Uptake of adhesive powders from lure stations by Mediterranean fruit fly (Dipt., Tephritidae)

Powders that are capable of adhering to insect cuticles can act as carrier particles when combined with insecticides, entomopathogens, or pheromones, for targeted insect control. One potential method of delivering the powder to an insect is to lure the insects to stations containing powder using a species‐specific attractant. Here, we report on the uptake of two different powders from lure stations (henceforth called ‘dispensers’) by the Mediterranean fruit fly, Ceratitis capitata, and the transfer of the powders to conspecifics during field studies in Portugal, as part of a research programme to develop lure‐and‐kill technologies based on adhesive powder. Uptake of an electrostatic wax powder, Entostat?, from dispensers was greater than uptake of a proprietary metallic powder, Entomag?, for both wild male C. capitata visiting field‐placed dispensers and laboratory‐reared males confined with dispensers in field cages. In agreement with field data, C. capitata also took up more Entostat than Entomag when artificially dosed on dispenser trays containing powder in the laboratory, and the quantities taken up were shown to be greater than that calculated from field experiments. Increasing the amount of Entostat powder in field‐placed dispensers resulted in greater uptake of powder by visiting male C. capitata. Laboratory‐reared male and female C. capitata were released in field cages in which were hung dispensers containing adhesive powder that were baited with the male attractant trimedlure. After 24 h, the powder was successfully extracted from all males and nearly all females collected, indicating that males probably transferred powder to conspecific females after visiting dispensers. The results underscore that a lure‐and‐kill system based on adhesive powder might have potential for controlling Mediterranean fruit fly and other flying insects.     

Sex roles in the ghost moth Hepialus humuli (L.) and a review of mating in the Hepialidae (Lepidoptera)

Recently it has been found that female Lepidoptera belonging to diverse families actively court their males, rather than play a merely passive role. Male and female Hepialus humuli have been reported to come together in three different ways: (1) females are attracted to groups or ‘leks’ of white, hovering males by visual stimuli; (2) females are attracted to the males by olfactory-substances produced on the hind-tibial brushes of the males; (3) males are attracted to sedentary females by olfactory stimuli. During my study I observed H. humili males flying on a total of 21 nights in two different parts of England. The males hovered in groups for about 20 min each evening, starting and stopping their flights in synchrony. Timing depended on light intensity, northern moths flying later in the summer evenings than southern moths. I observed a total of 18 matings. Normally, a female from outside a male lek flew into the group and up to one of the males. This male then usually followed her to a settling position, where mating took place. In a few cases females touched males; in one case a female struck a male in the air so that both fell to the ground and were copulating when examined. Photographs of hovering males show that their hind tibial brushes are fully everted in flight. The organs are folded against the body when the moth is mating, at rest or dead. Whilst hovering, the males are apparently emitting pheromones which function as primary attractants, rather than as the aphrodisiacs of many other lepidopteran males. The mating behaviour of hepialids is reviewed. It is concluded that all studied hepialids which have male brush organs (some Hepialus and Oncopera, Sthenopis, Zenophassus) exhibit similar flight and mating behaviour: males hover, sit or loop back and forth on the spot in leks; females fly into male aggregations and mate there (although some published observations suggest otherwise). In contrast, hepialids such as Fraus, Oxycanus and other Hepialus that lack male brush organs have mating behaviour in which the males are the active partner, a more standard lepidopteran method. In view of the controversies surrounding mating in hepialids, future systematic and behavioural work on Hepialidae throughout the world will be worthwhile.     

Alternative Male Spawning Tactics and Acoustic Signals in the Plainfin Midshipman Fish Porichthys notatus Girard (Teleostei,Batrachoididae)

