Diel patterns of reproductive activities in the tomato fruit fly, Neoceratitis cyanescens

We describe the diurnal temporal and spatial patterns of reproductive activities (mating and ovipositing) of adult tomato fruit flies, Neoceratitis cyanescens (Bezzi) (Diptera: Tephritidae), under laboratory, field‐cage and field conditions. The distribution pattern of flies in the different habitats (host and non‐host plants) showed temporal, physiological and sexual differences. Mature females were observed to frequent host plants preferentially, particularly in the late afternoon. By contrast, immature females preferred to rest on non‐host plants throughout the day. Males frequented mainly non‐host plants but some were observed on host fruits in the morning. Mating activity took place preferentially on non‐host plants, in the morning. In laboratory cages, females showed a strong daily propensity to engage in mating behaviour; the mean percentage of mating females was 88 ± 5%. Time allotted to this activity (4 h 52 min ± 1 h 23 min in laboratory cages and at least 3 h for the mating pairs observed in the field) appeared considerable compared to the time required to satisfy feeding or oviposition activities. Except for a few ovipositing females observed in the morning, oviposition activities occurred mainly in the late afternoon.     

Distribution of cabbage root flies in brassica crops

Changes in the distribution of adult cabbage root flies were determined in 0–05-0-4 ha brassica plots during 1970 and 1971 using yellow water-traps. The three generations of flies occurred at similar times in both years and the numbers caught generally decreased from the first to the third generation. In resident populations, the numbers of males and females declined by 70 and 40% respectively, between the second and third generations. When there was no decline, immigration presumably compensated for flies trapped or otherwise lost. The results confirmed that male cabbage root fly movement is trivial, that wind has little effect on distribution at a brassica plot and that females move into the crop independently of males. They failed to confirm that females aggregate markedly at hedgerows or fences, that they spend little time in the crop or that they regularly return to hedgerows. Differences in the dispersal behaviour of the male and female flies were reflected in the numbers caught. Some males dispersed actively during the first 2 weeks after emergence but many were displaced slowly downwind. Once at a plot, wind appeared to have little effect on distribution of young males, but older males tended to shelter just downwind of the hedge. Relatively few males were caught near hedges but they tended to aggregate along the crop-interfaces, particularly during the first generation. Plot size did not appear to influence the pattern of fly distribution greatly. Females dispersed more actively than males and were generally more uniformly distributed through the crops even during periods of immigration. Gravid females entered an isolated plot without accompanying males, inferring that they are migratory and mate near the site of emergence. Female distribution was not related to the stage of ovarial development except during maximum oviposition when those that had laid some eggs visited the hedgerow. Aged females became progressively easier to capture and usually died in the crop.     

Dispersal of the cabbage root fly

The dispersal rates of wild and culture cabbage root flies Erioischia brassicae were determined in release-recapture experiments at Wellesbourne in 1971–3. The experiments were concerned mainly with the first 7 days of adult life. The flies, released from nine locations in the area, were recaptured in yellow water-traps. Dispersal was affected by wind, rain and the terrain the flies were crossing. The flies least often recaptured were those released into the host crop when 6–12 days old. The results indicated the following pattern of behaviour. Flies moved little during the first 2 days of adult life but by the third day both sexes had dispersed to c. 100 m from the release point. Flies are known to mate about the fourth day and after this the males continued to disperse at c. 100 m per day for the three subsequent days. ‘Wild’ females from field-collected pupae carried out a ‘migratory’ flight, however, and dispersed at c. 1000 m per day during the fifth and sixth days, the days preceding the start of oviposition. Similar rates of dispersal were recorded from flies released across host crop and non-host crop areas. Some females did not stop at the first crop they encountered. The culture females from the laboratory-reared pupae dispersed only c. one-third of the distance of the wild females. There was considerable intermingling of local populations. The percentage recapture of young culture and wild flies released during the pre-oviposition period of this species was 38 ± 4 and 19 ±4 for males, and 15 + 2 and 8+1 for females, respectively. The dispersal range of the cabbage root fly is probably within a 2000–3000 m radius of the site of infestation.     

