied by heart activity beginning after the first 5–12 abdominal strokes. Simultaneous periods of abdominal movements and heartbeats lasted 2–5 min, while the intervening pause ranged from 40 to 72 min at 24°C. Experiments of forced heart activity showed that a slight external tactile stimulus (prodding once with a single hair), applied prior to an expected heartbeat bout, evoked abdominal movements followed soon afterwards by contractions of the dorsal vessel. Repeated prodding with a hair evoked body rotating movements (1–3 strokes) at any time with heartbeats starting at the first movement. We surmise that rhythmic body movements play an active role in the triggering of heart activity in pupae according to the principle of cardiac reflex response. This is a possible mechanism for synchronizing periods of heart activity with periods of rhythmic body movements. Haemolymph circulation in pupae is favoured when heartbeat bouts are accompanied by active body movements.     

The activity of abdominal stretch receptors during non-giant swimming in the crayfish<Emphasis Type="Italic"> Cherax destructor</Emphasis> and their role in hydrodynamic efficiency

Recordings were made from the nerve innervating the stretch receptors of the abdominal muscle receptor organs and slow extensor muscles of tethered crayfish, Cherax destructor, during so-called non-giant swimming. The stretch receptors were active during the flexor phase of swimming but the duration and pattern of activity varied from cycle to cycle. Their pattern of firing was modified by the activity of the large accessory neurons which make direct inhibitory synapses upon them. Neither the stretch receptors nor the accessory neurons were active during the extensor phase of the cycle. The timing and extent of tailfan movements during the period of stretch receptor activity were measured from video records before and after the stretch receptor nerves were cut in the second to fifth segments. The promotion of the tailfan during flexion was significantly delayed and the minimum angle to which the uropods were remoted at the end of flexion significantly larger in denervated animals. We propose that afferent information from the stretch receptors coordinates the timing and extent of tailfan movements according to variations in the positioning and movement of the abdominal segments such that the hydrodynamic efficiency of the tailfan is enhanced on a cycle by cycle basis during non-giant swimming.Abbreviations A# abdominal segment number - Acc accessory neuron - LUU large unidentified unit - MRO muscle receptor organ - NGS non-giant swimming - SEMN slow extensor motor neuron - SR stretch receptor neuron     

Patterns of nocturnal movement of the long-spined sea urchin<Emphasis Type="Italic"> Diadema antillarum</Emphasis> (Philippi) in Gran Canaria (the Canary Islands,central East Atlantic Ocean)

We tagged individuals of the sea urchin Diadema antillarum (Philippi) around the island of Gran Canaria (The Canary Islands) during winter 2001–2002 using a new technique, consisting of the insertion of a hook fastened to a fishing line into the aboral pole (periproctal membrane). This allowed individual identification of tagged sea urchins. The goals were: (1) to quantify nocturnal movements and the homing behaviour of this echinoid on shallow rocky bottoms, and (2) to assess short term spatial and temporal variability of these movements. Tagged sea urchins displayed clear homing behaviour. The mean distance travelled at night was 3.7±1.2 m (range 1.0–5.1 m). Mean speed of nocturnal movement was 33±26 cm h^–1 (range 5–110 cm h^–1). We observed greater movement at midnight than at the beginning and the end of the night.Communicated by H.-D. Franke     

Efficacy of four density gradient separation media to remove erythrocytes and nonviable sperm from canine semen