The plainfin midshipman Porichthys notatus has two male reproductive morphs, ‘Type I’ and ‘Type II’, which are distinguishable by their physical traits alone. Type I males are eight times larger in body mass than Type II males and have a six-fold larger relative sonic (vocal) muscle mass than Type II males. In contrast, the testicles of Type II males are seven times larger than those of Type I males. This study demonstrates morph-specific patterns of reproduction, including acoustic signals, for Type I and II males. Field censuses of nests showed that only Type 1 males maintained nests. Type II males and females transiently appeared in these nests in association with each other. Infra-red video and hydrophone recordings in aquaria showed that Type I males maintained nests and readily vocalized. Long-duration ‘hums’ and sequences of short-duration ‘grunts’ were produced during advertisement and agonistic contexts, respectively. Humming Type I males attracted females to their nests, pair-spawned, and then guarded egg clutches alone. By contrast, Type II males neither acoustically courted females nor maintained available nest sites, but rather ‘sneak-’ or ‘satellite-spawned’ at the nests of Type I males. Type II males infrequently produced low amplitude, short duration grunts that were similar in spectral, temporal and amplitude characteristics to the grunts of females. Type II males appear to be obligate sexual parasites of the nest-building, mate-calling, and egg-guarding Type I males. The dimorphic body and vocal muscle traits of the two male morphs in the plainfin midshipman are thus paralleled by a divergence in their reproductive tactics and the properties of their acoustic signals.     

A stag beetle’s life: sex-related differences in daily activity and behaviour of <Emphasis Type="Italic">Lucanus cervus</Emphasis> (Coleoptera: Lucanidae)

The study of behavioural ecology of rare and elusive species can be challenging, but it is an important issue for monitoring populations in planning conservation actions. This is especially true for arboreal insects inhabiting old-growth forests, whose habits are scarcely known and rely on many anecdotal data, such as the stag beetle Lucanus cervus (Linnaeus, 1758). In order to gather information on daily activity and behaviour of this species, a radio-telemetry study was conducted in a relict broadleaf forest surrounded by intensively cultivated land in northern Italy. During 2014 and 2015, from May to July, 55 stag beetles (34 males, 21 females) were radio-tagged. Most of them were caught with the aid of an entomological hand net, but nine of them were captured by emergence traps set up on deadwood. Visual contact analysis confirmed that stag beetle males were less elusive than females. Males were more frequently radio-tracked in flight, females mostly underground and in proximity of deadwood. Males were mostly observed flying at sunset, and resting or walking on standing trees during the day. The combination of air temperature and humidity determined the optimal weather conditions for male flights (temperature range: 20.7–26 °C, humidity range: 65.5–78.3%). Flying at sunset, mostly performed by males, significantly increased the detectability of the species and supported the selection of the ‘transect walk at dusk’ as the best monitoring method for the stag beetle.     

Intraspecific acoustic communication and mechanical sensitivity of the tympanal ear of the gypsy moth Lymantria dispar

Tympanal ears of female gypsy moths Lymantria dispar dispar (L.) (Lepidoptera: Erebidae: Lymantriinae) are reportedly more sensitive than ears of conspecific males to sounds below 20 kHz. The hypothesis is tested that this differential sensitivity is a result of sex‐specific functional roles of sound during sexual communication, with males sending and females receiving acoustic signals. Analyses of sounds produced by flying males reveal a 33‐Hz wing beat frequency and 14‐kHz associated clicks, which remain unchanged in the presence of female sex pheromone. Females exposed to playback sounds of flying conspecific males respond with wing raising, fluttering and walking, generating distinctive visual signals that may be utilized by mate‐seeking males at close range. By contrast, females exposed to playback sounds of flying heterospecific males (Lymantria fumida Butler) do not exhibit the above behavioural responses. Laser Doppler vibrometry reveals that female tympana are particularly sensitive to frequencies in the range produced by flying conspecific males, including the 33‐Hz wing beat frequency, as well as the 7‐kHz fundamental frequency and 14‐kHz dominant frequency of associated clicks. These results support the hypothesis that the female L. dispar ear is tuned to sounds of flying conspecific males. Based on previous findings and the data of the present study, sexual communication in L. dispar appears to proceed as: (i) females emitting sex pheromone that attracts males; (ii) males flying toward calling females; and (iii) sound signals from flying males at close range inducing movement in females, which, in turn, provides visual signals that could orient males toward females.     