Feeding behaviour,midgut distension and ovarian development in Phlebotomus papatasi (Diptera: Psychodidae)

Phlebotomus papatasi females were fed through membranes or from cotton wool soaked in blood, water, sucrose or sodium chloride solutions. In membrane-fed flies, all diets were routed to the midgut and not to the crop. Following the meals that went to the midgut, females showed ovarian development at least 3 times greater than in sucrose-fed, autogenous control flies. Neither small quantities of water arriving in the midgut following drinking from soaked cotton wool, nor piercing of a membrane without feeding, stimulated ovarian development. Flies exhibited different feeding behaviour namely, blood feeding, sugar feeding, and water drinking. The blood-feeding behaviour was typical of flies ingesting any of the experimental diets through membranes, or blood or saline from cotton wool. The other two types of behaviour were observed in flies which fed from soaked cotton wool. The type of behaviour was characterized by the depth of penetration of the mouthparts into the substrate, the deployment of the palps and the degree of contact between the palps and the surface. It is suggested that the stimuli which control the routing of meals to the crop or to the midgut are derived from these types of behaviour.     

Movement of adult pecan weevils Curculio caryae within pecan orchards

1 The pecan weevil Curculio caryae (Horn) (Coleoptera: Curculionidae) is an indigenous pest of pecan Carya illinoinensis (Wangenh.) K. Koch, in North America. Understanding the movement of this pest from the orchard floor to host trees could lead to pest management practices that exploit weevil behaviour and thus reduce insecticide application to the entire orchard canopy. Furthermore, no information exists on diel periodicity of pecan weevil movement. 2 Movement of adult pecan weevils crawling and flying to the host trunk, flying to the host canopy, crawling within the host canopy and flying between host trees was studied using four types of passive traps over four seasons. Each type of trap was used to capture weevils at different locations on or near the tree and to discriminate flying versus crawling behaviour. 3 More pecan weevils crawl to the trunk than fly and a proportion of the population flies directly from the orchard floor into the pecan canopy. The majority of this movement occurs at dusk. 4 The vertical distribution of weevils was generally uniform throughout the canopy but more weevils were captured in suspended traps nearest tree tops, rather than traps near the ground, when flying between trees and this was significantly so for two of 4 years. 5 The results of the present study are contrary to previous reports suggesting that most adult pecan weevils fly to the pecan trunk after emergence from the soil; however, our results did indicate that a proportion of the population flies directly from the orchard floor into the pecan canopy and thus would circumvent strategies that attempt to control weevils moving up the trunk.     

The pattern,rate, and range of within‐patch movement of a stem‐galling fly

1. Information on the movement of insects is critical to understanding the spatial spread, dynamics, and genetic structure of their populations, as well as their interactions with other species. With this in mind, the movement behaviour of the stem‐galling fly Eurosta solidaginis Fitch (Diptera: Tephritidae) was investigated. 2. Fluorescent‐marked adults were released at a single location within pure patches of the host plant, tall goldenrod Solidago altissima, and their distributions censused repeatedly throughout the day. 3. Following their release, male and female flies redistributed themselves in a manner that was well described by a simple‐diffusion model. The diffusion rate was independent of fly density and time since flies were released. 4. Female flies dispersed at a significantly faster rate, and therefore farther on average, than males. Based on the diffusion model, it was estimated that at 2.5–3.0 h post release, males and females had a median dispersal distance of only 2.0 and 2.5 m respectively. Furthermore, 95% of the males were estimated to have dispersed no more than 5.9 m, and females no more than 6.4 m. 5. Post‐release censuses suggested that flies were most active during mid morning, disappeared from the site at a rate of 10–15% per hour (most likely due to mortality), and survived for less than 2 days. Based on the rate of spread, diel activity, and liberal estimates of longevity in the field, 50% of the ovipositing females were predicted to have had a maximum lifetime range of movement within a patch of hosts of ≤ 51 m (95% were expected to have been limited to ≤ 130 m). 6. These data are used to assess whether the absence of a positive correlation between host‐plant preference and offspring performance in this system could be due to the limited scale of dispersal of this species relative to the spatial scale at which its oviposition behaviour has been studied.     