The objective was to compare four commercially available density gradient centrifugation (DGC) media (ISolate [Irvine Scientific; Santa Ana, CA, USA], Percoll [Pharmacia; Uppsala, Sweden], PureCeption [SAGE In-Vitro Fertilization, Inc.; Trumbull, CT, USA], PureSperm 100 [Nidacon International AB; Molndal, Sweden]) for their ability to separate viable, motile sperm from contaminant nonviable (immotile and/or dead) sperm and red blood cells (RBC). Pooled sperm-rich fractions from four healthy dogs were assessed using Spermvison SAR (Minitube of America). For this, 1 mL of the blood/sperm admixture was pipetted over 4 mL of DGC media: 50%/90% ISolate (Irvine Scientific), 45%/90% Percoll (Pharmacia), 40%/80% PureCeption (SAGE In-Vitro Fertilization, Inc.), and 40%/80% PureSperm 100 (Nidacon International AB). After centrifugation, five 1-mL fractions (A, B, C, D, and E) and the sperm pellet (bottom fraction F) were separated. Sperm morphology and red blood cell/sperm ratio (RBC/S) per fraction were determined on stained slides. All DGC media separated RBC from sperm; the highest red blood cell/sperm ratio was present in ISolate (Irvine Scientific) and Percoll (Pharmacia) fraction A (29.4 ± 29.7 and 28.2 ± 20.8, respectively), and in fractions A and B of both PureCeption (SAGE In-Vitro Fertilization, Inc.) (37.0 ± 22.8 and 39.6 ± 24.3, respectively) and PureSperm 100 (Nidacon International AB) (25.2 ± 5.9 and 23.0 ± 3.9, respectively). The fractions with the highest total sperm recovery, motile sperm recovery, as well as overall motility were ISolate (Irvine Scientific) and Percoll (Pharmacia) fraction D (33.9 ± 29.4%; 40.99 ± 27.9%; 71.2 ± 21.8% and 36.4 ± 14.5%; 39.3 ± 15.8%; 88.6 ± 2.3%, respectively), and for PureCeption (SAGE In-Vitro Fertilization, Inc.) and PureSperm 100 (Nidacon International AB), the sperm pellet, fraction F (78.8 ± 28.3%; 88.0 ± 17.4%; 70.2 ± 11.1% and 73.1 ± 21.0%; 75.4 ± 24.6%; 80.6 ± 17.1%, respectively). In the pellet for PureCeption (SAGE In-Vitro Fertilization, Inc.), more sperm and motile sperm were recovered than in ISolate (Irvine Scientific) and Percoll (Pharmacia) fractions D (P < 0.0163). Therefore, DGC media should be considered for canine semen purification when contaminated with blood or when separation of motile versus immotile sperm is needed.     

Multisegmental motor activity in the segmentally restricted gin trap behavior in Manduca sexta pupae

Stimulation of sensory neurons innervating hairs in the gin traps on the abdomen of Manduca sexta pupae evokes a rapid bending of the abdomen that is restricted to one or more of the three articulating posterior segments. However, electrical stimulation of the gin trap sensory nerve in an isolated abdominal nerve cord evokes characteristic motor neuron activity in every abdominal segment. To determine if the segmentally distributed motor activity also occurred in intact animals and how it contributed to the segmentally restricted reflex movement, mechanical stimulation of the sensory hairs in intact animals was used to evoke reflex responses that were recorded as electromyograms synchronized with video recordings of the behavior. Motor activity was monitored during movements to determine if there was activity in many segments when the movement was restricted to one segment. Coordinated muscle activity was evoked throughout the abdomen in response to stimulation of any of the three gin traps, even when movement was restricted to one segment. Differences in the timing of ipsilateral and contralateral motor activity among segments allowed the closing of gin traps to be segmentally restricted. These findings suggest that the neural circuit underlying the gin trap reflex is distributed throughout the abdominal nerve cord. This network generates a complex, yet coordinated, motor pattern with muscular activity in many abdominal segments that produces a localized bending reflex. Accepted: 10 January 1997     

Cyclic carbon dioxide release in the dampwood termite, Zootermopsis nevadensis (Hagen)

Real-time traces of CO₂ release of pseudergates of the dampwood termite, Zootermopsis nevadensis (Hagen) were obtained using flow-through respirometry. Traces were made at each of six temperatures, between 10 and 35°C. Termites released CO₂ in a cyclic pattern at each of the six temperatures. CO₂ release rate (as in ml h⁻¹) increased significantly with temperature and body mass. Rate of change in with temperature (or Q₁₀) was 2.11. Degree of cycling in CO₂ traces was estimable using the coefficient of variability. Coefficient of variability for both acyclic and cyclic traces declined exponentially with increasing temperature.     