The flexible lek: Phymatopus hecta the gold swift demonstrates the evolution of leking and male swarming via a hotspot (Lepidoptera: Hepialidae)

Leking moths present an evolutionary problem in their apparently simultaneous reversal of male–female attraction. The mating system of Phymatopus (or Hepialus) hecta embraces an unusually wide range of procedures. Both males and females use medium‐range olfactory attractants (also probably visual signals), and both sexes will lure, and both will approach, the other, either when flying or perched. This produces an ‘infinite variety’ which includes the classic moth mating procedure (males fly to sessile female); a typical lek procedure (female flies to sessile male); a mating swarm (hovering male follows passing female); and intermediates such as a mutual courtship dance. Male behaviour includes a flying display, two sessile displays, and an escalating war of attrition. The system is versatile, persistent, and probably evolutionarily stable. The lek site has a high density of perches suitable for copulation which facilitate predator‐escape by means of a dead drop. The whole supports a model for the evolution of resource‐based leks which commence with a concentration of females on a hotspot, leading to a concentration of males; and then an escalating process of sexual selection as males become increasingly attractive over a distance to females, and females use the males as a way of locating the resource. The main stabilizing pressure may be selection for efficient mate acquisition, and as with grouse leking systems, the precondition for evolution was probably having travelling females that actively sought a reproductive resource or a predator free space. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 184–201.     

Emergence,dispersal, and mate finding via a substrate‐borne sex pheromone in the parasitoid Metaphycus luteolus

Metaphycus luteolus Timberlake (Hymenoptera: Encyrtidae) is a facultatively gregarious parasitoid of soft scale insects. We conducted behavioral experiments to better understand the mating structure of this species. Emergence of male and female offspring is synchronized, beginning at the onset of photoperiod. Both sexes are able to disperse, although dispersal of males from natal patches appears to take longer than dispersal of females. We demonstrated the presence of a female‐produced contact pheromone using open arena bioassays and motion tracking software, testing residues deposited by walking females, and extracts of females. Males responded to the females' ‘chemical footprints’ and to acetone and hexane extracts of females by searching and arrestment on the chemical residues. Responses of males were dose dependent and diminished with time since the stimulus was deposited. Our findings support the hypothesis that non‐local mating among wasps emerging from different hosts may be quite common in these parasitoids. The implications of our results for the mating structure and previously documented sex ratio patterns of these parasitoids are described.     

Social Behaviour on the Wing in the False Vampire,Megaderma lyra

Social interaction occurs in bats. In a group of 10 Megaderma lyra (seven female and two male adults, as well as one female juvenile) held in captivity, two stereotyped flying behaviour patterns —the ‘grumbling flight’ and the ‘song flight’ — were observed and studied. The ‘grumbling flight’ is a social interaction in flight between at least two Megaderma lyra in which ‘grumble sequences’ are emitted. This behaviour is triggered by stress or arising aggression, and presumably attempts to avoid agonistic interactions with dangerous physical contact. The song flight was exclusively displayed by the dominant male bat and only directed at the non-lactating female members of the group, with a preference to alien females. This behaviour is composed of three behavioural stages, each accompanied by a specific ‘song strophe’. The song flight presumably aims at bonding the females to the male. During the grumbling flight and the song flight, M. lyra emits communication sounds in the ultrasonic range. The sounds consist of simple elements (FM_down, FM_up, CF), and are similar to types of sounds emitted for echolocation by various bat species.     

Sexual Strategies of a North American ‘Tarantula’ (Araneae: Theraphosidae)

Despite their general notoriety and popularity as pets, little is known of the behavioural ecology of ‘tarantulas’ or theraphosid spiders. We studied a theraphosid of the Arizona deserts, Aphonopelma sp., to determine behavioural events crucial to successful courtship and mating. Males search for spatially scattered females and, at short range, may detect females by substrate-borne cues. When two males are present with a single female, no direct competition such as aggression is observed. Both males may mate with a single female in rapid succession, with no evidence of post-copulatory mate guarding. Despite the potential for sexual cannibalism, courtship and mating behaviour patterns exhibit few aggressive elements and males nearly always survive sexual encounters with females. The mating system of this Aphonopelma species may best be described as a type of scramble-competition polygyny, in which the ability of males to locate receptive females is an important determinant of mating success in males. Multiple mating by females renders predictions concerning fertilization success uncertain, due to the possibility of sperm competition and ‘cryptic’ female mate choice.     