The diurnal activity, movement and trypanosome infection rates of Glossina fuscipes fuscipes （Diptera： Glossinidae） in Buvuma Island, Lake Victoria, Uganda

The diurnal activity patterns, trypanosome infection rates and movement of Glossinafuscipesfuscipes （Diptera： Glossinidae） were investigated in Buvuma Island, Lake Victoria, Uganda. Hourly trapping of tsetse flies was undertaken to determine their activity rhythm while a capture-mark-release-recapture method was conducted to assess the movement and dispersal of tsetse flies between lakeshore, hinterland and further inland sites along a transected area. Dissection of tsetse flies was also undertaken to determine the trypanosome infection rates in salivary glands, proboscis and mid-gut. Results indicated a bimodal diurnal activity profile for G. f fuscipes on the Island, both on the lakeshore and in the hinterland. Movement and dispersal of G. f fuscipes tsetse flies occurred between lakeshore, hinterland and further inland sites with a greater tendency of flies to move to the lakeshore. Trypanosome infection rates of 4.32% for Trypasoma vivax and 1.15% for 7. congolense were found in G. f. fuscipes.     

Effect of age and time of day on sex attraction and mating of the face fly, Musca autumnalis

A well-defined sexual attraction exists between mature virgin female and mature unmated male face flies, Musca autumnalis, tested in a Y-tube olfactometer. Attraction seems to be maximum when females are 5 to 6 days old and males are 4 to 8 days old post emergence. Gonadotrophic cycles probably play a rôle in the periodicity of the sexual attraction. Attraction as well as insemination reach a peak during the noon hours, gradually decline through the afternoon, and reach a minimum level in the evening. A daily rhythm of sex pheromone release and/or production is most probably present in virgin female flies.     

Diurnal periodicity in Daphnia reproduction

The diurnal periodicity of reproduction in a reservoir population of Daphnia galeata Sars was tested by two methods (egg age distribution analyses and direct observation of hatching). No significant diurnal synchronization of reproduction in the natural cladoceran population was found. A conspicuous shortage of older embryonic stages was parallely revealed by analyses of diurnal changes of egg age distribution. Suggested selective feeding of fish on gravid females of Daphnia brooding embryos with black eye pigmentation is discussed.     

Flower nitrogen status and populations of Frankliniella occidentalis feeding on Lycopersicon esculentum

Alighting and capture of wild olive fruit flies, Bactrocera oleae (Rossi) (Diptera, Tephritidae), on spheres of seven different colours was studied on Chios island, Greece. The 70-mm-diam plastic spheres, coated with adhesive, were suspended on olive trees. Yellow and orange spheres trapped the greatest number of males while red and black spheres trapped the greatest number of females. White and blue spheres were the least effective for both sexes. Peak captures occurred in the late afternoon and especially around sunset. Since mating takes place in the last hours of the photophase, the increased captures during that period may be related to the sexual behaviour of the fly. When red spheres were assessed against glass McPhail traps baited with 2% ammonium sulphate, which consist a standard tool for monitoring the olive fruit fly in Greece, there were no significant differences in male captures. However, spheres trapped almost three times as many females as McPhail traps. The possible mechanisms underlying colour discrimination, the motivation of alighting flies and the possible use of red spheres for monitoring and controlling B. oleae are discussed.     

Windows of opportunity and the temporal structuring of foraging activity in a desert solitary bee

1. Females of the desert solitary bee Anthophora pauperata collect nectar and pollen almost exclusively from Alkanna orientalis (Boraginaceae). The bee and plant are found together in the early spring, living in the bottom of steep-sided wadis (dry river valleys) at an altitude of 1500 m in Egyptian Sinai. 2. Female A. pauperata showed clear morning and afternoon peaks in foraging activity, separated by a 2–3 h midday period spent in their underground nests. This study analyses the following in order to identify the factors structuring this daily pattern: thermal aspects of the bee and its environment, temporal patterns of resource provision by the plant, and female nectar and pollen foraging behaviour. 3. Although A. pauperata can generate substantial heat endothermically, morning and evening ambient temperatures well below 10 °C defined a thermal window within which foraging occurred. Maximum air temperatures were moderate (25–30 °C), and examination of the physiology and behaviour of A. pauperata suggests that the midday reduction in flight activity was not due to thermal constraints. 4. Alkanna orientalis produces protandrous hermaphroditic flowers. Female A. pauperata collected pollen from male-phase flowers and harvested nectar preferentially from female-phase flowers. Although the nectar standing crop was relatively constant throughout the day, pollen availability peaked strongly in the early afternoon. 5. Female A. pauperata visited young male-phase flowers as soon as they opened, generating an early afternoon peak in pollen foraging activity and depleting the pollen standing crop rapidly. A morning peak in pollen foraging occurred when females gleaned remnant pollen from flowers that had opened the previous day. Pollen availability in the morning was far lower than in the early afternoon, and the time taken to collect a full pollen load in the morning was significantly longer. Collection of pollen in the morning despite very low resource availability suggests that pollen may be a limiting resource for A. pauperata. 6. In contrast to many existing examples of bimodal activity patterns in highly endothermic bees, the bimodal activity patterns of female A. pauperata appear to be driven not by thermal considerations but by daily patterns of pollen release from its principal food source.     