The unique pollination ofLeporella fimbriata (Orchidaceae): Pollination by pseudocopulating male ants (Myrmecia urens,Formicidae)

Leporella fimbriata is a self compatible orchid of southern Australia. It is dependant across its range on unique pollination by sexually attracted male winged antsMyrmecia urens, which pseudocopulate with the flower. Typical pollination sequences began with an initial circling then zig-zag flight to the flower. Vectors usually alighted on the inflorescence stem and quickly crawled to the flower where they adopted a copulatory position sideways along the wide labellum, pseudocopulatory probing immediately followed. In this position pollen carried on the thorax was deposited on the stigma. Departure from the labellum usually resulted in pollinium removal. Pollinator movements were restricted and the distribution leptokurtic with a mean of 3.141 ± 4.59 m. Pollination was widespread but variable from site to site and season to season with a maximum of 70% of all flowers being pollinated. Pollinator limitation is indicated. Traits essential for this pollination interaction include the coincidence of orchid and ant geographic distributions and the coincidence of flowering with the flight period of the ant. The production of pheromonelike substances and the distinctive floral morphology are also essential for attraction and manipulation of male ants. The ant mating system which the orchid can exploit is also important.     

Food Shortage Disrupts Social Organization: The Case of Red Squirrels in Conifer Forests

A recent conceptual model of spatial organization in vertebrates, based upon changes in home range overlap with habitat quality, ‘the space-use model’, predicts large and strongly overlapping home ranges and absence of territorial behaviour in habitats with poor food availability. We investigated whether the model can be extended to predict intra-population variation in space-use in a habitat with strong temporal variation in resource abundance. We studied space use of Eurasian red squirrels (Sciurus vulgaris) in subalpine conifer forests over 4 years with strong fluctuations in seed production. Norway spruce dominated forests are resource limited habitats for squirrels, and tree seed availability is gradually depleted in the year following a poor seed-crop. Male and female squirrels responded to spruce seed-crop failure in 2000 by strongly increasing their home ranges and core-areas in the following year (mean MCP ± SE in summer and autumn, respectively: 2001, 122.9±14.1 ha and 84.3±9.4 ha, against 43.0±6.4 ha and 20.8±3.8 ha in 2002, and 22.7±1.5 ha and 21.4±2.0 ha in 2003). In 2001, half of the animals, those with the poorest quality pre-dispersal home ranges, emigrated to areas with more larch. Residents had multi-nuclear core-areas. Also, intra-sexual core-area overlap (males by males, females by females) was higher in 2001 and summer 2002, than in autumn 2002 and in 2003 (means ± SE in summer and autumn, respectively: males by males, 2001, 52±8 and 44±7%, 2002, 80±30 and 15±11%, 2003, 37±19 and 26±14%; females by females 2001, 52±10 and 112±32%, 2002, 55±27 and 0±0%, 2003, 12±8 and 0.1±0%). Red squirrels responded to food shortage by moving to patches with other food resources and abandoned the spacing pattern of reduced core-area overlap among males and nearly exclusive core-areas among females, found in less variable habitats. After richer seed-crops in 2001 and 2002, it took squirrels about a year to reduce the size of their home ranges and core-areas and return to a spacing pattern of stable home ranges and intra-sexual territoriality of adult females. These results are consistent with the space-use model and show that spacing behaviour in red squirrels is a plastic, conditional strategy with individuals adapting the size and/or location of their home ranges in relation to local distribution and abundance of food resources.Co-ordinating editor: R. Härdling     

Ventilatory movements of the abdomen and branchial apparatus in dragonfly larvae (Odonata: Anisoptera)

Use of cine-photography in the analysis of ventilatory movements has necessitated a reappraisal of the ventilatory patterns previously described in dragonfly larvae. Movements of the abdominal exoskeleton, monitored by cine-photography, phototransistor and strain gauge, are discussed in relation to known muscular activity. Internal movements of the branchial apparatus, which consists of two chambers and three valves, are also illustrated in the intact animal. The mechanism of "gulping" ventilation ( V _a) in dragonfly larvae is discussed.