Upwind anemotaxis in response to cue-lure by the Queensland fruit fly, Bactrocera tryoni

Movements of mature male Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) were observed individually in a wind tunnel under conditions of ‘cue-lure with wind’, ‘cue-lure with no wind’, ‘wind only’ and ‘no wind or cue-lure’. Further observations were made using a dense foliage array in the wind tunnel and a structured plume of cue-lure. Patterns of walking or flying were essentially the same in all of the first four treatments except that in the ‘cue-lure with wind’ treatment, over half of the flies moved in a consistent track upwind for at least 400 mm at some time during the first 5 min of observation. With clean wind, only 10% of the flies did this. The result was that mean net upwind displacement after 5 min in the ‘cue-lure with wind’ treatment significantly exceeded that in the other three treatments, the results of which did not differ significantly from each other. The upwind tracks were accomplished by either walking or flying (with or without stops) or by a combination of both. When the wind tunnel was filled with a dense foliage array, the results with cue-lure laden wind were similar to those obtained with the equivalent treatment without foliage, except that upwind tracks were predominantly in short stages. When flies were exposed to a structured plume of cue-lure odour (without foliage present), they did not apparently alter their behaviour on leaving or entering the plume, but some did make consistent upwind tracks while they were in the plume.     

The Influence of Age on Male Mate‐Searching Behaviour in Thornbug Treehoppers

One prediction from life‐history theory is that males should increase investment in reproductive effort as they age because the opportunity for future reproductive events declines. However, older males may not be able to increase their reproductive effort if condition declines with age. The effect of age‐related changes in condition may be especially important for energetically costly activities such as moving within and between habitat patches while searching for mates. Although such searching is a component of many mating systems, the relationship between age and active mate searching has not been investigated. We investigated whether mate‐searching effort increased with age in the thornbug treehopper, Umbonia crassicornis (Hemiptera: Membracidae). In this species, males search for females using a ‘fly‐call‐walk’ strategy consisting of three phases: (1) flying from one plant to another; (2) walking and signalling while on a plant; and (3) close‐range courtship of encountered females. We measured several aspects of mate‐searching behaviour over the month‐long period of a male’s reproductive lifetime. Over the relevant period of male sexual activity (19–33 d), male condition remained stable. However, older males (25–33 d) did not search more actively than younger males as predicted; instead, younger males (19 d) had greater plant‐to‐plant flight activity and found females faster. Within‐plant walking rates and courtship duration did not differ among age classes. These results suggest that thornbug males may be investing so heavily in mate searching at younger ages that they are unable to increase investment in searching effort when they get older. As a result, older males are likely to be at a competitive disadvantage when active searching is required to locate sparsely distributed females.     

Time budget and activity rhythm of wild Great Bustard in winter

Instantaneous scanning and focal animal sampling methods were used to record the behaviors of nine wild Great Bustards (Otis tarda) and their time budget from November 2004 to January 2005. The results indicated that foraging and resting time accounted for as high as 46.40% and 39.12%, respectively, while the rest of the behaviors, such as walking, alerting, preening and flying, only shared 6.83%, 2.03%, 2.64% and 1.84%, respectively. There were significant differences in foraging time, alert time and walking time between males and females (P < 0.05). The preening and flying were closely related to males (Pearson’s correlation, r = 0. 932), and it was similar for resting and walking in females (Pearson’s correlation, r = 0. 716). Resting occurred mainly at dawn and dusk, and foraging appeared mostly from 9:30 to 11:30 and from 14:30 to 16:30. Non-parameter testing demonstrates that time budget among resting, flying and others in two temperature intervals (> −5 °C and ⩽ −5 °C) was significantly different.     

Oviposition site selection and avoidance of additional mating by females of the dragonfly,Cordulia aenea amurensis Selys (Corduliidae)

Females of an odonate species in which oviposition sites overlap with mating sites may adopt one or more of the following strategies when they lay eggs except when they ‘trade’ mating for access to suitable oviposition sites or for services (guarding, etc.) provided by males: (1) ovipositing at hidden places; (2) ovipositing at a time when males are neither patrolling nor watching; (3) indicating non-receptivity by a behavioral display. The density of ovipositing females of the dragonfly, Cordulia aenea amurensisSelys which was studied between 1970 and 1983 at a pond (Hôrai-numa, Sapporo, Hokkaido) had a high negative correlation with the distance from ‘entrance’ (a part of shore at which the arrival of most adults seems to have occurred). On the other hand, oviposition was rarely observed at a sector being distant from entrance in spite of the inference that larval survivorship was probably high at this sector. Most females oviposited among emergent vegetation in which approach of males to them was difficult, and they scarcely traveled across the open water in search of oviposition sites. Therefore, most females of the population studied were considered to adopt the first strategy. The second and third strategy were not adopted by the population studied. Finally, the influences of some environmental factors and traits possessed by a species on the adoption of these tactics or on the execution of the ‘trades’ were discussed.     