Flower and fruit volatiles assist host‐plant location in the Tomato fruit fly Neoceratitis cyanescens

Mature females of the tomato fruit fly Neoceratitis cyanescens can detect host fruit at a short distance using only visual stimuli, but little is known about the role of airborne volatile cues in the host searching strategy. A series of experiments is conducted in a laboratory wind tunnel, in which the behavioural responses of individual flies to volatiles from Solanaceae host plants (including tomato Lycopersicum esculentum Mill., bug weed Solanum mauritianum Scop. and Turkey berry Solanum torvum Sw.) are observed, according to some environmental (air speed) and physiological (age and mating status of females, time of day) factors. Mature females respond primarily to specific olfactory cues from blends of flowers or host fruit, preferentially unripe fruit for bug weed, as opposed to ripe fruit for Turkey berry or tomato. Males are also highly attracted by the odour of unripe fruit of bug weed. Wind plays a key role, as shown by the proportion of flies that reach the upwind section of the tunnel in the presence of both fruit odour and air flow (66.7%) and in the absence of either fruit odour (13.3%) or wind (36.7%). In response to fruit volatiles carried by wind, flies embark in a ‘plume tracking’ or ‘aim and shoot' flight, consistent with odour‐conditioned anemotaxis. Females respond to host fruit odour regardless of their age, egg load or mating status, and also more consistently in the afternoon, which is their preferential time of day for egg‐laying. Searching behaviour and response to host volatiles in N. cyanescens are discussed in the light of host‐finding and an adaptive strategy.     

Oddity in diel periodicity of Ditula angustiorana (Haw.) (Lepidoptera: Torticidae) female calling behaviour,pheromone titre and reproductive activity

The diel periodicity of female calling behaviour in Ditula angustiorana (Haw.) was studied under laboratory conditions. Observations throughout photophases and scotophases showed that D. angustiorana females presented a rare diurnal calling behaviour with two distinct calling periods, one at the first part of the photophase (L+5) and another one at the end (L+14). Mating only occurred during the second calling period (L+14). Five components were identified in extracts of the pheromone glands: Z10–14:Ac, 14:Ac, Z10–14:OH, Z8–14:Ac, Z8–14:OH in the average ratio of 74.1/18.2/4.8/2.2/0.6 respectively. The blend ratio remains similar throughout the photoperiod, whilst the total quantity of pheromone components reached a maximum during the calling period overlapping the mating period (L+14). Wind tunnel experiment confirms that the synthetic blend baited with the identified pheromone components was as attractive as calling females for males. The particular chemical structure of the components within Tortricidae and the particular occurrence of the two calling periods, one not matching any reproductive behaviour are discussed in terms of phylogeny and sexual selection.     

Behavioural switch of female Mediterranean fruit fly, Ceratitis capitata: mating and oviposition activity in outdoor field cages in Hawaii

1 Laboratory-reared normal, and wild female Mediterranean fruit flies, Ceratitis capitata (Wiedemann), were assayed in outdoor field cages to assess the impact of a mating-induced behavioural switch on mating and subsequent oviposition activity. 2 Virgin females preferred interactions with males leading to mating over attraction to, and oviposition in, artificial yellow spheres containing guava odour or green apples hung in a guava tree. Laboratory-reared females previously mated with either laboratory-reared normal males or laboratory-reared irradiated (sterile) males showed little interest in remating with males and instead, were much more likely to be found arrested on artificial and real fruit and ovipositing. Oviposition on artificial fruit was five times greater by females that had mated with either normal or irradiated males than by virgin females. Wild females showed similar qualitative changes in the mating-induced behavioural switch; however, oviposition activity was significantly less than for laboratory-reared females. 3 These results confirm that mating has a profound effect on the behaviour of female Mediterranean fruit flies and that irradiated males are functionally equal with normal males (lab-reared or wild) in their ability to alter female behaviour. These results are discussed in the context of the sterile insect technique for control of Mediterranean fruit flies in the field.     

Zur Tagesperiodik des Schlüpfens von Ephemeropteren und Plecopteren

Zusammenfassung Am Ufer des Mondsees, Oberösterreich, wurde die Tagesperiodik des Schlüpfens folgender Insektenarten in einem Schlüpfkasten experimentell untersucht: Nemoura cinerea: Schlüpft während des ganzen Tages mit Maxima zwischen 6.00 und 10.00 Uhr und nachmittags; Ephemerella ignita: Schlüpfmaximum von 18.00–20.00 Uhr; Leptophlebia marginata, Ephemera danica und Centroptilum luteolum: Schlüpfmaxima von 12.00–16.00 Uhr.Die Dauer des Subimaginallebens der untersuchten Ephemeropterenarten wurde durch Aufzucht zur Imago festgestellt. Die Bedeutung dieser temperaturabhängigen Entwicklungszeit für die Tagesperiodik der Imaginalhäutung wird diskutiert.

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Summary The diurnal periodicity of emergence of several stonefly and mayfly species was studied on the shore of lake Mondsee, Austria. Nemoura cinerea (Plecoptera) emerges during the whole day with peaks in the morning and afternoon. The Ephemeroptera Leptophlebia marginata, Ephemera danica and Centroptilum luteolum emerge as subimagines at noon and in the afternoon, Ephemerella ignita in the evening around sunset. The duration of the subimaginal life of these mayfly species was determined. The importance of the temperature-dependent length of subimaginal life for the diurnal periodicity of the imagoes' emergence from subimaginal stage is discussed.

     

Diurnal Variations in Cycling Kinematics

Physiological and biomechanical constraints as well as their fluctuations throughout the day must be considered when studying determinant factors in the preferred pedaling rate of elite cyclists. The aim of this study was to monitor the diurnal variation of spontaneous pedaling rate and movement kinematics over the crank cycle. Twelve male competitive cyclists performed a submaximal exercise on a cycle ergometer for 15 min at 50% of their W_max. Two test sessions were performed at 06:00 and 18:00 h on two separate days to assess diurnal variation in the study variables. For each test session, the exercise bout was divided into three equivalent 5‐min periods during which subjects were requested to use different pedal rates (spontaneous cadence, 70 and 90 rev min^?1). Pedal rate and kinematics data (instantaneous pedal velocity and angle of the ankle) were collected. The results show a higher spontaneous pedal rate in the late afternoon than in the early morning (p < 0.001). For a given pedal rate condition, there was a less variation in pedal velocity during a crank cycle in the morning than in the late afternoon. Moreover, diurnal variations were observed in ankle mobility across the crank cycle, the mean plantar flexion observed throughout the crank cycle being greater in the 18:00 h test session (p < 0.001). These results suggest that muscular activation patterns during a cyclical movement could be under the influence of circadian fluctuations.     

Sex pheromone components in oral secretions and crop of male Caribbean fruit flies, Anastrepha suspensa (Loew).

Chemical analysis of hexane extracts of the oral secretions from male Caribbean fruit flies, Anastrepha suspensa, resulted in identification of pheromone components including: anastrephin, epianastrephin, suspensolide, beta-bisabolene, and alpha-farnesene in a ratio of 63:396:4:8:1. Extracts of the crop from male flies contained these same components. No pheromone was detected in the extracts of female oral secretions. Bioassay of the oral secretions indicated that females were attracted to oral secretions from males but not from females. The amounts of anastrephin and epianastrephin in male oral secretions changed with age and time of the day, and were correlated with the amounts of volatile pheromone components released by male flies. The amounts of suspensolide, beta-bisabolene, and alpha-farnesene in the crop tissue were greater than those in the crop liquid, while amounts of anastrephin and epianastrephin in the crop tissue and crop liquid changed during the day. Generally, the amounts of suspensolide and beta-bisabolene decreased, and the amounts of anastrephin and epianastrephin increased from 9 am to 6 pm in both crop tissue and crop liquid. The amounts of anastrephin and epianastrephin from crop tissue or crop liquid incubated with suspensolide were significantly higher than those of control. The data show that oral secretions deposited on leaves by males contained terpenoid pheromone components that attract females and that suspensolide was converted to anastrephin and epianastrephin by enzymatic degradation in the crop of male flies.     