Genetic effects on flight capacity in the beet armyworm,Spodoptera exigua (Lep., Noctuidae)

The beet armyworm, Spodoptera exigua, is an important migratory insect pest in tropical and subtropical regions worldwide. The current study investigated genetic variation in the flight capacity of both female and male moths, using a quantitative genetics approach. The offspring–parent regression showed that parents had a significant influence on the flight duration of offspring, and the heritability estimated as 0.302. The upward selection increased mean flight duration from 123.7 to 284.6 min in females and from 113.9 to 254.0 min in males during 8 h of flight test; by contrast, downward selection decreased it from 123.7 to 65.6 min in females and from 113.9 to 29.8 min in males, while it did not change significantly in either females or males of the control line over eight generations. The mean realized heritability was estimated as 0.432 based on upward selection but 0.130 on downward selection, suggesting the asymmetry of response to selection on flight capacity. Reciprocal crosses between the two selected lines confirmed the dominance of ‘long‐flying genes’ in the inheritance of flight capacity. A positive genetic correlation was found between increased flight duration and pupal weight. The presence of such additive genetic variance and covariance for flight capacity and the fitness trait, pupal weight, in the population of S. exigua may have underpinned the evolution of its migratory behaviour.     

Orientation of male gypsy moths,Lymantria Dispar (L.), to pheromone sources: The role of olfactory and visual cues

The role of olfaction and vision in the close-ranging flying and walking orientation of male gypsy moths, Lymantria dispar(L.), to females was studied in the forest and in the laboratory. In the forest, feral males found an isolated pheromone source as readily as one supplemented with female visual cues; dead, acetonerinsed females deployed without pheromone received virtually no visitations. In flight tunnel choice experiments using cylinders as surrogate trees and pheromone in different spatial configurations, visual attributes of the female did not influence either the males' choice of landing site or the efficiency with which they located the female. Rather, the presence of pheromone on the cylinder was necessary to elicit orientation as well as landing and walking on the cylinder. When a female visual model was placed in various positions around a pheromone source, walking males oriented primarily to the chemical stimulus. There were, however, indications that males would alter their walking paths in response to female visual cues over short distances (<5 cm), but only if they continued to receive pheromone stimulation. When visual and chemical cues were abruptly uncoupled by altering the trajectory of the pheromone plume, most males responded to the loss of the odor cue rather than to visual cues from the female. Temporal pheromone stimulation patterns affected male walking orientation. When stimulated by pheromone, males oriented toward the source; loss of the odor cue prompted an arearestricted local search characterized by primarily vertical and oblique movements with frequent reversals in direction. Presumably these maneuvers enhance the likelihood of recontacting the plume or serendipitously encountering the female. The apparent lack of visual response to the female is discussed in light of morphological and behavioral evidence suggesting that gypsy moths were formerly nocturnal.     

Inheritance of vector competence by the thrips Ceratothripoides claratris (Shumsher) (Thysanoptera: Thripidae)

The thrips Ceratothripoides claratris is an efficient vector of the Capsicum chlorosis virus (CaCV). Transmission studies with a natural population of C. claratris found in a greenhouse ‘GH’ and a ‘colony’ derived from this ‘GH’ population by selection and inbreeding resulted in lowering the percentage of viruliferous individuals within the ‘colony’. After passing through approximately 20 generations, the ‘colony’ lost the ability to transmit the CaCV. When either viruliferous or non‐viruliferous virgin females reproduced parthenogenetically, 81% of F1 arrhenotokous males inherited their viruliferous status from their mothers, whilst, no viruliferous offspring arose from non‐viruliferous virgin mothers. Crosses between viruliferous and non‐viruliferous individuals suggest that the competence of the thrips C. claratris as a vector for this virus is probably a heritable trait controlled by a recessive allele.     