Dietary Modulation of Drosophila Sleep-Wake Behaviour

Background

A complex relationship exists between diet and sleep but despite its impact on human health, this relationship remains uncharacterized and poorly understood. Drosophila melanogaster is an important model for the study of metabolism and behaviour, however the effect of diet upon Drosophila sleep remains largely unaddressed.

Methodology/Principal Findings

Using automated behavioural monitoring, a capillary feeding assay and pharmacological treatments, we examined the effect of dietary yeast and sucrose upon Drosophila sleep-wake behaviour for three consecutive days. We found that dietary yeast deconsolidated the sleep-wake behaviour of flies by promoting arousal from sleep in males and shortening periods of locomotor activity in females. We also demonstrate that arousal from nocturnal sleep exhibits a significant ultradian rhythmicity with a periodicity of 85 minutes. Increasing the dietary sucrose concentration from 5% to 35% had no effect on total sucrose ingestion per day nor any affect on arousal, however it did lengthen the time that males and females remained active. Higher dietary sucrose led to reduced total sleep by male but not female flies. Locomotor activity was reduced by feeding flies Metformin, a drug that inhibits oxidative phosphorylation, however Metformin did not affect any aspects of sleep.

Conclusions

We conclude that arousal from sleep is under ultradian control and regulated in a sex-dependent manner by dietary yeast and that dietary sucrose regulates the length of time that flies sustain periods of wakefulness. These findings highlight Drosophila as an important model with which to understand how diet impacts upon sleep and wakefulness in mammals and humans.     

Spatial and temporal partitioning of behaviour by adult dacines: direct evidence for methyl eugenol as a mate rendezvous cue for Bactrocera cacuminata

Abstract. Resource use in adult Dacinae (Diptera: Tephritidae) is believed to be restricted to the host plant (i.e. the plant that provides fruit for oviposition and larval development). However, studies on Bactrocera cacuminata did not support this hypothesis. Thus, it was hypothesized that adult flies partition their diurnal activities between spatially separated resources (host plant, sugar, protein and methyl eugenol) as a function of the physiological status of the fly (immature, mature–unmated, mature–mated). In accordance with a priori predictions, the results of a field‐cage study show that there are significant diurnal patterns in abundance and behaviour, and that flies of different physiological status use resources differently. Immature flies spend most of their time foraging for sugar and protein to facilitate development. Sexually mature flies forage for sugars during the day, and at dusk, responded strongly to methyl eugenol and mate. The fact that polygynous males wait at methyl eugenol at dusk whereas the mated, monandrous females spend their time ovipositing in fruit and are nonresponsive to methyl eugenol, supports Metcalf's hypothesis that such phenylpropanoids serve as mate rendezvous cues.     

Diurnal variation of salivary cortisol in captive African elephants (Loxodonta africana) under routine management conditions and in relation to a translocation event

The present study assessed the diurnal variation in salivary cortisol in captive African elephants during routine management (baseline) and in relation to a potential stressor (translocation) to evaluate to what extent acute stress may affect diurnal cortisol patterns. Under baseline conditions, we collected morning and afternoon saliva samples of 10 animals (three zoos) on different days in two study periods (n = 3–10 per animal, daytime and period). Under stress conditions, we sampled the transported cow (newcomer) and the two cows of the destination zoo before and after the transport in the morning and afternoon (n = 3–9 per animal, daytime and transport phase), as well as after the first introduction of the newcomer to the bull (n = 1 per animal). Cortisol was measured in unextracted samples by enzyme immunoassay. Under baseline conditions, we observed the expected diurnal variation with higher cortisol levels in the morning than in the afternoon. Under stress conditions, neither a significant difference between pre- and posttransport, nor between morning and afternoon levels was found. The percentage difference between morning and afternoon cortisol after the transport, however, was remarkably lower than before the transport in the newcomer potentially indicating a stress response to familiarization. Saliva samples taken immediately after the introduction of the newcomer to the bull revealed a marked cortisol increase. Our findings indicate that stressors may disturb the diurnal cortisol rhythm. Furthermore, provided that samples can be collected promptly, salivary cortisol is a useful minimally invasive measure of physiological stress in the African elephant.