A NEW ACOUSTIC SIGNAL OF THE FRUIT-FLIES DROSOPHILA SIMULANS AND D. MELANOGASTER

ABSTRACT

We found a new acoustic signal in Drosophila simulans (si) and D. melanogaster (me). It is a ‘rejection signal’ (RS) produced by adult males and young males and females in response to the courting behaviour of mature males who emit ‘pulse songs’ (i.e. love song: LS). It occurs most frequently in si, less in adults me except if the interacting males belong to different chemical morphs (i.e. temperate or equatorial population). There are no differences in the LS characteristics directed to various sexes and ages. The RSs produced by adult males or by young animals do not differ significantly either. They are emitted by neither virgin nor fecundated adult me females but a few times by virgin adult si females. The RS (like the LS) is a multipulse signal but intervals between pulses are about twice those of LS, around 90 ms for si and 80 ms for me. They are very irregular, as is the distribution of energy along the bandwidth mainly between 300 and 800 Hz for si and 200 and 600 Hz for me. The sound level of the RS is from 10 to 20 dB less than the LS. The RS seems to be linked to the ‘flicking’ behaviour produced by both wings, while the LS always corresponds to the so-called ‘wing vibration’.     

Effects of pheromone plume structure and visual stimuli on the pheromone-modulated upwind flight of male gypsy moths (Lymantria dispar) in a Forest (Lepidoptera: Lymantriidae)

The pheromone-modulated upwind flight ofLymantria dispar males responding to different pheromone plume structures and visual stimuli designed to mimic trees was video recorded in a forest. Males flying upwind along pheromone plumes of similar structure generated tracks that were similar in appearance and quantitatively similar in almost all parameters measured, regardless of the experimentally manipulated visual stimuli associated with the pheromone source. Net velocities, ground speeds, and airspeeds of males flying in point-source plumes were slower than those of males flying in the wider, more diffuse plumes issuing from a cylindrical baffle. The mean track angle of males flying in plumes issuing from a point source was greater (oriented more across the wind) than that of males flying in plumes issuing from a transparent cylindrical baffle. Males flying in point-source plumes also turned more frequently and had narrower tracks overall than males responding to plumes from a cylindrical baffle. These data suggest thatL. dispar males orienting to pheromone sources (i.e., calling females) associated with visible vertical cylinders (i.e., trees) use predominantly olfactory cues to locate the source and that the structure of the pheromone plume markedly affects the flight orientation and the resultant track.     

The Hilltopping Mating System of Leschenaultia adusta (Loew) (Diptera: Tachinidae)

Males of the tachinid fly Leschenaultia adusta perch on small trees and shrubs on the highest parts of Usery Peak in central Arizona. Individuals select twig perches on the downwind side of these plants and fly out spontaneously from time to time or in response to another passing insect. Conspecific males elicit chases that on occasion escalate into elaborate, high-speed pursuit flights that go back and forth near the plant for several minutes. Although several males sometimes perch together briefly in the same plant, typically only one individual remains at a site for more than an hour on any given day. These site-faithful males can be considered territorial residents; they constituted about one-quarter of the males marked during the study. More than half of these residents returned to the same perch plant for two or more days. Perch plants varied in their attractiveness to male flies; male preferences were largely consistent across two years of study. Given that females were occasionally observed mating at male-occupied plants, we place the mating system of L. adusta within the hilltopping territorial category in which males compete for landmark perching sites attractive to receptive females. As is true for other hilltopping insects, receptive females of L. adusta appear to be rare and widely distributed.     

Do Female Zebra Finches,Taeniopygia guttata,Choose Their Mates Based on Their ‘Personality’?

A major challenge in behavioural and evolutionary ecology is to understand the evolution and maintenance of consistent behavioural differences among individuals within populations, often referred to as animal ‘personalities’. Here, we present evidence suggesting that sexual selection may act on such personality differences in zebra finches (Taeniopygia guttata), as females seem to choose males on the basis of their exploratory behaviour per se, while taking into account their own personality. After observing a pair of males, whose apparent levels of exploration were experimentally manipulated, females that exhibited low‐exploratory tendencies showed no preference during mate choice for males that had appeared to be either ‘exploratory’ or ‘unexploratory’. In contrast, intermediate and highly exploratory females preferred apparently exploratory males over apparently unexploratory ones. Our results suggest that behavioural or genetic compatibility for personality traits might be important for mate choice, at least for exploratory individuals.